Zoological Journal of the Linnean Society, 2011, 162, 351–442.

With 219 figures

Revision and phylogeny of the bee genus

Paratetrapedia Moure, with description of a new genus
from the Andean Cordillera (Hymenoptera, Apidae,
Tapinotaspidini)
ANTONIO J. C. AGUIAR1* and GABRIEL A. R. MELO2
1
Departamento de Zoologia, Universidade de Brasília, Brasília, DF, Brazil
2
Laboratório de Biologia Comparada de Hymenoptera, Departamento de Zoologia, Universidade
Federal do Paraná, Caixa Postal 19020, Curitiba, PR Brazil

Received 15 January 2010; revised 17 May 2010; accepted for publication 24 May 2010

The bee genus Paratetrapedia represents a commonly collected group of bees and is especially diverse in forested
areas of the Neotropics. Its taxonomy has remained poorly understood because of a lack of modern revisionary work
and numerous species described as Tetrapedia whose type specimens have not been re-examined in recent times.
Here, a comprehensive study was carried out to review the taxonomy of the genus Paratetrapedia and to
investigate cladistically the relationships amongst its species. Eighteen new species of Paratetrapedia are
described, giving a total of 32 species in the genus. A phylogenetic analysis of the species of Paratetrapedia was
carried out using 61 morphological characters for 41 terminal taxa. The phylogenetic results confirmed the
monophyly of Paratetrapedia and allowed the recognition of five species groups: the lugubris, moesta, bicolor,
lineata, and flavipennis groups. Nasutopedia gen. nov., recognized as the sister group of Paratetrapedia and with
its distribution restricted to the western forested portions of the Andean highlands, is proposed based on distinct
morphology, its placement in the phylogenetic tree, and biogeographical patterns. Species of Paratetrapedia are
especially diverse in the Amazon Forest; the eastern Brazilian Atlantic Forest contains four endemic species, and
one species is endemic to the Cerrado of central Brazil. Paratetrapedia shows a biogeographical pattern similar to
other Neotropical groups of bees and birds, with wide distribution and high diversity in lowland forests and whose
sister taxon occurs on highlands of north-western portions of the Andean cordillera. Identification keys for males
and females of all species are provided, as well as distribution maps and illustrations of general external
morphology and genitalia.

© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442.
doi: 10.1111/j.1096-3642.2010.00678.x

ADDITIONAL KEYWORDS: Apinae – Neotropical – oil bees – Tetrapedia.

INTRODUCTION the largest group, being commonly collected on

Tapinotaspidini is the most diverse group of oil-
collecting bees by both number of genera and biologi-
cal habits. Ninety-four species distributed in 12
genera are recognized within the tribe (Aguiar, 2007).
Amongst them, the genus Paratetrapedia represents
*Corresponding author. Current address: Departamento de
Zoologia, Universidade de Brasília, 70910-900, Brasília, DF,
Brazil. E-mail: ajcaguiar@gmail.com
flowers of Malpighiaceae and presenting specialized
morphological and behavioural features for oil and
pollen collecting (Vogel, 1974; Albuquerque & Rego,
1989; Alves-dos-Santos, 2003).
Despite a current increase in our knowledge of the
taxonomy of the tribe Tapinotaspidini, its phylogeny
and biogeography are still poorly understood. Revi-
sionary work of the taxonomy and phylogeny of this
group is fundamental for the reconstruction of the
evolution of oil-floral rewards in the tribe, and also for

© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442 351
352 A. J. C. AGUIAR and G. A. R. MELO
a better understanding of the biogeographical rela-
tionships of the Neotropical areas of endemism.
The genus Paratetrapedia was proposed by Moure
(1941) when he recognized that the old genus Tetra-
pedia included many distinct unrelated lineages that
could be easily distinguished by the number of labial
palpomeres. In that study, Moure transferred to
Paratetrapedia nine species described in Tetrapedia.
Later, when revising groups previously included in
Exomalopsini, Michener & Moure (1957) broadened
the scope of Paratetrapedia to include seven subgen-
era: Paratetrapedia sensu stricto, Xanthopedia, Tropi-
dopedia, Lophopedia, Amphipedia, Arhysoceble, and
Trigonopedia. Trigonopedia and Arhysoceble were
originally proposed as genera by Moure (1941, 1948),
a status revalidated more recently by Roig-Alsina
(1997) based on phylogenetic analyses of the tribe
Tapinotaspidini. Recently the subgenus Amphipedia
was synonymized under Lophopedia through exami-
nation of its type species (Aguiar & Melo, 2005). In
the present study, the subgenera of Paratetrapedia
are treated as genera following the views of Moure
(1994) and Aguiar (2007). Classifications in which
genus level status has been given to many groups of
Neotropical bees, as adopted for the stingless bees
(e.g. Pedro & Camargo, 2003; Rasmussen & Cameron,
2010) and Tapinotaspidini bees (e.g. Roig-Alsina,
1997, 1999), are supported mainly by phylogenetic
studies. These classifications provide better and more
precise concepts for communicating other biological
information relating to these taxa, such as morpho-
logical and behavioural comparative studies, biogeog-
raphy, etc. Some large groups of bees (e.g. Megachile,
Xylocopa, Ceratina) are traditionally subdivided into
subgenera only because of poor knowledge of their
internal phylogenetic relationships.
The phylogenetic analysis for Tapinotaspidini pre-
sented by Roig-Alsina (1997) was not intended to
investigate the relationships amongst the Paratetra-
pedia lineages. It included only one species of
Lophopedia and one of Paratetrapedia as at that time
the relationships amongst the lineages related to
Paratetrapedia were still unknown. Recently, Aguiar
& Melo (2007a) presented a review of the genus
Tropidopedia that included a phylogenetic analysis
with representatives of the closest groups of Paratet-
rapedia. That analysis supported Paratetrapedia as
monophyletic, with Tropidopedia and Lophopedia as
sister groups.
Very little is known about the biology of Paratetra-
pedia species. Females and males have specialized
morphological apparatuses to collect floral oils. The
oil-collecting apparatus is composed mainly of the fore
basitarsus, which is compressed with the posterior
surface concave and the anterior surface convex. The
concave surface supports dense plumose setae and the
© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
outer margin possesses a comb of simple setae (Vogel,
1974; Buchmann & Buchmann, 1981; Neff &
Simpson, 1981). Sazima & Sazima (1989) and Mick-
eliunas, Pansarin & Sazima (2006) also reported the
use of mid legs by Paratetrapedia for oil collecting.
This behaviour is supported by the presence of spatu-
late setae on ventral surface of tibia on females
of Paratetrapedia, Lophopedia, and Tropidopedia
(Aguiar & Melo, 2007a).
The first records of Paratetrapedia species visiting
oil flowers can be attributed to Fritz Müller in his
letter to Charles Darwin (Darwin, 1877) reporting
bees of the genera ‘Tetrapedia’ and Epicharis gnawing
glands on the calyces of flowers of Bunchosia gaud-
ichadiana (Malpighiaceae). The idea of the associa-
tion of these bees with the glands of Malpighiaceae
were reinforced by the notes of Friese (1899), when he
described Tetrapedia bunchosiae (junior synonym of
Paratetrapedia fervida) based on some bees sent from
Brazil by Herman Müller, brother of Fritz Müller.
Vogel (1974) suggested that Paratetrapedia species
are primarily pollinators of plants with epithelial oil
glands like Orchidaceae (e.g. Stigmatostalix, Ornitho-
phora, Oncidium, Zygostates) and secondarily Mal-
pighiaceae. Species of Paratetrapedia seem to be
especially important as pollinators of Malpighiaceae
with small flowers (Steiner, 1985; Vogel, 1990).
The nesting biology of Paratetrapedia is only known
for one species. Camillo, Garófalo & Serrano (1993)
described one aggregation consisting of ten nests of
Paratetrapedia lugubris (Cresson) [cited as Paratet-
rapedia gigantea (Schrottky)]. These nests were found
in a dead branch of Cedrella sp. (Meliaceae). The
nests were composed of horizontal tunnels in the
same orientation as the wood fibres, some of the
tunnels branching into secondary tunnels, each with
series of two to eight cells.
The goal of the present work is a phylogenetic study
and taxonomic revision of the genus Paratetrapedia
sensu Moure (1994), including identification keys to
the species, distribution maps, illustrations of termi-
nalia, and discussion of its patterns of biogeographi-
cal distribution.
MATERIAL AND METHODS
The terminology used in the descriptions of morphol-
ogy follows Michener (1944), Urban (1967), and
Michener (2000), except for the use of metapostnotum
(Brothers, 1976) instead of propodeal triangle and
gonapophyses (Smith, 1970) instead of valva of the
male genitalia. The flagellomeres of the antenna are
indicated as F1, F2, etc.; terga and sterna, respec-
tively, as T1 to T7 and S1 to S8. The deeply impressed
area above the antennal fovea is referred to as the
antennal scrobe. The tergal and sternal bands of
 SYSTEMATICS OF THE GENUS PARATETRAPEDIA
hairs mentioned in the descriptions are located at the
posterior margins of these sclerites. The carina on the
vertex, posterior to the ocelli, is here designated as
the post-ocellar carina instead of pre-occipital carina
because of its uncertain homology with the limits of
the occipital area.
The labels of the specimens examined were tran-
scribed in the sections Type material and Additional
examined material (Appendix) in the following way:
one inverted bar (\) to indicate different lines on the
label, two inverted bars (\\) to indicate that the
inscription is on the back of the label, and quotation
marks to indicate different labels on the same
specimen.
All measurements are in millimetres. The density
of the punctures is relative to their diameters and the
abbreviation pd is used for puncture diameter (e.g.
< 2 pd: distance between the punctures less than two
times the diameter of the punctures). The size of the
punctures was classified into four types: minute, fine,
coarse, and very coarse following Aguiar & Melo
(2007a).
The distribution maps were produced in the soft-
ware ArcView GIS 3.2. The inner contour of the
Andean cordillera, represented by the elevation of
1000 m above sea level, was plotted in grey using the
digital elevation model ETOPO5 in the software
MICRODEM 7.0 (Guth, 2003) and ArcView GIS 3.2.
The geographical coordinates of the localities were
obtained from the internet pages of Centro de Refer-
ência em Informação Ambiental (http://splink.cria.
org.br/geoloc), GeoNet Names Server (http://
geonames.nga.mil/ggmagaz), and Global Gazetteer
(http://www.fallingrain.com/world/). The Cerrado
limits shape file was derived from the ecoregions
presented in Olson & Dinerstein (2002), and it was
obtained from http://worldwildlife.org/science. Three
additional ecoregions were included adjacent to the
Cerrado in the grey-shaded area in Figure 70: ‘Mara-
nhão Babaçu Forest’, ‘Campos rupestres’, and ‘Beni
savanna’.
The acronyms of museums cited in the text and
their respective curators are: AMNH, American
Museum of Natural History, New York City, New
York, USA, Dr Jerome Rozen Jr.; ANSP, Academy
of Natural Sciences of Philadelphia, Philadelphia,
Pennsylvania, USA, Dr Jason Weintraub; ASC, Dr
John Ascher Private Collection; BLCU, Bee Biology
and Systematics Laboratory, Utah State University,
Logan, Utah, USA, Dr Terry Griswold; BMNH, The
Natural History Museum, London, UK, Mr George
Else; CLAUS, Dr Claus Rasmussen private collection;
CUIC, Cornell University, Ithaca, New York, USA, Dr
Rick Hoebeke; DBAI, Departamento de Zoologia, Uni-
versidade de Brasília, Distrito Federal, Brasília,
Brazil, Dr Antonio Aguiar; DZUP, Departamento de
© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
353
Zoologia da Universidade do Paraná, Coleção Pe. J.S.
Moure, Paraná, Curitiba, Brazil, Dr Gabriel Melo;
IBUSP, University of São Paulo, São Paulo, São
Paulo, Brazil, Dra. Isabel Alves dos Santos; IBUV,
Vogel Collection, Institute für Botanik und Botanis-
cher Garten der Universität Wien, Wien, Austria, Dr
Stefan Vogel; FCVZ, Dr Fernando Zanella private
collection; INBP, Museo Nacional de la Historia
Natural del Paraguay, San Lorenzo, Paraguay,
Mr Bolivar Garcete; IMLA, Universidad Nacional
de Tucumán, Fundación y Instituto Miguel Lillo,
Tucumán, Argentina; INHS, Illinois Natural History
Survey, Champaign, Illinois, USA, Dr Colin Favret;
LEA, Universidade Federal do Maranhão, Maranhão,
São Luís, Brazil, Dr Márcia Rego; MRSN, Museo
Regionale di Scienze Naturali, Torino, Piemonte,
Italy, Dr Guido Pagliano; MNHP, Muséum National
d’Histoire Naturelle, Paris, France, Dr Claire Ville-
mant; MNRJ, Museu Nacional, Rio de Janeiro, Rio de
Janeiro, Brazil, Dr Miguel Monné; MPEG, Museu
Paraense Emílio Goeldi, Pará, Belém, Brazil, Dr
Orlando Silveira; MSML, Museo de Historia Natural,
Universidad Nacional Mayor de San Marcos, Lima,
Peru, Dr Gerardo Lamas; MZSP, Museu de Zoologia
da Universidade de São Paulo, São Paulo, Brazil, Dr
Beatriz Coelho and Msc. Rodrigo Gonçalves; NHMV,
Naturhistorisches Museum Wien, Wien, Austria, Dr
Stefan Schoedl; NHRS, Naturhistoriska Riksmuseet,
Estocolm, Sweden, Dr Niklas Jönsson; RPSP, Univer-
sidade de São Paulo, Ribeirão Preto, São Paulo,
Brazil, Dr João M.F. Camargo; SM, Departamento de
Biologia, Universidade de São Paulo, Ribeirão Preto,
São Paulo, Brazil, Dr Sidnei Mateus; Entomology
Division, Natural History Museum, Kansas,
Lawrence, USA, Dr Zachary Falin; UENF, Univer-
sidade Estadual do Norte Fluminense, Campos de
Goitacazes, Rio de Janeiro, Brazil, Dr Maria Cristina
Gaglianone; UFMG, Coleções Taxonômicas da Univer-
sidade Federal de Minas Gerais, Belo Horizonte,
Minas Gerais, Brazil, Dr Fernando Silveira; UNAN,
Universidad Nacional Autônoma de Nicarágua, Léon,
Nicarágua, Dr Jean-Michel Maes; ZMB, Museum für
Naturkunde der Humbolt-Universität zu Berlin,
Berlim, Germany, Dr Frank Koch; ZSMC, München
Zoologische Staatssammlung, Munich, Germany, Dr
Benjamin Bembé. Most of the specimens from Collec-
tion Pe. J.S. Moure (DZUP) are listed without explicit
indication of the depository collection; this can be
easily deduced by transcription of the label containing
the DZUP register number. Only the type material is
listed in the main text, with the additional examined
material listed in the Appendix because of the large
number of specimens examined.
Type material of most species originally described
in the genera Tetrapedia and Chalepogenus was
examined in order to determine if they belong in
354 A. J. C. AGUIAR and G. A. R. MELO
Paratetrapedia. Type material of six species described
by Curt Schrottky (1909) from Paraguay could not be
located: Tetrapedia gigantea Schrottky, 1909; Tetra-
pedia trigonaeformis Schrottky, 1909; Tetrapedia
amalthea Schrottky, 1909; Tetrapedia anisitsi Schrot-
tky, 1909; Tetrapedia sapucayensis Schrottky, 1909;
Tetrapedia melanopus Schrottky, 1909. In addition,
the holotype of Tetrapedia elongata Friese, 1899 was
not found. Their original descriptions do not permit
assignment if they belong to Tapinotaspidini or Tet-
rapedia. Unfortunately, the type material of most of
the species described by Schrottky (1909) after his
move to Paraguay should be considered lost (Bolivar
Garcete, pers. comm.). In the collection of ZMB there
are specimens of Paratetrapedia with Friese’s hand-
written labels indicating new species dated 1904;
however, these appear to be unpublished manuscript
names (cf. Rasmussen & Ascher, 2008).
A cladistic analysis of the genus Paratetrapedia and
its related groups was carried out to test its mono-
phyly and reconstruct the phylogenetic relationships
amongst the species. The relationships within the
species groups could not be properly reconstructed
because of rampant homoplasy and numerous incom-
parable characters that could not be coded amongst
the set of species.
Characters were constructed from the study of
external morphology of dried specimens and also the
morphology of male terminalia. The ingroup was com-
posed of the 32 species of Paratetrapedia and the
outgroup was made up of nine species of related taxa
of Tapinotaspidini. Series of specimens of each species
were examined to verify possible intraspecific varia-
tion of character states.
The character matrix was edited in the software
WINCLADA 1.0 (Nixon, 1999). Inapplicable charac-
ters are indicated by ‘-’ in the matrix, and missing
data by ‘?’. The analyses were carried out using the
software NONA (Goloboff, 1999) implemented by
WINCLADA with all characters treated as non-
additive. A heuristic search was performed with 1000
replications using the following commands: hold 1000
trees in memory; hold 100 starting trees in memory;
perform tree bisection-reconnection (TBR) branch
swapping [multiple TBR + TBR (mult*max*)]. An
additional search of cladograms was carried out
through implied weighting of characters in PIWE 3.0
(Goloboff 1997; commands hold10000, hold/20,
mult*1000, max*), where the characters are weighted
according to their consistency index during the tree
search (Goloboff, 1993).
PHYLOGENY OF PARATETRAPEDIA
Some of the characters included in the analyses to
evaluate the monophyly of the genus Paratetrapedia
© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
were constructed based on the taxonomic studies of
Michener & Moure (1957) and of Melo (in Silveira,
Melo & Almeida, 2002). Some of them have already
been evaluated cladistically in the studies of Tapino-
taspidini (Roig-Alsina, 1997) and of Tropidopedia
(Aguiar & Melo, 2007a). Some characters, such as the
pattern of sculpture of the metapostnotum, shape of
male pygidial process, shape of epistomal suture,
colour of wing membrane and face marks are impor-
tant in the identification of the species groups and
species within groups, but were not used because of
their continuously varying nature. The following 61
characters were constructed for the phylogenetic
analyses:
1. Setae on the first labial palpomere (females): (0)
short and straight; (1) long, apically curved
(Fig. 1). The females of Paratetrapedia and
species of the pallidipennis group of Tropidopedia
(except Tropidopedia nigrita; see Aguiar & Melo,
2007a) have long and curved setae on the first
labial palpomere.
2. Pubescence on lower margin of mandibles (males;
Melo in Silveira et al., 2002); (0) simple; (1)
plumose.
3. Setae on prementum (females): (0) short and
straight; (1) long and wavy (Alves-dos-Santos,
2002: fig. 11); Alves-dos-Santos (2002) reported
that curved setae on the prementum and labial
palpi of two species of Paratetrapedia are adap-
tations for pollen collecting.
4. Second preapical tooth of mandible (females): (0)
absent; (1) large, present. The species of the lugu-
bris group and Nasutopedia gen. nov. have large
conspicuous pre-apical teeth. The pre-apical teeth
on the mandible of Lophopedia pygmaea are very
close to each other, and in the remaining species
of the genus they seem to have been fused. In
species of the flavipennis and bicolor groups, the
mandible has one minute, vestigial, second pre-
apical tooth. In remaining species of Paratetrape-
dia the inner margin of the mandible is uniformly
concave, without any indication of the second
pre-apical tooth.
5. Epistomal suture (males): (0) convex; (1) straight.
6. Paraocular area (Michener & Moure, 1957; Roig-
Alsina, 1997; Silveira et al., 2002: identification
key for Tapinotaspidini genera): (0) flat (Fig. 11);
(1) convex, extending above the level of the mid
portion of frons, reaching the vertex margin; (2)
convex, extending at most to the level of the mid
portion of frons. The convex surface of the
paraocular area is more easily observed in males
of Xantohopedia, Lophopedia, and Tropidopedia.
7. Surface between disc of frons and upper paraocu-
lar area: (0) strongly concave (Figs 6, 8, 9, 10); (1)
 SYSTEMATICS OF THE GENUS PARATETRAPEDIA 355

Figures 1–5. Fig. 1. Paratetrapedia connexa, female: labial palpi; scale bar = 0.2 mm. Fig. 2. Paratetrapedia fervida,
female: mesoscutum pubescence; scale bar = 0.05 mm. Figs 3–5. Paratetrapedia connexa, female; 3: outer surface of fore
basitarsus, showing the hair comb for oil gathering; 4: detail of setae on second tarsomere, with the two distinct stout,
hook-like setae; scale bar = 0.2 mm; 5: mesepisternum, in lateral view, with acute carina on omaulus; scale bar = 0.5 mm.

weakly concave, almost flat (Figs 7, 11). The state
(1) is a synapomorphy for the moesta species
group.
8. Distal portion of antennal scape (males): (0) sym-
metric; (1) outer lateral portion strongly convex,
dilated (Figs 9, 46).
9. Sculpture of clypeus and supraclypeal area (modi-
fied from Roig-Alsina, 1997: character 3; Silveira
et al., 2002): (0) fine, diameter of punctures
slightly larger than setae arising from them; (1)
coarse punctures on disc and lateral margins of
supraclypeal area and clypeus, punctures more
than twice the diameter of the setae arising from
them.
10. Vertex, behind ocelli (males; Roig-Alsina, 1997;
character 4): (0) swollen, declivous; (1) carinate
(post-ocellar carina present).
11. Vertex behind ocelli (females; Roig-Alsina, 1997;
character 4): (0) swollen, declivous to narrowly
carinate (Fig. 210); (1) carinate (post-ocellar
carina present).
12. Post-ocellar carina [males; applicable only to
taxa codified as state (1) for character 10]: (0)
short, restricted to the post-ocellar portion of
vertex; (1) occupying the entire post-ocular
portion of vertex; (2) long, extending to genal
area lateral to eyes, below the post-ocular
portion of vertex.
13. Lower third of frons, above supraclypeal area
(females): (0) mostly flat and smooth; (1) with a
conspicuous sulcus.
14. Eye margin on lower paraocular area: (0) flat or
with one very short carina, not extending over
the height of upper margin of clypeus; (1) with
one long and sharp carina, extending over the
height of the upper margin of clypeus (Fig. 7).
15. Area between the lateral ocelli and medial
ocellus (Moure, 1994; Silveira et al., 2002:

© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
356 A. J. C. AGUIAR and G. A. R. MELO

© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
 SYSTEMATICS OF THE GENUS PARATETRAPEDIA 357

Figures 6–11. Head, anterior view; 6: Paratetrapedia lugubris, female (Brazil, Minas Gerais, Serra do Salitre); 7:
Paratetrapedia leucostoma, male (Brazil, Mato Grosso, Cáceres); arrows indicating the weak concave area between upper
paraocular area and disc of frons, and portion of thin carina on lower paraocular area; 8: Paratetrapedia punctata sp.
nov., male (Brazil, Minas Gerais, Serra do Salitre); 9: Paratetrapedia testacea, male (Brazil, Pará, Óbidos), arrows
indicating convex surface on paraocular area; 10: Paratetrapedia lineata, female (Brazil, Mato Grosso, Cáceres); 11:
Paratetrapedia volatilis, female (Brazil, Paraná, Piraquara); scale bar = 0.5 mm.
䉳

fig. 8.128): (0) with a conspicuous sulcus
between ocelli; (1) convex.
16. Notaulus: (0) without indication externally; (1)
indicated by a thin sulcus externally.
17. Pronotal collar (Roig-Alsina, 1997, character 18
modified): (0) lamellate dorsally, forming a con-
spicuous gutter between the main lamella and
the mesoscutum; (1) appressed against the
mesoscutum along its entire extension.
18. Pronotal collar laterally (males; character 14 in
Aguiar & Melo, 2007a, b): (0) low, not raised,
confluent with lateral portions of pronotal collar
(Figs 18–21, 210); (1) strongly raised and closed,
extending backwards to mesoscutum.
19. Mesoscutum: (0) unicolour; (1) with two yellow
stripes on central portion and lateral margins
(Figs 54–63).
20. Omaulus: (0) convex; (1) carinate (Fig. 5).
21. Omaular carina (applicable only to taxa coded
as state (1) for character 20): (0) restricted to
upper portion of mesepisternum; (1) extending
below the upper half of the mesepisternum
(Fig. 5).
22. Pygidial plate I (females): (0) with curved
margins, with one distinct triangular basal
portion and apical portion usually rectangular
(Figs 38–41); (1) with subparallel margins,
pygidial plate triangular.
23. Pygidial plate II [females; applicable only to
species coded as state (0) for character 22]: (0)
basal portion confluent with apical portion
(Fig. 39); (1) apical portion separated from basal
portion (Figs 38, 40, 41).
24. Pygidial plate II, apex of basal portion [appli-
cable only to taxa coded as state (1) for char-
acter 23; Silveira et al. 2002; in key for
Paratetrapedia]: (0) obtusely angled; (1) acutely
angled.
25. Pygidial plate III (males) (Roig-Alsina, 1997:
character 38 and 39 modified; Silveira et al.,
2002): (0) weak, with only lateral margins dis-
tinct; (1) absent, with only pygidial process.
26. Apical portion of fore basitarsus (females): (0)
convex, without projection; (1) acute, with dis-
tinct projection.
27. Pubescence of second tarsomere of fore leg,
outer margin (females; Silveira et al., 2002, key
to Paratetrapedia): (0) with similar setae; (1)
with one or two stout hooked setae, distinct
from remaining setae.
28. Apex of setae on ventral surface of mid tibia
(females): (0) acute, not wider on apex; (1) wider
on apex.
29. Pronotal lobe: (0) oval to rectangular; (1) trian-
gular.
30. Hind basitarsus, proximal portion of ante-
rior margin (males): (0) straight, without
tooth; (1) with one tooth or raised carina
(Figs 32, 33).
31. Basitibial plate, central portion (females;
Silveira et al., 2002): (0) flat; (1) convex, oval-
elongated (Figs 34, 36); (2) reniform (Fig. 35).
Females of the genus Nasutopedia possess a
weakly reniform basitibial plate.
32. Distal margin of hind tibia (females): (0) almost
straight angled; (1) weakly declivous.
33. Basitibial plate (males; Silveira et al., 2002): (0)
completely distinct; (1) indistinct on upper third;
(2) reduced, identifiable only by a short carina
on the lower third (Fig. 37).
34. Basitibial plate (females): (0) completely haired
(Fig. 34); (1) with glabrous and smooth margins
(Figs 35, 36).
35. Inner hind tibial spur (females; Roig-Alsina,
1997, character 30; Silveira et al., 2002):
(0) serrate, similar to outer spur; (1) finely
pectinate.
36. Pubescence on mandibles, coxae, trochanteres,
and lower portion of mesepisternum (males): (0)
fine and long, longer than width of mandibular
base; (1) short.
37. Colour of terga (males): (0) yellow; (1) dark
brown to black; (2) bicoloured, yellow and
brown.
38. Terga sculpture (females): (0) smooth; (1) with
fine reticulate lines (Fig. 14).
39. T1, marginal hair band (females): (0) absent; (1)
short bands of setae laterally.
40. T4, marginal hair band (females): (0) short
bands of setae laterally; (1) complete band of
setae on entire margin.
41. T4, marginal hair band (males): (0) short bands
of setae laterally; (1) complete band of setae on
entire margin.
42. T5, marginal hair band (males): (0) absent; (1)
complete to almost complete band of setae.

© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
358 A. J. C. AGUIAR and G. A. R. MELO

Figures 12–17. Figs 12–14. Paratetrapedia connexa, female; 12: vertical surface of terga 1 (T1) with distinct short
pubescence composed of dense microsetae; 13: detail of microsetae on T1; scale bar = 0.2 mm; 14: microreticulate lines on
disc of T3; scale bar = 0.05 mm. Fig. 15. Sterna 2–6 (S2–S6), Paratetrapedia connexa, male; scale bar = 0.5 mm. Fig. 16:
Paratetrapedia leucostoma, male, lateral view of S6; scale bar = 0.2 mm. Fig. 17: Paratetrapedia fervida, male, ventral
view of metasoma, with concave area of S3 covered by dense short plumose pubescence; scale bar = 0.5 mm.

43. S2, margin (males): (0) with dense row of
plumose setae throughout its margin; (1) with
only single row of simple setae, throughout its
margin; (2) with one triangular tuft of plumose
setae (Silveira et al., 2002; key to Paratetrape-
dia).
44. S2, row of setae on margin [males, applicable
only to taxa coded as state (1) for character 43]:
(0) single; (1) clustered on mid portion.
45. S2 [males; applicable only to taxa coded as state
(1) for character 43]: (0) one mid tuft; (1) two
mid tufts.
46. Row of setae along margin of S2 [males; appli-
cable only to taxa coded as state (1) for char-
acter 43]: (0) row not interrupted, continuous
(Figs 24, 25); (1) interrupted by two gaps
(Figs 26, 27) or three gaps (Figs 28, 29).
47. S3, margin (males): (0) mostly glabrous (Fig. 15);
(1) with one row of simple setae (Figs 24, 25); (2)
with one continuous row of plumose setae; (3)
deeply concave and ‘U’-shaped, covered by dense
short plumose pubescence.
48. Short and simple pubescence on pregradular
surface of S4–S5 (males): (0) covering its entire
surface; (1) restricted to lateral portions of the
sclerite.
49. Lateral margins of S6 (males): (0) without tufts of
stout plumose setae (Fig. 83); (1) with tuft of stout
plumose setae (Figs 77–82, 84).
50. Pubescence on apical portion of S7 (males): (0)
absent, or with few short stout setae; (1) with
numerous setae along margin and surface of apical
portion.
51. Apical margin of S7 (males): (0) simple; (1) folded.
52. Proximal portion of gonostylus ventrally: (0)
homogeneous, without lamellate expansion; width
almost equal throughout its length (Figs 181, 185,
187, 193, 195); (1) proximal portion with one
lamellate projection [basal expansion of gonosty-
lus (beg); Figs 147, 149, 153, 155, 159, 171, 173].
53. Lamellate projection of gonostylus [applicable only
to taxa coded as state (1) for character 48]: (0) oval,
convex (Fig. 147); (1) acute with apex directed
posteriorly (Figs 153, 157).
54. Gonapophyses: (0) without a distinct fold on inner
surface; (1) with one lamellate projection on mid
portion of inner surface [lamella of gonapophyses
(lg); Figs 148, 154].

© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
 SYSTEMATICS OF THE GENUS PARATETRAPEDIA 359

Figures 18–23. Figs 18–21. Lateral portion of pronotal collar, dorsal view; 18: Paratetrapedia flaveola sp. nov.;
female; 19: Paratetrapedia lineata, female; 20: Paratetrapedia duckei, male; 21: Paratetrapedia testacea, male; scale
bar = 0.2 mm. Figs 22–23. Lateral portion of metapostnotum, female; 22: P. lineata; 23: P. flaveola sp. nov.; scale
bar = 0.25 mm.

55. Lamella on inner surface of gonapo- 57. Spatha (males): (0) about two times wider than
physes [applicable only to taxa coded as state long; (1) about three times wider than long
(1) for character 54]: (0) not produced (Figs 150, 154).
(Fig. 187); (1) with apical portion acutely 58. Apodeme of S8 (males): (0) inconspicuous, almost
angle (lbg, Fig. 156); (2) lobed (lbg; absent; (1) large and long.
Fig. 184). 59. Dorsal projection of gonapophyses (males): (0)
56. Apodeme of gonapophyses (males): (0) with one glabrous; (1) with numerous short stout setae,
large projection extending over the gonocoxite usually on inner surface.
margin ventrally (ea; Fig. 157); (1) convex, without 60. Apex of gonocoxite dorsally (males): (0) short,
projection. obtuse; (1) long, acute.

© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
360 A. J. C. AGUIAR and G. A. R. MELO

Figures 24–29. Metasoma, ventral view, male; 24: Paratetrapedia volatilis; 25: Paratetrapedia testacea; 26: Paratetra-
pedia punctata sp. nov.; 27: Paratetrapedia lineata; 28: Paratetrapedia flaveola sp. nov.; 29: Paratetrapedia duckei;
scale bar = 0.5 mm.

Figures 30–33. Figs 30–31. Metasoma, ventral view, female; 30: Paratetrapedia lineata; 31: Paratetrapedia flaveola
sp. nov.; scale bar = 0.5 mm. Figs 32–33. Fore margin of hind basitarsus, male; 32: Paratetrapedia volatilis; 33:
Paratetrapedia fervida; scale bar = 0.5 mm.

© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
 SYSTEMATICS OF THE GENUS PARATETRAPEDIA 361

Figures 34–41. Figs 34–36. Basitibial plate, female; 34: Tropidopedia nigrita; 35: Paratetrapedia connexa; 36: Lophope-
dia pygmaea; scale bar = 0.25 mm. Fig. 37. Basitibial plate, male, Paratetrapedia testacea; scale bar = 0.25 mm. Figs 38–
41. Pygidial plate, female; 38: T. nigrita; 39: Arhysoceble dichroopoda; 40: L. pygmaea; 41: Paratetrapedia fervida; scale
bar = 0.2 mm.

61. Parapenial lobe ventrally (males): (0) with stout
setae (Fig. 216); (1) glabrous.
The search for the most parsimonious trees under
equal weighting of characters resulted in four cla-
dograms [Fig. 209; strict consensus; length: 126
steps; consistency index (CI): 54; retention index
(RI): 83], whereas the search under implied weight-
ing returned one cladogram (Fig. 210; total fitness:
455.2; length: 123; CI: 56; RI: 84). The genus
Paratetrapedia was recovered as monophyletic in
both analyses, being supported by at least 11 char-
acters in the tree resulting from implied weighting
(Fig. 210) and 13 characters in the tree from equal
weighting (Fig. 209).
The main differences between the two analyses
involve the relationships amongst the major clades
within the species groups. In both analyses, five
main clades within Paratetrapedia were recovered.
These five main clades are each recognized here
as nominal species groups: (1) lugubris group:
Paratetrapedia lugubris, Paratetrapedia connexa,
Paratetrapedia hyalinata; (2) moesta group: Paratet-
rapedia moesta, Paratetrapedia leucostoma,
Paratetrapedia scaposa, Paratetrapedia volatilis;
(3) bicolor group: Paratetrapedia bicolor, Paratetra-
pedia testacea, Paratetrapedia calcarata,
Paratetrapedia basilaris, Paratetrapedia bifrons;
(4) lineata group: Paratetrapedia lineata, Parate-
trapedia flavifrons, Paratetrapedia romani,
Paratetrapedia tocantinensis, Paratetrapedia
alsinai, Paratetrapedia fervida, Paratetrapedia
punctata; (5) flavipennis group: Paratetrapedia fla-
vipennis, Paratetrapedia duckei, Paratetrapedia
flaveola, Paratetrapedia camargoi, Paratetrapedia
atlantica, Paratetrapedia rufa, Paratetrapedia choc-
oensis, Paratetrapedia albilabris, Paratetrapedia
andina, Paratetrapedia albopilosa.
Paratetrapedia flavescens and Paratetrapedia mexi-
cana have somewhat isolated positions and were not
placed in any of the groups above. Owing to their
distinct morphologies, they deserve recognition as
monotypic species groups if future analyses confirm
the present results.
PARATETRAPEDIA MOURE, 1941
Paratetrapedia Moure 1941: 517; type species: Ancy-
loscelis lineata Spinola, 1853, by original designa-
tion. Moure (1948): 337. Michener (1954): 114.
Michener & Moure (1957): 395. Michener (1997):
46. Aguiar (2007): 620.
Chalepogenoides Michener 1942: 279; type species
Chalepogenus leucostoma Cockerell 1923a, by origi-
nal designation. Michener (1997): 14.
Comments and diagnosis
The genus Paratetrapedia can be recognized by the
following combination of characters: body length

© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
362 A. J. C. AGUIAR and G. A. R. MELO

© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
 SYSTEMATICS OF THE GENUS PARATETRAPEDIA 363

Figures 42–53. Head, anterior view; 42: Paratetrapedia flavescens sp. nov., male (holotype); 43: Paratetrapedia
flavifrons sp. nov., male (holotype); 44: Paratetrapedia mexicana sp. nov., male (holotype); 45: Paratetrapedia
bifrons sp. nov., male (holotype); 46: Paratetrapedia calcarata, male (Mexico, Los Tuxtlas); 47: Paratetrapedia
flaveola sp. nov., male (holotype); 48: Paratetrapedia punctata sp. nov., male (holotype); 49: Paratetrapedia lineata,
female (Brazil, Pará, Óbidos); 50: Paratetrapedia rufa sp. nov., male (holotype); 51: Paratetrapedia leucostoma, male
(Suriname, DZUP 021739); 52: Paratetrapedia scaposa sp. nov., male (holotype); 53: Paratetrapedia moesta, male
(Mexico, San Luís Potosí). Scale bar = 0.5 mm.
䉳

about 6.0 to 12.0 mm; integument black, brown, pale
yellow, orange yellow, or bicoloured, usually with
white or yellow marks on face, thin yellow stripes
on mesoscutum, terga usually monochromatic or
bicoloured (reddish brown and yellow), wing mem-
brane hyaline, brown, yellow or bicoloured. Pubes-
cence mainly sparse and short; first labial
palpomere, prementum and galea with numerous
apically curved setae (character 1:1; Fig. 1 – more
conspicuous in females); paraocular area with very
short plumose pubescence; vertex and upper
paraocular area with erect stout setae; mesoscutum
and scutellum with dense short velvet pubescence
(Fig. 2), intermingled with sparse long setae. Terga
mainly smooth and shiny, with bands of hairs on
hind margins: composed mainly of simple hairs in
females, and plumose hairs in males; T1–T3 with
marginal hair band occupying less than one-third of
margin laterally (for T1 – character 39:1); on T4–T6,
marginal hair bands complete along entire margin
(character 40:1), occupying one-third of margin lat-
erally or absent. Sterna of males with pregradular
area covered by dense short plumose pubescence,
complete on S3–S4 and covering only the lateral
third of S5–S6. Margins of sterna of males with
very distinct pubescence: posterior margin of S2
with row of erect setae, varying on different species
as a single continuous row or small rows; S3 with
long plumose hairs laterally; mid portion of S3 com-
pletely glabrous (Figs 24–26, 29), with one row of
straight simple setae or deeply concave, ‘U’ shaped,
covered by short plumose hairs (Figs 17, 27, 28); S4
with band of decumbent long plumose hairs, forming
long converging fringes laterally; S5 mostly glabrous
with few erect stout plumose hairs laterally; disc of
S6 with sparse short plumose hairs and two con-
spicuous rows of stout plumose hairs on lateral
margins (Figs 24–29). Second tarsomere of fore leg
of females with one or two stout curved setae,
clearly distinct from remaining setae (character
27:1; Fig. 4).
Males and females possess specialized morphologi-
cal features for oil collecting. The fore basitarsus is
flattened, with the anterior surface convex and the
posterior surface concave, and tarsomeres 2–4
bearing curved stout setae. The outer margin of the
fore basitarsus has one comb of simple setae and
the concave surface is covered by dense stout
plumose hairs (Vogel, 1974; Neff & Simpson, 1981;
Fig. 3). The females also have numerous short stout
flattened setae on the inner surface of mid and hind
basitarsi, and stout spatulate setae forming one
comb on inner surface of mid tibia. Mandible of
males with only one large preapical tooth; mandible
of females with one or two preapical teeth
(character 4:1, Fig. 6). Vertex carinate on postocular
portion, not reaching genal area. Pronotal collar
lamellate dorsally, with lateral portions curved pos-
teriorly (convex in dorsal view; Figs 18–21). Lamella
of pronotal collar mostly sharp, except in a few
species in which the lateral portions are obtuse;
mesoscutum without indication of notaular sulcus.
Pygidial plate of female differentiated in apical and
basal portions; basal portion with margins almost
carinate and apex conspicuously acute (character
24:1; Fig. 41); pygidial plate absent in males, apex
of tergum with only pygidial process. Basitibial
plate of females with smooth glabrous margins and
central portion convex reniform (character 31:2;
Fig. 35); basitibial plate of males obsolete, with only
one very small carina on lower third (character 33:2,
Fig. 37); hind basitarsus of males usually with
one carinate tooth on anterior margin (Fig. 32–33);
inner hind tibial spur serrate, similar to outer
spur (character 35:0). Head about 1.2¥ wider than
long; eyes weakly divergent toward vertex, lower
interocular distance about 0.8¥ the upper interocular
distance. S6 of males with two conspicuous tufts of
plumose hairs on lateral margins (character 49:1);
S7 of males with apical portion of membrane folded
(character 51:1) and glabrous, or with few short
stout setae (character 50:0); apodeme of S8 long
and large (character 58:1); genital capsule of males
with parapenial lobe glabrous (character 61:1); gono-
styli lamellate with ventral surface concave, usually
with one large expansion on basal portion – beg
(character 52:1; Figs 147, 153, 157); gonapophyses
with one lamellate projection on mid portion of
inner surface (character 54:1); dorsal, inner, expan-
sion of gonapophyses with many short stout setae
(character 59:1); parapenial lobe glabrous (character
61:1).

© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
364 A. J. C. AGUIAR and G. A. R. MELO

THE LUGUBRIS SPECIES GROUP
The lugubris group contains the largest species in
the genus, usually more than 10 mm in body length.
Its species can be identified by the following char-
acters: integument mostly black; mandible of female
with two conspicuous preapical teeth (character 4:1);
epistomal suture of upper margin of clypeus
strongly curved; lamella of pronotal collar acute
throughout its length; hind basitarsus of male with
a large tooth on proximal portion of anterior margin
(character 30:1); metapostnotum with dense minute

© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
 SYSTEMATICS OF THE GENUS PARATETRAPEDIA 365

Figures 54–65. Mesosoma, dorsal view; 54: Paratetrapedia flavipennis, female (Brazil, Amazonas, Tefé); 55: Paratetra-
pedia duckei, female (Brazil, Pará, Serra Norte); 56: P. duckei, male (holotype of Chalepogenus xanthaspis); 57:
Paratetrapedia chocoensis sp. nov., female (paratype; Costa Rica); 58: Paratetrapedia tocantinensis sp. nov.,
female (paratype; Brazil, Tocantins, Buriti do Tocantins); 59: Paratetrapedia romani, male (Brazil, Amazonas, Manaus);
60: Paratetrapedia scaposa sp. nov., male (dark brown form; Brazil, Acre, Cruzeiro do Sul); 61: P. scaposa sp. nov.,
male (holotype, pale orange yellow form); 62: Paratetrapedia rufa sp. nov., male (paratype; Bolivia); 63: Paratetra-
pedia basilaris sp. nov., female (paratype; Brazil, Amazonas, Manaus); 64: Paratetrapedia bicolor, female (Brazil,
Espírito Santo, Linhares); 65: Paratetrapedia albilabris, female (Costa Rica). Scale bar = 0.5 mm.
䉳

punctures, almost contiguous; omaulus carinate
(Fig. 5; character 20:1); terga of female with micros-
culpture of fine reticulated lines (Fig. 14; character
38:1); T1 of female with dense fine punctures on
vertical surface; S3 of male with posterior margin
glabrous on mid portion (Fig. 15; character 47:0); S6
of male with apical portion long and acute (Fig. 15);
T7 of male with pygidial process narrow and long;
S7 of male with numerous short stout setae on
apical portion; hind basitarsus of male with one
large acute tooth on anterior margin; apical portion
of the lamella on inner surface of gonapophyses
acutely angled (character 55:1; Fig. 148).
Paratetrapedia hyalinata is the only species of the
lugubris group with a restricted distribution, being
found only in the northern portion of the Amazon
Forest. The other two species are widely distributed
over most of the Neotropical region. Paratetrapedia
connexa and P. lugubris are not found south of the
Tropic of Capricorn (23°S) and do not occur in the
coastal Atlantic Forest of south-eastern Brazil. Only
the form of P. connexa with dark infumate wings is
found in the coastal Atlantic Forest of north-eastern
Brazil.
PARATETRAPEDIA LUGUBRIS (CRESSON, 1878)
(FIGS 66, 85, 116, 147, 148)
Tetrapedia lugubris Cresson, 1878: 135; lectotype
male, Mexico (ANSP, examined), designated by
Cresson (1916: 122).
Tetrapedia amplipennis Smith, 1879: 128; lectotype
male, designated by Aguiar (2007), Brazil: Amazo-
nas, Tefé (BMNH, examined).
Tetrapedia gigantea Schrottky, 1909: 227; holotype
female, Brazil: São Paulo (depository unknown).
Tetrapedia bombitarsis Vachal, 1909: 27; holotype
male, locality unknown (MNHP, examined).
Tetrapedia gigantea Friese, 1910: 63; lectotype
female, designated by Aguiar (2007), Brazil: São
Paulo, Jundiaí (ZMB, examined). Junior homonym
of Tetrapedia gigantea Schrottky, 1909.
Tetrapedia dentiventris Friese, 1921: 90; lectotype
male, designated by Aguiar (2007), Costa Rica: San
José, San José (ZMB, examined).
Tetrapedia lugubris; Cresson, 1879: 228. Dalla Torre,
1896: 299. Friese, 1899: 281. Cockerell, 1899: 16.
Ducke, 1901: 55. Cockerell, 1905: 326. Cockerell,
1906: 98. Lutz & Cockerell, 1920: 569. Schwarz,
1934: 14.
Tetrapedia amplipennis; Dalla Torre, 1896: 299. Cock-
erell, 1905: 326. Ducke, 1910a: 364.
Tetrapaedia [sic] amplipennis; Schrottky, 1902: 539.
Tetrapedia gigantea Friese; Ducke, 1910a: 364.
Tetrapedia gigantra [sic] Friese; Friese, 1910: 62.
Tetrapedia bombitarsis; Cockerell, 1912a: 31.
Chalepogenus lugubris; Cockerell, 1923a: 450.
Chalepogenus amplipennis; Cockerell, 1923a: 450.
Paratetrapedia amplipennis; Moure, 1941: 518. Rozen
et al., 2006: 10.
Chalepogenoides lugubris; Michener, 1942: 281.
Chalepogenoides amplipennis; Michener, 1942: 281.
Paratetrapedia gigantea (Schrottky); Michener, 1954:
116. Rozen & Michener, 1988: 7. Santos, Carvalho
& Silva, 2004: 323.
Paratetrapedia lugubris; Michener, 1954: 115 (erro-
neous identification of P. connexa, as indicated by
drawings of male genitalia, figs 68–70). Michener
& Brooks, 1984: 46, fig. 92i–j. Silveira, 1995: 428,
fig. 5a (terminal taxon in phylogenetic analyses).
Michener, 2000: 667, figs 106–1e, 106–4d. Aguiar,
2007: 623.
Paratetrapedia (Paratetrapedia) amplipennis;
Michener & Moure, 1957: 416. Silveira et al., 2002:
136. Aguiar et al., 2004: 80.
Paratetrapedia (Paratetrapedia) gigantea (Schrottky);
Michener & Moure, 1957: 416. Lucas-de-Oliveira,
1962: 1, figs 1–3 (description of pupa). Lucas-de-
Oliveira, 1966: 430. Rozen, 1984: 3. Camillo et al.,
1993: 151, figs 4–7 (description of nest). Pedro,
1996: 251. Pedro & Camargo, 1999: 202. Silveira
et al., 2002: 136
Paratetrapedia (Paratetrapedia) lugubris; Michener
& Moure, 1957: 416. Ayala, Griswold & Yanega,
1996: 461.
Paratterapedia gigantea Schr. [sic]; Steiner, 1985:
1940 [flower record: Spachea membranacea (Mal-
pighiaceae)].
Paratetrapedia gigantea (Fabricius) [sic]; Camillo
et al., 1993: 145; Melo & Zanella, 2003: 2919.

© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
366 A. J. C. AGUIAR and G. A. R. MELO

IDENTIFICATION KEY TO SPECIES OF PARATETRAPEDIA

Females
1. Omaulus carinate (Fig. 5); body usually more than 10 mm in length ........................................................ 2
– Omaulus not carinate; body usually less than 10 mm in length ............................................................... 4
2. Carina on omaulus long, extending more than two-thirds of mesepisternum laterally (Fig. 5); lateral margins of
clypeus with a single yellow dot; frons uniformly convex; terga completely black, without yellow marks; vertical
surface of T1 with silvery reflections (Fig. 12), covered with numerous short spatulate setae (Fig. 13); wings yellow
infumate or dark brown infumate (Bolivia; Brazil: Amazonas, Bahia, Distrito Federal, Goiás, Mato Grosso, Mato
Grosso do Sul, Maranhão, Minas Gerais, Pará, Paraíba, Pernambuco, Rondônia, Roraima, Tocantins; Costa Rica;
Ecuador; Guatemala; Guyana; French Guiana; Panama; Peru) ........................................ P. connexa (Vachal)
– Carina on omaulus short, restricted to about upper half of mesepisternum; clypeus completely black; frons weakly
concave in middle portion; T5–T6 usually with yellow marks laterally; T1 completely black, without silver
reflections and differentiated setae; wings mostly dark brown infumate.....................................................3
3. Fore wing membrane with basal one-third dark brown infumate and apical one-third white infumate; T4 with
marginal hair band occupying entire margin; terga completely black, without yellow spots laterally (Brazil:
Amapá; Guyana)...............................................................................................................P. hyalinata
– Fore wing membrane mostly dark brown infumate, weakly light brown on apical third; T4 with marginal
hair band occupying less than one third of margin laterally; T5–T6 usually with yellow spots on lateral portions
(Bolivia; Brazil: Acre, Amazonas, Distrito Federal, Goiás, Mato Grosso, Minas Gerais, Pará, Rondonia, São Paulo;
Colombia; Costa Rica; Mexico; Guatemala; Guyana; Panama; Peru; Venezuela; Trinidad and Tobago) .............
..........................................................................................................................P. lugubris (Cresson)
4. Lower third of paraocular area with sharp carina, extending to height of upper margin of clypeus (Fig. 7);
prementum with numerous setae with apex wavy; metapostnotum with sparse fine punctures (> 2 pd); T4 with
marginal hair band occupying about one-quarter of margin laterally ........................................................ 5
– Lower paraocular area with short carina, usually obtuse, and occupying less than one-third of paraocular area;
prementum with setae straight or slightly curved; metapostnotum with dense punctures, usually with fine
punctures intermingled amongst coarser (1–2 pd); T4 usually with marginal hair band complete, occupying entire
margin...........................................................................................................................................7
5. Body length usually more than 8.0 mm; frons with sparse fine punctures (> 2 pd); pronotal collar with
lamella obtuse on lateral portions (Costa Rica; Guatemala; Honduras; Mexico; Nicaragua; Panama)................
............................................................................................................................P. moesta (Cresson)
– Body length less than 7.0 mm; frons with dense fine punctures (1 pd), intermingled with a few sparse coarser
punctures on disc; pronotal collar with lamella mostly sharp, weakly obtuse on lateral portions....................6
6. Integument mostly orange yellow; integument on frons and mesoscutum yellow orange or dark brown, with two
thin yellow stripes on disc and lateral margins of mesoscutum (Figs 60–61); pubescence mostly pale yellow
(Bolivia; Brazil: Acre, Amazonas; Ecuador; Peru).......................................................................P. scaposa
– Integument mostly dark brown, terga and sterna dark brown or pale brown, almost yellow; mesoscutum usually
completely dark brown, rarely with pale yellow stripes on disc and margins; pubescence mostly dark brown
(Bolivia; Brazil: Amapá, Goiás, Maranhão, Mato Grosso, Mato Grosso do Sul, Pará, Roraima; Guiana; French
Guiana; Paraguay; Suriname) ............................................................................ P. leucostoma (Cockerell)
7. Pubescence mostly yellow; integument of mesosoma mostly orange yellow, or bicoloured; terga completely yellow
or bicoloured (yellow and brown) ........................................................................................................ 8
– Pubescence mostly brown; integument of mesosoma mostly black; terga dark brown..................................18
8. Wing membrane mostly yellow infumate; microtrichia orange yellow, except apical margin with dark
brown microtrichia (except P. chocoensis, from Choco region, Ecuador, which has the wing membrane dark
brown)...........................................................................................................................................9
– Wing membrane milky, almost hyaline; some specimens with membrane of radial cell yellow infumate, micro-
trichia white hyaline to pale yellow, with few microtrichia dark brown on apical margin; or wing membrane
homogeneous brown or yellow infumate with dark brown microtrichia dispersed on whole membrane...........14
9. Lamella of pronotal collar with lateral portions obtuse; genal area with thin yellow stripe along eye margin,
usually with a large gap on mid third (except P. flavipennis, which has genal area completely orange yellow);
metapostnotum with dense fine punctures on lateral portions and dense coarser punctures on disc; epistomal
suture straight, above upper margin of clypeus; row of hairs along margin of S2–S3 arranged in ‘V’ shape on
mid portion (Fig. 31); supraclypeal area weakly convex, almost flat, with dense coarse punctures (< 1 pd)
(Fig. 47)........................................................................................................................................10
– Lamella of pronotal collar acute throughout its length; genal area completely yellow; metapostnotum with dense
fine homogeneous punctures, or heterogeneous, with sparse minute punctures intermingled with coarse punctures;

© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
 SYSTEMATICS OF THE GENUS PARATETRAPEDIA 367

epistomal suture weakly curved above upper margin of clypeus; row of hairs along margin of S2–S3 arranged in
weak curve on mid portion (Fig. 30); supraclypeal area mainly smooth and strongly convex, mostly with sparse
coarse punctures (> 2 pd) (Figs 42, 43, 49) .......................................................................................... 12
10. Mesoscutum completely orange yellow, with thin yellow stripes on disc and lateral margins (Fig. 54); genal area
completely orange yellow; body length more than 10.0 mm (Brazil: Amazonas; Peru)........P. flavipennis (Smith)
– Mesoscutum with disc black, with thin yellow stripes on disc and lateral margins (Figs 55–59); genal area
dark brown with thin orange yellow stripe, usually interrupted on mid portion; body length less than 10.0 mm.
...................................................................................................................................................11
11. Mesoscutum black with yellow stripes on lateral margins not enlarged (Fig. 57); lateral portions of metanotum
dark brown, in some specimens with small yellow dot on central portion; metapostnotum varying from orange
yellow to black; head maximum width c. 2.2 mm (Costa Rica; Ecuador; Panamá) ........................ P. chocoensis
– Mesoscutum black with yellow stripes on lateral margins contacting yellow stripes of central portion (Fig. 56);
lateral portion of metanotum completely yellow; head maximum width c. 3.0 mm (Bolivia; Brazil: Acre, Amapá,
Amazonas, Mato Grosso, Pará, Rondônia; French Guiana; Guyana; Peru; Suriname) ............. P. duckei (Friese)
12. Metapostnotum with heterogeneous punctures, sparse minute punctures (> 2 pd) intermingled amongst sparse
coarse punctures (2 pd); terga with marginal zone black; frons with mid line slightly sulcated; integument bright
deep yellow (Mexico)..........................................................................................................P. flavescens
– Metapostnotum with homogeneous dense and minute punctures (1–2 pd); marginal zone of terga weakly dark
brown; frons convex, with mid line inconspicuous; integument orange yellow............................................13
13. Frons with yellow ellipsoid spot on mid line, c. 1.0 ¥ F2 diameter (Brazil: Amazonas, Para) ... P. romani (Friese)
– Frons with large yellow spot completely occupying most of disc (Fig. 43) (Brazil: Amazonas, Pará, Rondônia)....
......................................................................................................................................P. flavifrons
14. Integument mostly orange yellow, a few specimens have disc of mesoscutum brown; terga completely orange
yellow; wing membrane brown or yellow infumate with brown microtrichia..............................................15
– Integument bicoloured; terga yellow with marginal zones dark brown; wing membrane milky, slightly pale yellow,
almost hyaline, microtrichia pale yellow to white, some specimens with radial cell yellow infumate ............. 16
15. Lamella of pronotal collar with lateral portions obtuse (Fig. 18); metapostnotum with dense fine punctures on
lateral portions (1 pd), and coarser punctures on disc; terga without microsculpture of fine reticulated lines;
epistomal suture straight above the upper margin of clypeus; vertical surface of T1 smooth; body length more than
8.0 mm (Brazil: Pará; Bolivia).....................................................................................................P. rufa
– Lamella of pronotal collar acute throughout its length (Fig. 21); metapostnotum with fine homogeneous punctures
(c. 2 pd); terga usually with microsculpture of fine reticulated lines (Fig. 14); epistomal suture curved above upper
margin of clypeus; vertical surface of T1 with dense minute punctures; body length less than 8.0 mm (Brazil: Acre,
Amazonas, Pará, Goiás, Tocantins; Peru; Guyana; French Guiana; Suriname).....................P. testacea (Smith)
16. Genal area with one fine yellow stripe along eye margin, width c. 0.8 ¥ F2 diameter, usually interrupted on mid
portion; yellow stripe on paraocular area with upper portion acutely sinuous, similar to that of male (Fig. 47);
metapostnotum completely black, with dense fine punctures on lateral portions (0.5–1 pd) and coarser punctures
on disc (Fig. 23) (Bolivia; Brazil: Acre, Amapá, Amazonas, Bahia, Distrito Federal, Goiás, Maranhão, Mato Grosso,
Mato Grosso do Sul, Minas Gerais, Pará, Rondônia, Roraima, São Paulo, Tocantins; Colombia; Ecuador; Guyana;
Peru; Suriname) .................................................................................................................. P. flaveola
– Genal area with a wide yellow stripe, wider than F2 diameter, usually occupying whole genal area; yellow stripe
on paraocular area with upper portion widely obtuse (Figs 43, 49); metapostnotum completely yellow or partially
yellow, with homogenous dense fine punctures .................................................................................... 17
17. Propodeum and metapostnotum yellow except for thin brown stripes on lateral margins of metapostnotum;
metanotum yellow laterally; mesepisternum mostly yellow, in some specimens with lower portion and posterior
margin weakly reddish brown; mesoscutum with yellow stripes of lateral margins in contact with yellow stripes
of disc anteriorly (Fig. 58); lamella of pronotal collar with lateral portion obtuse; supraclypeal area with dense
coarse punctures (1 pd). Metapostnotum with dense fine punctures, smooth and shiny integument between the
punctures. Fore wing membrane milky hyaline, without dark brown microtrichia on apical margin (Brazil: Goiás,
Maranhão, Piauí, Tocantins) ........................................................................................... P. tocantinensis
– Propodeum bicoloured, yellow with large brown areas on lateral portions; metapostnotum with disc yellow and
lateral portions brown; dorsolateral areas of metanotum brown; mesepisternum mostly brown with large yellow
spot on omaular area; yellow stripes on lateral margins of mesoscutum rarely in contact with yellow stripes of disc;
lamella of pronotal collar acute throughout its length (Fig. 19); supraclypeal area with sparse coarse punctures
(2 pd), integument smooth and shiny between the punctures (Fig. 49). Metapostnotum with dense fine punctures;
surface between the punctures dull with microsculpture of fine reticulate lines (Fig. 22). Fore wing membrane with
microtrichia dark brown on apical margin (Bolivia; Brazil: Acre, Amazonas, Mato Grosso, Pará, Rondônia;
Colombia; Ecuador; Peru)..........................................................................................P. lineata (Spinola)

© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
368 A. J. C. AGUIAR and G. A. R. MELO

18. Supraclypeal area with upper portion strongly projected, with trapezoidal aspect, similar that of male (> 1 pd;
Figs 8, 48); terga with dense fine punctures (1 pd); legs with pubescence mostly black; hind tibia with tuft of white
hairs on apical portion (Brazil: Amapá, Goiás, Mato Grosso, Mato Grosso do Sul, Maranhão, Minas Gerais, Pará,
Paraná, Rondônia, São Paulo, Tocantins)................................................................................P. punctata
– Upper portion of supraclypeal area weakly convex, not produced; terga with surface largely impunctate, with few
scattered punctures laterally............................................................................................................19
19. T4 with one complete band of setae occupying whole margin or with three short bands distributed on mid portion
and laterally; body length about 6.0–8.0 mm.......................................................................................20
– T4 with marginal hair band occupying less than one third of margin laterally; body length more than 8.0 mm .
...................................................................................................................................................26
20. Frons biconvex, with mid line strongly sulcated (Fig. 45) (Costa Rica; Mexico).................................P. bifrons
– Frons convex, mid line not sulcated .................................................................................................. 21
21. Frons and mesoscutum completely black, without yellow stripes; pronotal lobe oval; metapostnotum with
heterogeneous punctures, with coarse punctures intermingled with fine punctures; body length about 9.0–
10.0 mm; first labial palpal segment with numerous long stout setae apically curved.................................22
– Frons with yellow stripe along mid line; mesoscutum with two yellow stripes on disc; pronotal lobe triangular;
metapostnotum with homogeneous dense fine punctures (1–2 pd); body length less than 9.0 mm..................23
22. Frons with dense coarse punctures (1–2 pd) intermingled with fine punctures; lamella of pronotal collar conspicu-
ously acute along its entire extension; wings with membrane pale brown infumate; hind leg with pubescence
usually dark brown, with some plumose white plumose hairs intermingled, or completely pale yellow
(Brazil: Bahia, Espírito Santo, Minas Gerais, Paraná, Rio de Janeiro, Santa Catarina, São Paulo; Paraguay)....
..............................................................................................................................P. fervida (Smith)
– Frons with sparse coarse punctures (> 2 pd); lamella of pronotal collar with lateral portions obtuse; wing
membrane dark brown infumate; hind leg with pubescence completely black (Mexico)...................P. mexicana
23. Labrum yellow; paraocular area with yellow stripe occupying more than lower half; terga dark brown with
margins translucent hyaline (Brazil: Amapá, Amazonas, Pará; French Guiana; Guyana).................P. basilaris
– Labrum weakly brown; paraocular area with weak yellow stripe; terga completely pale brown to yellow, in some
specimens with margins translucent..................................................................................................24
24. Mesoscutum black, with inconspicuous yellow stripes (Fig. 64); terga with microsculpture of dense reticulate lines
on whole surface (as in Fig. 14) (Brazil: Bahia, Espírito Santo, Minas Gerais, Paraíba, Pernambuco, São Paulo,
Rio de Janeiro)..........................................................................................................P. bicolor (Smith)
– Mesoscutum black with conspicuous yellow stripes (Figs 63, 65); terga mostly smooth, in some specimens with
microsculpture of dense reticulated lines only on disc of T3–T4..............................................................25
25. Omaulus acutely angled in upper third, almost carinate; lateral surface of mesepisternum behind omaular
carina weakly concave; terga brownish yellow; sterna pale brown; mesoscutum with conspicuous yellow stripes
(Colombia; Costa Rica; Ecuador; Guatemala; Honduras; Mexico; Nicaragua; Panama; Venezuela) ....................
.........................................................................................................................P. calcarata (Cresson)
– Omaulus right-angled in upper third; lateral surface of mesepisternum adjacent and posterior to omaular carina
flat or convex; terga dark brown, only T2 disc slightly pale yellow; sterna weakly pale yellow; mesoscutum with
faint yellow stripes (Brazil: Acre, Mato Grosso; Peru) ................................................................... P. vogeli
26. Lamella of pronotal collar acute; surface between disc of frons and upper paraocular area weakly concave (Fig. 11);
metapostnotum with heterogeneous punctures, with coarse punctures intermingled amongst fine punctures;
epistomal suture, above upper margin of clypeus, curved; terga black, usually with yellow oval spots on T3–T5
laterally; hind tibia and basitarsus with golden yellow pubescence; wing membrane black infumate; body length
more than 10 mm (Argentina: Tucumán; Brazil: Bahia, Espirito Santo, Minas Gerais, Paraná, Rio de Janeiro, Rio
Grande do Sul, Santa Catarina, São Paulo) .................................................................. P. volatilis (Smith)
– Lamella of pronotal collar with lateral portions obtuse; surface between disc of frons and upper portion of
paraocular area strongly concave; epistomal suture straight, above upper margin of clypeus; metapostnotum with
fine punctures in the lateral portions and coarse punctures on central portion; terga completely pale yellow to dark
brown, without yellow oval spots on T3–T5 laterally; hind leg pubescence mostly dark brown; wing membrane pale
brown to pale yellow infumate; body length less than 10.0 mm .............................................................. 27
27. Mesoscutum with two conspicuous yellow stripes on disc, axilla yellow on upper portion (Fig. 65); omaulus almost
right-angled in upper third; omaular area on lateral mesepisternum weakly concave (Costa Rica; Mexico)........
.......................................................................................................................... P. albilabris (Friese)
– Mesoscutum with two weak yellow stripes on the disc; axilla completely brown or black; omaular area on lateral
mesepisternum convex .................................................................................................................... 28
28. Posterolateral portions of scutellum with pubescence dark brown; T1 with dense minute punctures on vertical and
horizontal surfaces; transition zone between metapostnotum and propodeum with dense fine punctures, without
a conspicuous smooth area (Brazil: Espírito Santo, Rio de Janeiro, São Paulo)..............................P. atlantica

© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
 SYSTEMATICS OF THE GENUS PARATETRAPEDIA 369

– Posterolateral portions of scutellum with pubescence of white plumose hairs, similar to P. albilabris (Fig. 65); T1
completely smooth; transition zone between metapostnotum and propodeum with narrow smooth area, width
c. 0.5 ¥ F2 diameter (Brazil: Goiás, Maranhão, Pará) .............................................................. P. albopilosa

Males
1. S3 with wide concave area on posterior margin, covered by dense short plumose pubescence (Figs 17, 27–28) ...
.................................................................................................................................................... 2
– S3 with posterior margin not sinuous, without concave area; margin completely smooth or with one row of simple
setae adjacent to margin (Figs 15, 24–26)...........................................................................................16
2. Hind basitarsus with conspicuous tooth or carina on proximal third portion of anterior margin (Figs 32–33) ... 3
– Hind basitarsus without tooth or carina .............................................................................................. 7
3. Integument mostly dark brown or black .............................................................................................. 4
– Integument mostly yellow ................................................................................................................. 5
4. Clypeus dark brown with yellow stripe usually occupying entire lower margin; supraclypeal area dark brown or
black, in some specimens with small yellow spot on disc; scape completely reddish brown, in some specimens with
one yellow spot on lower portion; tarsomeres dark brown or weakly reddish brown; wing membrane pale brown
infumate; metapostnotum with dense fine punctures (1 pd); lamella of pronotal collar acute (Brazil: Santa
Catarina, Paraná, São Paulo, Minas Gerais, Rio de Janeiro, Espírito Santo, Pernambuco, Paraíba; Paraguai) ...
..............................................................................................................................P. fervida (Smith)
– Clypeus and supraclypeal area completely yellow (Fig. 44); scape mostly yellow; tarsomeres yellow; wing mem-
brane dark brown infumate; metapostnotum with sparse fine punctures (2 pd); lamella of pronotal collar weakly
obtuse on lateral portions (Mexico) ....................................................................................... P. mexicana
5. Wing membrane yellow infumate; mesoscutum black with thin yellow stripes on disc, and usually very wide
orange yellow stripes on lateral margins, occupying most of lateral portions, similar to P. duckei (Fig. 55); terga
completely orange yellow (Brazil: Amazonas) ............................................................................. P. alsinai
– Wing membrane mostly hyaline, weakly milky white; mesoscutum mostly black, with thin yellow stripes on disc
and lateral margins (Figs 57–58); terga with marginal zone completely dark brown .................................... 6
6. Metanotum yellow laterally. Metapostnotum with fine punctures, surface between punctures smooth and shiny.
Mesepisternum yellow laterally; metapostnotum and propodeum completely yellow; hind basitarsus with one long
acute tooth on anterior margin; epistomal suture straight above upper margin of clypeus (Brazil: Goiás, Piauí,
Tocantins, Maranhão) ....................................................................................................P. tocantinensis
– Metanotum reddish brown laterally. Metapostnotum with dense fine punctures, surface between punctures dull,
with microsculpture of dense fine sulci (Fig. 22); mesepisternum laterally mostly reddish brown, usually with one
yellow spot on omaulus; metapostnotum reddish brown with one yellow spot on disc; propodeum completely
reddish brown, some specimens with large yellow spots on anterior margin; hind basitarsus with one short acute
tooth on anterior margin; epistomal suture strongly curved above upper margin of clypeus (Bolivia; Brazil: Acre,
Mato Grosso, Pará, Rondônia; Peru)............................................................................P. lineata (Spinola)
7. Integument mostly orange yellow; terga completely yellow or yellow with wide dark brown stripe.................8
– Integument mostly brown; terga brown..............................................................................................13
8. Outer margin of mandible, lower portion of genal area, coxa, trochanters and mesepisternum ventrally with long
fine white hairs (length c. 2 ¥ the width of mandibular base); S2 with one isolated small row of simple stout setae
on mid portion of margin, similar to P. lineata (Fig. 25) (Brazil: Amazonas, Rondônia, Pará) .......... P. flavifrons
– Outer margin of mandible, lower portion of gena, coxa, trochanters and lower portion of mesepisternum with short
hairs, c. 1 ¥ the width of mandibular base; S2 with continuous row of simple setae adjacent to margin (Figs 24,
25), or with two small rows of setae on mid portion (Figs 26, 27).............................................................9
9. S2 with continuous row of simple erect setae adjacent to posterior margin; integument of supraclypeal area and
disc of frons smooth and shiny, with punctures coarse and sparse (> 2 pd) (Fig. 42); metapostnotum mostly smooth
and shiny, with minute punctures intermingled amongst sparse fine punctures (> 2 pd); gena completely yellow;
frons with large yellow spot occupying whole disc (Mexico).......................................................P. flavescens
– S2 with two small rows of erect simple setae on mid portion of apical margin; supraclypeal area and frons with
dense coarse punctures (1–2 pd); metapostnotum with dense fine punctures (1 pd); genal area reddish brown with
thin yellow stripe (c. 1.2 ¥ F2 diameter), usually interrupted on mid portion; frons with one thin yellow stripe
along mid line, c. 0.5–1.5 ¥ F2 diameter ............................................................................................. 10
10. Wing membrane hyaline with pale microtrichia; a few dark brown microtrichia on distal margin of fore wing;
T3–T6 with disc reddish brown and anterior and posterior margins yellow (Bolivia; Brazil: Acre, Amapá,
Amazonas, Bahia, Distrito Federal, Goiás, Maranhão, Mato Grosso, Mato Grosso do Sul, Minas Gerais, Pará,
Rondônia, Roraima, São Paulo, Tocantins; Ecuador; Guiana; Peru; Suriname) ................................ P. flaveola

© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
370 A. J. C. AGUIAR and G. A. R. MELO

– Wing membrane orange yellow infumate; distal portion of fore wing membrane dark with brown microtrichia;
terga mostly yellow, slightly dark brown on posterior margin.................................................................11
11. Frons and mesoscutum mostly orange yellow; frons with one thin yellow stripe on mid line (Fig. 50); disc of
mesoscutum dark brown with lateral portions reddish orange (Fig. 62); margins of T4–T6 completely glabrous
(Bolivia; Brazil: Pará) ............................................................................................................... P. rufa
– Frons and mesoscutum mostly dark brown with thin yellow stripes; frons dark brown with one thin yellow stripe
along mid line; mesoscutum black with thin yellow stripes on disc and lateral margins (Figs 55–59); T4–T6 with
marginal hair bands occupying whole margin......................................................................................12
12. Mesoscutum with disc dark brown and lateral portions orange yellow; four thin yellow stripes on disc and lateral
margins, similar to P. duckei (Fig. 55); yellow stripe on paraocular area with upper portion largely obtuse (similar
to P. lineata, Fig. 49); maximum head width more than 2.5 mm (Brazil: Acre, Amapá, Amazonas)....P. camargoi
– Mesoscutum dark brown to black without orange yellow lateral portions, with only thin yellow stripes on the disc
and lateral margins (Fig. 57); yellow stripe on paraocular area with upper portion narrowly acute (similar to P.
flaveola, Fig. 47); maximum head width less than 2.5 mm (Costa Rica; Ecuador; Panamá) ............ P. chocoensis
13. Posterior margin of scutellum with dense white plumose pubescence laterally (Fig. 65) .............................. 14
– Posterior margin of scutellum with sparse dark brown plumose hairs laterally (Fig. 62).............................15
14. Labrum, clypeus, and paraocular area completely dark brown; mesoscutum with conspicuous thin yellow stripes
on disc (Costa Rica; Mexico).....................................................................................P. albilabris (Friese)
– Labrum yellow, clypeus with at least lower margin yellow, paraocular area with yellow stripe occupying more than
lower half of paraocular area; mesoscutum with faint yellow stripes on disc (Brazil: Goiás, Maranhão, Pará)....
.....................................................................................................................................P. albopilosa
15. T5 with marginal hair band occupying about one-third of margin laterally; frons with conspicuous yellow stripe
along the mid line; genal area with one yellow stripe on lower third (except in the specimen from Ecuador, in which
the stripe is absent) (Bolivia; Ecuador).....................................................................................P. andina
– T5 with margin completely glabrous, some specimens with few hairs along margin laterally; frons dark brown,
with faint yellow stripe along mid line; genal area completely dark brown (Brazil: Espírito Santo, Rio de Janeiro,
São Paulo) ........................................................................................................................ P. atlantica
16. Mesepisternum with one acute carina on omaular area (Fig. 5); margin of S3 mostly glabrous, with only two small
tufts of long hairs laterally (Fig. 15)..................................................................................................17
– Mesepisternum with omaular area convex, without carina; S3 with row of simple setae along margin (Figs 24–
26) .............................................................................................................................................. 19
17. Omaular carina extending over more than upper half of mesepisternum (Fig. 5); margin of S6 weakly sinuous on
apex with small rows of short hairs between the apical and anterolateral rows (Fig. 15) (Bolivia; Brazil: Amazonas,
Bahia, Distrito Federal, Goiás, Mato Grosso, Mato Grosso do Sul, Maranhão, Minas Gerais, Pará, Paraíba,
Pernambuco, Rondônia, Roraima, Tocantins; Costa Rica; Ecuador; Guatemala; Guyana; French Guiana; Panama;
Peru).....................................................................................................................P. connexa (Vachal)
– Omaular carina occupying less than upper half of mesepisternum; margin of S6 not sinuous with only apical and
lateral rows of hairs ....................................................................................................................... 18
18. Fore wing membrane dark brown infumate, some specimens with apical portion of wing slightly hyaline; T3–T5
with marginal hair bands occupying more than one-third of margins laterally; margin of T6 nearly glabrous, some
specimens with few hairs laterally (Bolivia; Brazil: Acre, Amazonas, Distrito Federal, Espírito Santo, Goiás, Mato
Grosso, Minas Gerais, Pará, Rondônia, São Paulo; Colombia; Costa Rica; Guatemala; Mexico; Panama; Peru;
Venezuela; Trinidad and Tobago)..............................................................................P. lugubris (Cresson)
– Fore wing membrane brown infumate on proximal third and white infumate on apical third; T4–T6 with marginal
hair bands occupying entire tergal margins (Brazil: Amapá; Guyana).........................................P. hyalinata
19. Frons biconvex; mid line strongly sulcated (Fig. 45) (Costa Rica; Mexico)........................................P. bifrons
– Frons convex; mid line not sulcated .................................................................................................. 20
20. Supraclypeal area strongly projected, with lateral portions almost carinate (Figs 8, 48); supraclypeal area, central
portion of frons and mesepisternum with dense very coarse punctures; terga with dense fine punctures; S2 with
tuft of short hairs on central portion (Fig. 26); margin of S3 mostly glabrous on mid portion or with two very small
rows of decumbent hairs (Brazil: Amapá, Bahia, Distrito Federal, Goiás, Maranhão, Minas Gerais, Paraná, Rio de
Janeiro, Rondônia, São Paulo) .............................................................................................. P. punctata
– Supraclypeal area convex; supraclypeal area, central portion of frons and mesepisternum with coarse punctures;
terga mostly smooth (except P. volatilis); S2 with continuous row of setae along margin, or with two small rows
of setae on mid portion; margin of S3 row continuous row of simple setae ............................................... 21
21. Terga black with yellow spots on lateral portions of T3–T5; area between disc of frons and upper portion
of paraocular area weakly concave; T5 with marginal hair band occupying about one third of margin laterally;

© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
 SYSTEMATICS OF THE GENUS PARATETRAPEDIA 371

margin of S2 with continuous row of setae, clustered on middle portion (Fig. 24); hind basitarsus with one long
digitiform tooth on anterior margin (Fig. 32).................................................................P. volatilis (Smith)
– Terga with homogeneous colour or yellow with marginal zone dark brown; area between disc of frons and upper
portion of paraocular area strongly concave; T5 with marginal hair band complete or completely absent; margin
of S2 with one row or two small rows of simple setae on middle portion; hind basitarsus without tooth on anterior
margin (except in P. romani)............................................................................................................22
22. Lower paraocular area with acute lamellate carina occupying more than lower third of paraocular area (Fig. 7);
mid basitarsus very slender, about three times longer than wide............................................................23
– Lower paraocular area without carina; mid basitarsus about two times longer than broad..........................25
23. Integument mostly orange yellow; pubescence mostly pale yellow; mandible with short simple setae on outer
margin, length of hairs less than width of mandibular base; scape short and very broad (c. two times longer than
broad; Fig. 52); S4 without tufts of long convergent hairs on lateral portions (Brazil: Amazonas, Acre; Ecuador;
Peru).................................................................................................................................P. scaposa
– Integument mostly dark brown; pubescence mostly brown; mandible with long hairs on outer margin, length
c. 1.5 ¥ width of mandibular base; scape c. four times longer than broad; S4 with tufts of long convergent plumose
hairs on lateral portions..................................................................................................................24
24. Mid basitarsus wider on apical portion than proximal portion; clypeus and supraclypeal area completely yellow;
paraocular area with one yellow spot occupying only lower half (Fig. 53); margin of S2 with row of erect hairs,
clustered on middle portion; S6 with a row of plumose hairs oblique to lateral margins, and one row of plumose
hairs on mid line of apical portion; S6 with surface convex, without tooth; body length usually more than 8.0 mm
(Costa Rica; Guatemala; Honduras; Mexico; Nicaragua) ................................................. P. moesta (Cresson)
– Mid basitarsus homogeneously long and thin (width less than one third of length); clypeus and supraclypeal area
varying from completely black to partially yellow, but never completely yellow (Fig. 51); yellow stripe on lower
paraocular area usually thin with upper portion acute; margin of S2 with continuous row of simple setae; S6 with
one apical projection on middle portion of apex (Fig. 16); body length less than 8.0 mm (Bolivia; Brazil: Amapá,
Goiás, Maranhão, Mato Grosso, Mato Grosso do Sul, Pará, Roraima; Guyana; French Guiana; Paraguay;
Suriname).......................................................................................................P. leucostoma (Cockerell)
25. Apical portion of scape normal or weakly dilated; S6 without two rows of stout plumose hairs along margins
laterally (Fig. 83) ........................................................................................................................... 26
– Apical portion of scape conspicuously dilated (Figs 9, 46); S6 with two rows of plumose hairs along margins
laterally (Fig. 81) ........................................................................................................................... 28
26. Integument mostly orange yellow (Brazil: Acre, Amapá, Amazonas, Goiás, Maranhão, Pará, Roraima, Tocantins;
Guyana; French Guiana; Peru; Suriname).....................................................................P. testacea (Smith)
– Integument mostly dark brown.........................................................................................................27
27. Genal area with one broad yellow stripe along eye margin; central portion of supraclypeal area yellow; omaular
angle slightly acute, almost carinate; surface behind omaular carina slightly concave (Colombia; Costa Rica;
Ecuador; Guatemala; Honduras; Mexico; Nicaragua; Panama; Venezuela).......................P. calcarata (Cresson)
– Genal area and supraclypeal area completely brown; mesepisternum with omaular angle obtuse, convex; surface
behind omaular angle convex (Brazil: Bahia, Espírito Santo, Minas Gerais, Paraíba, Pernambuco, São Paulo, Rio
de Janeiro) ............................................................................................................... P. bicolor (Smith)
28. Body length less than 9 mm; margin of T5 completely glabrous ............................................................. 29
– Body length more than 9 mm; margin of T5 with hair band occupying entire margin ................................ 30
29. Scape normal; margins of terga translucent hyaline, with weak silvery reflections (Brazil: Amapá, Amazonas,
Pará; Guyana; French Guiana) ............................................................................................. P. basilaris
– Apical portion of scape weakly dilated laterally; terga with integument homogeneous, without silvery reflections
along margin (Brazil: Acre, Mato Grosso; Peru) ........................................................................... P. vogeli
30. S2 with small row of setae on mid portion of margin; terga mostly orange yellow; outer margin of mandible, lower
portion of genal area, coxae, and trochanters with thin long pale hairs (length about two times width of
mandibular base); hind basitarsus with acute tooth on anterior margin (Brazil: Amazonas, Pará) ...................
..............................................................................................................................P. romani (Friese)
– S2 with two small row of setae on mid portion; outer margin of mandible, lower portion of genal area, coxa, and
trochanters with short hairs (length about same width as mandibular base); hind basitarsus without tooth on
anterior margin ............................................................................................................................ 31
31. Mesoscutum and gena completely orange yellow (Fig. 54) (Brazil: Amazonas; Peru)..........P. flavipennis (Smith)
– Mesoscutum black with lateral portions orange yellow (Fig. 56); gena dark brown with yellow stripe along eye
margin (Bolivia; Brazil: Acre, Amapá, Amazonas, Mato Grosso, Pará, Rondônia; Guyana; French Guiana; Peru;
Suriname).................................................................................................................P. duckei (Friese)

© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
372 A. J. C. AGUIAR and G. A. R. MELO
Figure 66. Geographical distribution of Paratetrapedia lugubris (Cresson) and P. hyalinata sp. nov.
Comments and diagnosis: Paratetrapedia lugubris
differs from P. connexa mainly by a short omaular
carina, less than upper half of mesepisternum; T5 of
female usually with yellow spots on lateral portions;
surface of terga of female with fine microsculpture of
lines forming minute circles and fine reticulated
stripes on marginal zones (Fig. 14). Paratetrapedia
lugubris differs from P. hyalinata by the wing mem-
brane being completely dark brown infumate and
T4–T5 with marginal hair bands occupying less than
one-quarter of margin laterally.
Variation: The yellow marks on the integument can
be completely absent or very conspicuous. A few male
specimens have frons completely smooth or with
punctures very sparse. Variation in distribution of
punctures on mesoscutum, between sparse (> 2 pd)
and dense (0.5 pd) occurs in specimens from distant
localities and specimens collected at the same place.
The omaular carina can vary in length and angle
(sharpness), from short and obtuse to long and acute,
but it never occupies more than the upper half of the
mesepisternum. Most specimens examined have the
pubescence on hind legs mostly black, with few white
plumose hairs on the lower third of tibia. Some speci-
mens from Espírito Santo (Brazil) have the pubes-
cence of hind tibia and basitarsus completely pale
yellow. This pattern of variation in colour of hind leg
pubescence and wing membrane is also observed in
other species of Tropidopedia and Paratetrapedia (e.g.
© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
P. connexa). Some variation in shape of the tooth of
the male hind basitarsus was also observed: speci-
mens from Costa Rica and Panamá have the tooth
almost at right angles to main margin with apex
weakly acute; specimens from Mato Grosso have it
weakly sinuous and apex long and obtuse; specimens
from Trinidad and Tobago and Venezuela have it
acute and long. Most of the male specimens examined
have margins of T5 and T6 completely glabrous;
however, some specimens from Mato Grosso, Trinidad
and Tobago, and Venezuela have short hair bands
with few hairs on less than one third of margins
laterally. Few specimens have apical margin of S8
completely glabrous.
Male. Body length: 11.0–12.3; maximum head
width: 3.2–3.4; fore wing length (including the
tegula): 11.2. Female. Body length: 11.0–12.5;
maximum head width: 3.8–4.0; fore wing length
(including the tegula): 10.8–11.7.
Distribution: Bolivia; Brazil: Acre, Amazonas, Distrito
Federal, Espírito Santo, Goiás, Mato Grosso, Minas
Gerais, Pará, Rondônia, São Paulo; Colômbia; Costa
Rica; Guatemala; Mexico, Panama; Peru; Venezuela;
Trinidad and Tobago (Fig. 66).
Type material
Tetrapedia lugubris Cresson; lectotype male (ANSP):
‘Mex.’ ‘Lectotype\ 2375’.
 SYSTEMATICS OF THE GENUS PARATETRAPEDIA
Tetrapedia bombitarsis Vachal; holotype male
(MHNP): ‘Holotype’ ‘Tetrap. 씹\ bombitarsis\ Vach.’
‘Museum Paris\ Coll. J. Vachal 1911’ ‘Tetrapedia\
bombitarsis\ Vachal’.
Tetrapedia amplipennis Smith; lectotype male
(BMNH), ‘Ega\\ 56\ 84’ ‘Paratetrapedia\
amplipennis\ (Sm)\ Pe. JS Moure 1972’.
Tetrapedia gigantea Friese; lectotype female (ZMB),
‘Brazil\ Jundiahy\ 2-1901’ ‘Tetrapedia\ gigantea\
씸 1908 Friese det.\ Fr.’; paralectotypes: 1 female
(ZMB), ‘Bolivia\ Tarata\ 1900’ ‘Tetrapedia\
gigantea\ 씸 1908 Friese det.\ Fr’ ‘Type’ ‘Zool.
Mus.\ Berlin’; 1 female (ZMB), ‘Bolivia\ Tarata\
1900’ ‘Tetrapedia\ gigantea\ 씸 1908 Friese det.\
Fr’ ‘Typus’.
Tetrapedia dentiventris Friese; lectotype male (ZMB),
‘Costa Rica\ San José\ 1913’ ‘Tetrapedia\ dentiven-
tris\ 씹 1920 Friese det.\ Fr.’; paralectotype female:
‘Costa Rica\ San José\ 1913’ ‘Tetrapedia\ dentiven-
tris\ F 1920 Friese det.\ Fr.’.
In MZSP there is one female specimen marked by
Schrottky as Tetrapedia gigantea with the following
labels: ‘12.868’ ‘Tetrapedia\ gigantea F\ Schrottky\
C. Schrottky det. 1910’ ‘96945’ ‘Paratetrapedia\
gigantea\ (Schrottky) F\ Pe. J.S.Moure det. 1988’;
information associated with number 12868 in the
register book of MZSP: ‘Tetrapedia (= gigantea Schrot-
tky) F; Anthophyla; det. Schrottky 1910; coll. O.
Dreher; Franca (SP)’. Under this record, above the
name gigantea is written fuliginosa Schrott. in the
hand of Ducke. This specimen was identified after the
description and thus cannot be a type.
Additional material
See Appendix.
PARATETRAPEDIA CONNEXA (VACHAL, 1909)
(FIGS 1, 3, 5, 12, 13, 14, 35, 67, 86, 117, 149, 150)
Tetrapedia connexa Vachal, 1909: 30; holotype male,
Bolivia: La Paz, Ampere (MNHP, examined).
Chalepogenus hypoleucus Cockerell, 1923a: 449; holo-
type female, Guyana: Pomeroon-Supenaam, Isso-
roro (BMNH, examined).
Tetrapedia mayarum Cockerell, 1912a: 30; holo-
type male, Guatemala, Izabal, Quirigua (AMNH,
examined).
Chalepogenus mayarum; Cockerell, 1914: 320. Lutz &
Cockerell, 1920: 562. Cockerell, 1942: 562.
Chalepogenus hypoleucus; Michener, 1942: 281.
Tetrapedia mayarum; Cockerell, 1946: 204.
Paratetrapedia lugubris; Michener, 1954: 115,
figs 68–70 (erroneous identification of P. connexa,
suggested by drawings of male genitalia).
Paratetrapedia connexa; Aguiar, 2007: 622.
© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
373
Comments and diagnosis: Paratetrapedia connexa is
very similar to P. lugubris in pattern of colour, pubes-
cence, and integument sculpture. The distribution of
hairs on sterna of the males is identical to that of P.
lugubris except for the apex of S6 weakly sinuous
with small rows of short hairs (Fig. 15). Paratetrape-
dia connexa differs from P. lugubris mainly in the
omaular carina, which occupies more than the upper
half of the mesepisternum. The female of P. connexa
also differs from P. lugubris by completely black terga,
without yellow marks laterally.
Variation: Paratetrapedia connexa exhibits colour
variation in wing membrane, sterna of male, and
pubescence of hind leg. Most of the specimens exam-
ined from the states of São Paulo and Minas Gerais
(Brazil) have wing membrane yellow hyaline with
microtrichia dark brown on apical portion. The speci-
mens from the states of Paraíba and Pernambuco
(Brazil), Peru, Costa Rica, Guatemala, and Panama
have the wing membrane black infumate. Specimens
from other regions have intermediate pattern of
colour of wing membrane, between yellow hyaline and
black infumate. A few specimens have the setae on
the first labial palpal segment not curved. The pubes-
cence on the hind leg can vary between completely
black to pale yellow. Most of the male specimens
examined have sterna completely black with small
yellow marks laterally. Some specimens from
Rondônia (Brazil) have S4–S6 completely yellow.
Some specimens from the south-western Amazon
basin are smaller than the majority of specimens
studied.
Male. Body length: 7.8–11.5; maximum head width:
2.7–3.4; wing length (including tegula): 8.1–11.2.
Female. Body length: 8.9–9.2; maximum head width:
3.0–3.7; fore wing length (including tegula): 8.0–9.4.
Distribution: Bolivia; Brazil: Amazonas, Bahia, Dis-
trito Federal, Goiás, Mato Grosso, Mato Grosso do
Sul, Maranhão, Minas Gerais, Pará, Paraíba, Per-
nambuco, Rondônia, Roraima, Tocantins; Costa Rica;
Ecuador; Guatemala; Guyana; French Guiana;
Panama; Peru (Fig. 67).
Type material: Tetrapedia connexa Vachal; holotype
(MNHP): ‘Bolivia\ Mapiri’ ‘씹’ ‘Holotype’ ‘Tetrap.\
connexa\ Vach.’ ‘Museum Paris\ Coll. J. Vachal 1911’
‘Tetrapedia\ connexa\ Vachal’.
Chalepogenus hypoleucus Cockerell; holotype
(BMNH): ‘Type’ ‘B.M. Type’\ Hym.\ 17.B.881’ ‘Chale-
pogenus\ hypoleucus\ Ckll. Type’ ‘Issororo\ N.W.D.\
B. Guiana\ June 1915’ ‘Press. by\ Imp. Bur. Ent.\
Brit. Mus.\ 1923-21’ ‘G.E. Brodkin\ Collector’.
374 A. J. C. AGUIAR and G. A. R. MELO
Figure 67. Geographical distribution of Paratetrapedia connexa (Vachal).
Tetrapedia mayarum Cockerell; holotype (AMNH):
‘Quirigua\ Guatemala\ (W.P. Ckll.)’ ‘Tetralonia
pedia\ mayarum\ Ckll. Type’ ‘Ac 33583’.
Additional material: See Appendix.
PARATETRAPEDIA HYALINATA SP. NOV.
(FIGS 66, 77, 87, 118, 151, 152)
Comments and diagnosis: Paratetrapedia hyalinata is
very similar to P. lugubris with regard to body size,
colour, and shape. Paratetrapedia hyalinata possesses
an omaular carina over less than upper half of
mesepisternum, and the same pattern of pubescence
as on sterna of P. lugubris. Paratetrapedia hyalinata
differs from P. lugubris mainly by T4–T6 of males and
females with marginal band of plumose hairs
throughout the entire margin, and fore wing mem-
brane dark brown with apical portion white infumate.
The holotype is partially damaged, with margins of
fore wings slightly torn, and most of pubescence of
head and mesosoma absent.
Description
Holotype male: Body length: 10.5; maximum head
width: 3.2; fore wing length (including tegula): 9.9.
Colour. Integument mostly black. Mandible with one
yellow spot on base; labrum and clypeus with yellow
marks on lateral margins; scape reddish brown, with
basal portion yellow; pedicel and flagellomeres
© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
reddish brown; genal area with one thin yellow stripe
on lower portion. Wing membrane mostly dark brown
infumate; fore wing membrane with apical third
white infumate; veins and pterostigma dark brown;
microtrichia dark brown. Tibial spurs dark brown.
Pubescence. (pubescence of head and mesosoma
mostly absent). Pubescence mostly dark brown.
T4–T6 with dense marginal band of plumose hairs on
entire margin; S2 with continuous row of simple
setae; margin of S3 glabrous on mid portion, with
long plumose hairs laterally. Legs with pubescence
mostly black, with few sparse white plumose hairs on
the distal half of the hind tibia. Integument sculp-
ture. Clypeus with sparse coarse punctures (1–2 pd)
intermingled amongst sparse minute punctures
(> 2 pd); supraclypeal area with central portion
mostly smooth, with some coarse punctures laterally
(1–2 pd); disc of frons with heterogeneous punctures,
with sparse coarse punctures (2 pd) intermingled
amongst fine sparse punctures (1–3 pd); antennal
scrobe with dense fine punctures (< 1 pd). Mesoscu-
tum and scutellum with dense fine punctures (1 pd),
intermingled amongst sparse coarse punctures
(> 1 pd); metapostnotum with heterogeneous punc-
tures, contiguous coarse punctures (< 0.5 pd) inter-
mingled amongst dense fine punctures (1 pd);
metapostnotum with fine lamellas adjacent to margin
of metanotum; mesepisternum with dense coarse
punctures laterally (0.5–2 pd). Structure. Lamella of
pronotal collar acute throughout its length with
 SYSTEMATICS OF THE GENUS PARATETRAPEDIA
lateral portions diverging from mesoscutum. Scutel-
lum strongly biconvex, mid line sulcate. Hind basi-
tarsus with acute tooth on anterior margin. Head
about 1.18¥ broader than long (2.7 : 3.2); ratio of
lower to upper interocular distances: 0.8 (1.4 : 1.8);
clypeus about 1.5¥ broader than long (1.45 : 0.95);
scape: length 0.91; maximum width 0.25; length
F1–F3: 0.23, 0.16, 0.25; diameter of F2: 0.21.
Paratype female: Body length: 11.1; maximum head
width: 3.8; wing length (including tegula): 10.5.
Colour. Similar to male, except for absence of yellow
marks on terga and sterna. Pubescence. Mostly dark
brown, except margins of sterna with pubescence pale
white; mesoscutum and scutellum with very short
dark brown pubescence; scutellum with one erect
setae laterally (c. 0.14 mm in length); T4–T6 with
marginal hair band throughout margin. Legs pubes-
cence mostly black, except hind basitarsus with
pubescence mostly white. Integument sculpture.
Punctuation denser and finer than on males; clypeus
and supraclypeal area with dense coarse punctures
(0.5–2 pd); frons with dense fine punctures (0.5–1 pd),
with few sparse coarse punctures on disc; mesoscu-
tum and scutellum with dense minute punctures
(0.5–1 pd), intermingled amongst sparse coarse punc-
tures (0.5–3 pd); metapostnotum with dense to con-
tiguous minute punctures (< 0.5 pd), intermingled
amongst coarse punctures on central portion (0.5–
2 pd). Mesepisternum laterally with dense coarse
punctures (0.5–1 pd), intermingled amongst fine
punctures (0.5–2 pd). Vertical surface of T1 with
microsculpture of dense minute punctures; disc of
T2–T6 with microsculpture of fine lines forming
minute circles; marginal zone of terga with microscu-
lpture of fine reticulated lines. Structure. Similar to
male, except that scutellum weakly biconvex. Head
about 1.3¥ broader than long (3.8 : 2.9); ratio of lower
to upper interocular distances: 0.88 (1.95 : 2.2);
clypeus about 1.8¥ broader than long (1.75 : 0.95);
scape: length 1.1, maximum width 0.25; length of
F1–F3: 0.32, 0.17, 0.22; diameter of F2: 0.27.
Etymology: The species name refers to the milky
hyaline apex of the fore wing membrane.
Distribution: Brazil: Pará; French Guiana (Fig. 66).
Type material: Holotype male (ZMB), ‘Brazil\ Para\
30.6.1900\ Ducke’ ‘Tetrapedia\ basalis\ Fri.\ det. H.
Friese 1924’. Paratypes: 1 female (SEMC), ‘French
Guiana\ 65 km, S. Cayenne\ 23 February 1977\ C.D.
Michener, G.\ Otis, M. Winston’; 1 female (SEMC),
© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
375
‘R. Cumina-mirim\ (Trombetas)\ 13-10-’13 Ducke’
‘Brazil\ Estado do Pará’ ‘Tetrapedia\ basalis\ Fri.\
det. H. Friese 1924’.
THE MOESTA SPECIES GROUP
Species of the moesta group are small to medium
sized bees (6.0–9.0 mm of body length), characterized
by integument orange yellow or black, surface
between disc of frons and upper paraocular area
weakly concave, almost flat (Figs 7, 11; character 7:1);
T4 of female with marginal band of hairs restricted to
about one-quarter of margin laterally (character
40:0); margins of S2–S3 of males with continuous row
of setae, without gaps; lamellate projection of gono-
stylus with acute apex ventrally (character 53:1; beg;
Figs 147, 153, 157); spatha about two times broader
than long (character 57:1). Except for P. volatilis, the
species of the moesta group possess the following
characteristics: prementum of female with numerous
long stout wavy setae (character 3:1); lower paraocu-
lar area with thin lamellate carina along ocular
margin (Fig. 7; character 14:1); T4 of males with
marginal hair band complete (character 41:1); meta-
postnotum and mesepisternum with fine sparse punc-
tures (> 3 pd); mid basitarsus of male narrow;
apodeme of gonapophysis projected medially below
the margin of gonocoxite ventrally (ea; character 56:0;
Fig. 157). Species of the moesta group possess a very
distinct terminalia, mainly the seventh sternum and
some features of gonostylus and spatha that are
highly differentiated from those of the remaining
species of Paratetrapedia. The seventh sternum of the
males of P. moesta, P. scaposa, and P. leucostoma has
the apical portion reduced (Figs 89–91), not lamellate
as in the remaining species of Paratetrapedia. The
genitalia is mainly distinct because of its elongate
spatha; also the gonostylus has one lateral acute
projection on proximal portion (basal expansion of
gonostylus, beg, character 53:1; Figs 153, 157, 169).
The distribution of the species of the moesta group is
mostly allopatric (Fig. 69): P. moesta is distributed in
Central America, P. scaposa in the western Amazon,
P. leucostoma mainly in the eastern Amazon, and P.
volatilis in the Atlantic Forest of south-eastern Brazil.
PARATETRAPEDIA MOESTA (CRESSON, 1878)
(FIGS 53, 69, 90, 120, 157, 158)
Tetrapedia moesta Cresson 1878: 135; lectotype,
Mexico (ANSP, examined), designated by Cresson,
1916: 124.
Tetrapedia maesta [sic]; Dalla Torre, 1896: 300. Cock-
erell, 1899: 16. Friese, 1899: 288. Ducke, 1910a:
369. Cockerell, 1912b: 31 [floral record: Pontederia
cordata (Pontederiaceae)].
376 A. J. C. AGUIAR and G. A. R. MELO
Tetrapedia moesta; Cockerell, 1906: 98. Cockerell,
1912b: 282.
Chalepogenus maesta [sic]; Cockerell, 1923a: 451.
Cockerell, 1923b: 4.
Chalepogenus moestus [sic]; Cockerell, 1914: 320.
Lutz & Cockerell, 1920: 562. Cockerell (1932): 12
[floral record: Tamonea curasavica (Verbenaceae)].
Chalepogenus moesta; Cockerell, 1923b: 4.
Chalepogenoides moestus; Michener, 1942: 281.
Paratetrapedia (Lophopedia) maesta [sic]; Neff &
Simpson, 1981: 111, figs 35–36 (morphology of the
oil collecting apparatus).
P. moesta; Heithaus, 1979: 195.
Paratetrapedia maesta [sic]; Simpson & Neff, 1981:
316; Simpson, 1989: 31 (oil flower record: Krameria
ixine).
Paratetrapedia (Paratetrapedia) moesta; Ayala et al.,
1996: 461. Ayala, 1988: 403; Yañez-Ordoñez &
Hinojosa-Diáz, 2004: 193.
Paratetrapedia moesta; Aguiar, 2007: 624.
Comments and diagnosis: Paratetrapedia moesta is
the only species of the moesta group found in Central
America. The male of P. moesta has, similarly to P.
leucostoma, the labrum yellow, clypeus partially or
completely yellow, supraclypeal area yellow, paraocu-
lar area with wide yellow stripe, usually on its lower
third, scape yellow (Fig. 53), posterior margin of S2
with cluster of setae on mid portion. Paratetrapedia
moesta differs from P. leucostoma by larger body size;
basitarsus of mid leg of male wider in apical third
than in proximal third; S6 of male with dense rows of
stout plumose setae oblique to lateral margins and
along mid line of apical portion, wing membrane
black infumate. The female possesses a completely
black integument, without yellow marks.
Friese (1899) and Cockerell (1906) suggested that
P. moesta is synonymous with Tetrapedia atripes
Smith 1854. The holotype of T. atripes was examined
for the present work and the species truly belongs in
Tetrapedia.
Variation: Some specimens of P. moesta are quite
small. The wing membrane can vary between com-
pletely black infumate, with dense black microtrichia,
and black infumate with apical third white infumate.
A geographical pattern in the variation of these two
characters was not observed, with specimens from a
single locality presenting variation in both body size
and wing colour.
Male. Body length: 6.7–10.0; maximum head width:
2.6–3.5; fore wing length (including the tegula): 7.1–
9.4. Female. Body length: 6.0–9.3; maximum head
width: 2.35–3.0; fore wing length (including tegula):
6.5–8.6.
© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
Distribution: Costa Rica; Guatemala; Honduras;
Mexico: Campeche, Chiapas, Jalisco, Michoacán,
Morelos, Oaxaca, San Luís Potosí, Veracruz, Yucatan;
Nicarágua (Fig. 69).
Type material: Tetrapedia moesta Cresson; lectotype:
‘Mex.’ ‘Lectotype\ 2376’; paralectotype: ‘Mex’ ‘씹’
‘Allotype\ 2376’.
Additional material: See Appendix.
PARATETRAPEDIA VOLATILIS (SMITH, 1879)
(FIGS 11, 24, 32, 88, 119, 153, 154)
Tetrapedia volatilis Smith, 1879: 128; holotype male,
Brazil: Rio de Janeiro, Teresópolis [‘Constancia’]
(BMNH, examined).
Tetrapedia maculata Friese, 1899: 291; lectotype
male, Brazil (NHMV, examined), designated by
Aguiar (2007): 625.
Tetrapedia fuliginosa Schrottky, 1902: 551; lectotype
male, Brazil: São Paulo, Botucatu (MZSP, exam-
ined), designated by Aguiar (2007).
Tetrapedia bimaculata Schrottky, 1902: 547;
holotype male, Brazil: São Paulo, Jundiaí (MZSP,
examined).
Tetrapedia volatilis; Dalla Torre, 1896: 300.
Tetrapaedia [sic] volatilis; Schrottky, 1902: 552.
Tetrapaedia [sic] maculata; Schrottky, 1902: 550.
Paratetrapedia maculata; Moure, 1941: 518; Moure,
1944b: 109. Sazima & Sazima, 1989: 107 [floral
record: Heteropterys aceroides (Malpighiaceae)].
Wittmann & Hoffman, 1990: 25. Roig-Alsina, 1997:
4 (terminal taxon in phylogenetic analyses).
Paratetrapedia (Paratetrapedia) maculata; Michener
& Moure, 1957: 416; Gonçalves & Melo, 2005: 564.
Silveira et al., 2002: 136.
Paratetrapedia (Paratetrap.) [sic] maculata; Alves-
dos-Santos, 1999: 204.
Paratetrapedia volatilis; Gonçalves & Melo, 2005:
560; Aguiar, 2007: 625.
Comments and diagnosis: Paratetrapedia volatilis
differs from the remaining species of the moesta
group by absence of stout setae with wavy apex in the
female prementum, and absence of carina on lower
portion of paraocular area. Paratetrapedia volatilis is
similar to P. lugubris by yellow marks on terga lat-
erally, and is distinct from this species by omaulus
convex, not carinate.
In its area of distribution, P. volatilis can also be
easily confused with P. fervida because of the very
similar pattern of structure, integument sculpture,
size, and colour. The female of P. volatilis differs from
P. fervida mainly by first labial palpi with few curved
setae, area between disc of frons and paraocular area
 SYSTEMATICS OF THE GENUS PARATETRAPEDIA
weakly concave, post-ocellar carina with lateral por-
tions weakly acute, T4 with marginal band of hairs
occupying about one third of margin laterally, and
T3–T5 usually with yellow marks laterally. The male
of P. volatilis is easily distinguished from P. fervida by
the presence of a long digitiform tooth in the anterior
margin of the hind basitarsus, row of setae on poste-
rior margin of S2 clustered on middle portion, S3 with
simple continuous row of setae (P. fervida presents S3
deeply concave with dense plumose hairs), and apical
portion of S6 widely rectangular.
Variation: Females have T3–T5 with small yellow
marks laterally or terga entirely black without
marks. Males have the marginal hair band on T5
throughout the entire margin or restricted to lateral
one-third.
Male. Body length: 9.0–10.0; maximum head
width: 2.7–3.4; fore wing length (including tegula):
9.0–11.0. Female. Body length: 8.3–11.0; maximum
head width: 3.2–3.5; fore wing length (including
tegula): 8.6–10.0.
Distribution: Argentina; Brazil, Bahia, Espírito
Santo, Minas Gerais, Paraná, Rio de Janeiro,
Rio Grande do Sul, Santa Catarina, São Paulo
(Fig. 69).
Type material: Tetrapedia volatilis, holotype male
(BMNH), ‘Type\ H.T.’ ‘B.M. Type\ Hym.\ I.M.B. 891’
‘Tetrapedia\ volatilis\ (Type) Sm.’ ‘Constancia\
January 1857\ J. Gray\\ 5757’.
Tetrapedia maculata, lectotype male (NHMV):
‘Beske Bras. 848’ ‘T. maculata M det. Friese 1898’;
paralectotype male (NHMV): ‘Beske Bras. 848’ ‘Tetra-
pedia maculata det. Friese 1898’. In the ZMB collec-
tion, there is one male specimen of P. lugubris with
undetermined label of collecting locality, identified by
Friese as P. maculata, which could be from the type
series: ‘Rio\ vs. Offers I’ ‘488’ ‘Tetrapedia 씹\ macu-
lata\ det. Friese 1898 n.sp.’ ‘Type’ ‘13’.
Tetrapedia fuliginosa Schrottky 1902, lectotype
male (MZSP): ‘17.842’ ‘fuliginosa’ ‘96935’; information
related to record number ‘17842’ in accession book:
‘Tetrapedia fuliginosa Schross.; det. Ducke 13; coll.
Hempel; Vict. Botucatú’.
Tetrapedia bimaculata Schrottky 1902; holotype
male (MZSP): ‘17.840’ ‘씹’ ‘96914 fuliginosa’; informa-
tion related to number ‘17.840’ in accession book:
‘xi.1900; Tetrapedia maculata Fr. (= bimaculata
Schoss. 씹 Type!) F, Anthophila; Ducke 13 det.;
Schrottky; Jundiahy (S.P.)’.
Additional material: See Appendix.
© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
377
PARATETRAPEDIA LEUCOSTOMA (COCKERELL,
1923a) (FIGS 7, 16, 51, 69, 89, 121, 169, 170)
Chalepogenus leucostoma Cockerell, 1923a: 450; lec-
totype male, designated by Aguiar (2007), Guyana:
Pomeroon-Supenaam, Issororo (ZMB, examined).
Chalepogenoides leucostoma; Michener, 1942: 279.
Paratetrapedia (Paratetrapedia) leucostoma;
Michener & Moure, 1957: 416.
Paratetrapedia sp. 1; Alves-dos-Santos, 2003: 260,
figs 11–12.
Paratetrapedia leucostoma; Aguiar, 2007: 623.
Comments and diagnosis: Paratetrapedia leucostoma
is easily differentiated from the remaining species of
the moesta group by small body size, brown integu-
ment and pattern of male pubescence. The males
have long simple setae on proximal portion of man-
dibles, S2 with a simple row of stout setae, and
bands of setae throughout the margins of T4–T6.
Paratetrapedia leucostoma is distinct from P.
scaposa by dark brown integument, and from P.
moesta by frons with dense fine punctures (< 1 pd)
and body size smaller (about 8.0 mm). The male of
P. leucostoma is easily identified by S6 with one
conspicuous projection on middle portion (Fig. 16),
and mid basitarsus very thin, about three times
longer than broad.
Variation: The colour of tibial spurs can vary between
black to almost white. The male specimens from
southern Brazil and Paraguay have a dark integu-
ment, mostly black, without yellow marks in the
head. The male specimens from the northern portion
of the distribution (Brazil: Maranhão, Pará; Suri-
name) have the terga completely yellow and large
yellow marks on the face. One specimen from Mara-
nhão possesses the clypeus, lower paraocular area,
and supraclypeal area completely yellow, and mesos-
cutum black with thin yellow stripes on disc.
Male. Body length: 5.7–7.6; maximum head width:
2.3–2.5; fore wing length (including tegula): 5.6–6.5.
Female. Body length: 6.4–8.2; maximum head width:
2.4–2.7; fore wing length (including tegula): 6.6–6.9.
Distribution: Bolivia; Brazil: Amapá, Goiás, Mara-
nhão, Mato Grosso, Mato Grosso do Sul, Pará,
Roraima; Guyana; French Guiana; Paraguay; Suri-
name (Fig. 69).
Type material: Chalepogenus leucostoma Cockerell,
lectotype female (MBNH): ‘Type’ ‘B.M. Type\ Hym\
17B.889’ ‘Chalepogenus\ leucostoma\ Ckll. Type’
‘B. Guiana\ Issidoro N.W.D.\ -XII.19\ G.E. Brodkiw
coll.’ ‘Press. by.\ Imp. Bur. Ent.\ Brit. Mus\ 1923-21’.
Additional material: See Appendix.
378 A. J. C. AGUIAR and G. A. R. MELO
PARATETRAPEDIA SCAPOSA SP. NOV.
(FIGS 52, 60, 61, 69, 79, 91, 122, 197, 198)
Comments and diagnosis: Paratetrapedia scaposa is
distinguished from the remaining species of the
moesta group by its mostly orange yellow integument
and frons with dense fine punctures (< 1 pd). The
male of P. scaposa is easily recognized by the short
and wide antennal scape (Fig. 52).
Distribution: Bolivia; Brazil: Acre, Amazonas;
Ecuador; Peru (Fig. 69).
Description
Holotype male: Body length: 6.1; maximum head
width: 2.1; wing length (including tegula): 6.0. Colour.
Integument mostly orange yellow. Mandible yellow
with apex black; disc of frons, upper paraocular area
and vertex black; scape yellow, pedicel and flagellom-
eres reddish brown (Fig. 52). Mesoscutum orange
yellow with two thin yellow stripes on disc and lateral
margins; scutellum yellow (Fig. 61); axilla dark brown
laterally; propodeum spiracle with margins black.
Terga orange yellow; tergal disc pale yellow, marginal
zone dark brown. Wing membrane dark brown infu-
mate, with dense dark brown microtrichia; veins and
pterostigma orange yellow. Tibial spurs pale white.
Pubescence. Mostly pale yellow, except marginal
bands of hairs on T2–T5 and hind leg with dark
brown pubescence. Scape without long conspicuous
setae. Mesoscutum and scutellum with dense short
pubescence golden yellow; scutellum with simple
erect setae, c. 0.08 mm in length; metapostnotum
with fine plumose hairs, c. 0.2 mm in length. T4–T6
with marginal hair bands throughout the margin; T6
with band of dense plumose golden yellow hairs.
S2–S3 with marginal row of long simple setae; row of
setae on S3 with short interruption on mid portion;
S4 with marginal band of plumose hairs, not forming
long curved fringes laterally. S6 with two dense rows
of stout setae on lateral portions of margin, almost
contacting one another; margin of apical portion with
short plumose hairs. Integument sculpture. Clypeus
and supraclypeal area with dense coarse punctures
(< 0.5 pd), intermingled amongst sparse minute punc-
tures (2 pd); frons with dense fine punctures (0.5–
1 pd). Mesoscutum and scutellum with dense fine
punctures (0.5–1 pd), intermingled amongst slightly
coarser sparse punctures (> 2 pd); metapostnotum
with sparse minute punctures (> 3 pd); mesepister-
num with sparse fine punctures laterally (> 3 pd).
Structure. Lower paraocular area with acute lamel-
late carina on ocular margin. Surface between upper
paraocular area and disc of frons slightly concave.
Scape very wide, about three times longer than broad.
Lamella of pronotal collar thin and acute throughout
© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
its length, with lateral portions low. Disc of scutellum
biconvex, with mid line concave. Mid basitarsus
slender and long, about three times longer than
broad. Head about 1.3¥ broader than long (2.1 : 1.6);
ratio between lower interocular distance and upper
interocular distance: 0.85 (0.95 : 1.15); clypeus about
1.6¥ broader than long (0.8 : 0.5). Scape: length 0.5,
maximum width 0.24; length of F1–F3: 0.08, 0.16,
0.21; diameter of F2: 0.16.
Paratype female: Body length: 7.2; maximum head
width: 2.4; fore wing length (including tegula): 6.7.
Colour. Mostly orange yellow, similar to male, except
that mesoscutum dark brown with two thin yellow
stripes on disc and lateral margins. Pubescence.
Similar to male, except that marginal hair band on T4
on less than one third of margin laterally; T5–T6 with
marginal hair band occupying throughout margin.
Prementum with numerous long curved setae with
apex wavy. Integument sculpture. Similar to male,
but denser and finer; frons, paraocular area and
vertex with dense fine punctures (< 0.5 pd). Structure.
Similar to male, except that mid basitarsus less than
three times longer than broad. Head about 1.2¥
broader than long (2.4: 2.0); ratio between interocular
lower distance and interocular upper distance: 0.78
(1.1 : 1.4); clypeus about 1.8¥ broader than long;
scape: length 0.67; maximum width 0.16; length of
F1–F3: 0.19, 0.10, 0.12; diameter of F2: 0.17.
Variation: Most specimens examined have integu-
ment orange yellow, but some specimens from differ-
ent localities have frons and mesoscutum partially or
completely dark brown (Figs 60–61).
Etymology: The species name refers to the enlarge-
ment of the male antennal scape.
Type material: Holotype male, ‘DZUP\ 022591’
‘Cruzeiro do Sul\ Acre-Brazil II-63\ M. Alvarenga’.
Paratypes: BOLIVIA: 1 male (ASC), ‘Bolivia: Beni\
Rurrenabaque, 3.5 km N\ 14°26′03′S 67°29′58W\ 10
July 1998\ K. B. Miller colr’, 1 male (ASC), idem
except ‘mimic of\ Trigona\ chancham\ ayensis’; 1
male and 1 female (SEMC); ‘Bolivia Beni\ Rurrena-
baque\ 175 m., 1/10\ October 1956’; 1 female,
‘DZUP\ 021922’ ‘Coleção\ Campos Seabra’ ‘Riber-
alta\ Pando-Bolívia\ XI-956, Fritz’; BRAZIL: Acre: 1
male, ‘DZUP\ 023547’ ‘Cruzeiro do Sul\ Acre-Brazil
II-63\ M. Alvarenga’, 4 males and 2 females (DZUP),
idem except ‘022595’, ‘022701’, ‘022699’, ‘022594’,
‘022596’; 1 male (RPSP), ‘Brazil-Acre\ RBr-UFAC/
PZ.\ Data 08/09/1995’ ‘A.H. Machado\ E.M. Santos’
‘1348’; 1 female (RPSP), ‘Brazil-Acre\ Rio Branco
UFAC’ ‘Data 05/IX/1995\ M.L. Oliveira’; 2 females
(INPA), ‘Brazil, Acre\ Rio Branco,\ 09°58′S67°48′W\
 SYSTEMATICS OF THE GENUS PARATETRAPEDIA 379

25/IV/2002’ ‘Em lanterneira\ (Lophantera\ lacte- 1946\ Alt. 670 m.’ ‘W. Weyrauch\ Coll. Donor\ Wm.
scens)\ Oliveira et al. leg.’; 1 female (INPA), ‘Brazil- Proctor’; Locality unknown: 1 male (INHS), ‘INHS\
Acre\ Rio Branco\ 01/maio/2001’ ‘Em lanterneira\ Insect Collection\ 53.055’.
(Lophantera\ lactescens)\ Oliveira et al. leg.’ ‘Paratet-
rapedia’ ‘P. (Paratetrapedia)\ testacea\ (Smith, THE BICOLOR SPECIES GROUP
1854)’; 1 male (RPSP), ‘RIO BRANCO-AC\ BRAZIL\
The species of the bicolor group have a small body
20/05/1993\ T.D. Tojal; Amazonas: 6 males and 1
size (body length about 6.0–8.0 mm), integument
female (RPSP), ‘Benjamin Constant\ AM. Brazil,
mostly dark brown (except P. testacea with integu-
SB-19,70-4c’ ‘13,14-I-1977, Camargo\ M.Mazucato
ment orange yellow); mesoscutum with two pale
leg.’; 1 male (RPSP), ‘Carauari, r. Juruá-AM\ Brazil,
yellow stripes (on some species these stripes are faint,
20-24.07.1993\ 66°54′W, 4°53′S/933169’ ‘Camargo,
almost inconspicuous); pronotal collar with lamella
Pedro\ Mazucato, leg.’, 1 male and 1 female (RPSP),
acute throughout its length and pronotal lobe trian-
idem except ‘932711’, ‘932954’; 1 female (RPSP),
gular with apical portion conspicuously acute; male
‘Sta. Eutália, r. Juruá-AM\ Brazil. 25-26. 07.1993\
antennal scape with apical portion dilated laterally
66°35′W, 4°31′S\ 433166’ ‘Camargo, Pedro\
(Figs 9, 46; except P. bifrons and P. basilaris; charac-
Mazucato, leg.’; 3 males (RPSP), ‘Fonte Boa. AM.
ter 8:1); margin of male T4 mostly glabrous (character
Brazil\ SA-19,66-3f.XII’ ‘24,25-I-1977, Camargo\
49:0); posterior margins of male T5 and T6 completely
M.Mazucato leg.’; 1 female (RPSP), ‘São Paulo de
glabrous (character 42: 0); male pygidial process short
Olivença\ AM.Brazil.SA-19,69-4b’ ‘19,20-I-1977,
and obtuse, almost absent; gonapophysis of male
Camargo\ M.Mazucato leg.’; 1 female, ‘DZUP\
genitalia with inner lamella forming an oval lobe.
022588’ ‘TEFE Amazonas\ Brazil XII-61\ F.M.
Males of P. calcarata, P. testacea, and P. bicolor
Oliveira’, 4 males (DZUP), idem except ‘022589’,
have the S6 without tufts of plumose hairs on lateral
‘022590’, ‘022592’, ‘022593’; 1 male (RPSP), ‘Vendava-
margins (character 49:0) and S7 with the apical
l.AM.Brazil\ SA-19,69-3h-VI’ ‘16,18-I-1977, Cama-
portion strongly acute laterally. Males of the bicolor
rgo\ M. Mazucato leg.’; ECUADOR: 1 male (AMNH),
group are similar to males of the moesta group in
‘Ecuador, Coca\ on Rio Napo, Napo\ Pastaza Prov.\
S2–S3 with continuous row of setae along margin and
V-1965’ ‘L.E.Pena\ Collector’; 3 males (SEMC),
hind basitarsus without tooth on anterior margin.
‘Ecuador Oriente\ 00°24′S, 76°36′W\ Limoncocha\
Except for P. testacea, which is widely distributed in
26 July 1970\ M.G. Naumann, #’; 1 male and
the Amazon basin, the remaining species of the
4 females (SEMC), ‘Ecuador Oriente\ 00°24′S,
bicolor group have allopatric distributions (Fig. 68): P.
76°36′W\ Limoncocha\ 25 July 1970\ M.G.
bifrons has a few records in Central America; P.
Naumann, #’; 1 female (SEMC), ‘Ecuador: Napo\
calcarata occurs in Central America and in areas
Coca, Rio Napo, 270 m\ 0°28′48′S,77°0′24′W\ 25
western of the Andes in Ecuador, Venezuela, and
MAR 1999’; R.Brooks\ ECU1B99 034\ ex: misc. col-
Colombia; P. basilaris occurs in the northern Amazon;
lecting’, ‘SM0 158889\ KUNHM-ENT’ ‘Paratetrape-
P. vogeli occurs in the western Amazon; and P. bicolor
dia\ det. R.W. Brooks 19’; 1 female (SEMC), ‘Ecuador:
occurs in the Atlantic Forest of south-eastern and
Sucumbios\ Sacha Lodge, 0.5°S\ 76.5°W, 270 m,
north-eastern Brazil, from the state of Rio de Janeiro
22-II-4-III\ 1994, Hibbs, ex: malaise’; PERU: 1 male
to Pernambuco.
(INHS), ‘INHS\ Insect Collection\ 53.054’ ‘Pilco-
pata\ Cuzco II-X-68’; 2 males (CLAUS), ‘PERU,
LO, Maynas,\ Quistococha 120 m asl\ S0350/ PARATETRAPEDIA BICOLOR (SMITH, 1854)
W7319.VII.2004\ Claus Rasmussen leg.’; 1 male (FIGS 64, 68, 83, 95, 124, 161, 162)
(SEMC), ‘Peru: Huanuco Department.\ Rio Tetrapedia bicolor Smith, 1854: 366; holotype female,
Pumahuasi, 930 m\ 9°11′33′S, 75°57′24′W\ 11 OCT Brazil: Bahia (BMNH, examined).
1999\ R. Brooks\ D. Bizoska, PERU1B99004’ Tetrapedia nasuta Smith, 1854: 366; holotype male,
‘SM0147509\ KUNHM-ENT’; 1 male (AMNH), ‘Peru, Brazil: São Paulo, Santos (BMNH, examined).
Quincemil,\ on branch R.Manu\ Madre de Dios Tetrapedia bicolor; Dalla Torre, 1896: 299. Friese,
Prov.\ X-16-31-1962’ ‘L.E.Pena\ Collector’; 1 female 1899: 294.
(AMNH), ‘Tingo Maria\ Huan. Peru\ oct. 14.1946\ Tetrapedia nasuta; Dalla Torre, 1896: 300. Friese,
Alt. 2200 ft.’ ‘J.C. Pallister\ Coll. Donor\ Frank 1899: 295. Ducke, 1901: 50 (misidentification, based
Johnson’ ‘A.M.N.H.’; 2 males (AMNH), ‘Tingo Maria\ on geographical distribution).
Huan. Peru\ oct.6.1946\ Alt. 2200 ft.’ ‘J.C. Pallister\ Tetrapaedia [sic] bicolor; Schrottky, 1902: 553.
Coll. Donor\ Frank Johnson’; 1 male (AMNH), ‘Tingo Tetrapaedia [sic] nasuta; Schrottky, 1902: 553.
Maria\ Huan. Peru\ oct.23.1946\ Alt. 2200 ft.’ ‘J.C. Paratetrapedia bicolor; Moure, 1941: 518 (misidenti-
Pallister\ Coll. Donor\ Frank Johnson’ ‘A.M.N.H.’; 1 fication, based on geographical distribution). Aguiar
female (AMNH), ‘Tingo Maria\ Huan. Peru\ October (2007): 623.

© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
380 A. J. C. AGUIAR and G. A. R. MELO

Figure 68. Geographical distribution of the species of the bicolor group.

Figure 69. Geographical distribution of the species of the moesta group.

Paratetrapedia (Lophopedia) bicolor; Michener &
Moure, 1957: 415.
Paratetrapedia (Paratetrapedia) nasuta; Albu-
querque & Rego, 1989: 168; Rego & Albuquerque,
1989: 185 (misidentification of P. leucostoma and P.
albopilosa).
Comments and diagnosis: The female of P. bicolor
differs from other brown species of the bicolor group
mainly by terga with microsculpture of thin reticu-
lated lines (similar to P. connexa; Fig. 14) and T1 with
dense minute punctures. The male of P. bicolor is
distinct from P. vogeli and P. basilaris by antennal

© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
 SYSTEMATICS OF THE GENUS PARATETRAPEDIA
Figure 70. Geographical distribution of Paratetrapedia fervida and Paratetrapedia punctata sp. nov.
scape strongly dilated laterally on apical portion and
S6 without lateral rows of plumose setae on margin
(Fig. 83).
Paratetrapedia bicolor is structurally identical to P.
testacea. These species are mainly distinct in the
colour of integument and pubescence and geographi-
cal distribution. Paratetrapedia bicolor occurs in the
Atlantic Forest in south-eastern and north-eastern
Brazil, whereas P. testacea occurs mainly in the
eastern Amazon.
Variation: A few specimens have faint, almost incon-
spicuous, yellow stripes on mesoscutum and frons.
Most female specimens do not have lateral portions of
marginal hair band on T4. It is possible, however,
that they represent worn out specimens, because in
some specimens the marginal hair band on T4 is
complete.
Male. Body length: 6.5–8.0; maximum head width:
3.1; fore wing length (including tegula): 7.1–8.1.
Female. Body length: 6.0–8.1; maximum head width:
2.7; fore wing length (including tegula): 7.5–8.0.
Distribution: Brazil: Bahia, Espirito Santo, Minas
Gerais, Paraíba, Pernambuco, São Paulo, Rio de
Janeiro (Fig. 68).
Type material: Tetrapedia bicolor Smith; holotype
female (BMNH): ‘Type’ ‘B.M. Type\ Hym\ 17B.886’
© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
381
‘Tetrapedia\ bicolor Sm.’ ‘Type F. Sm. Coll.\ 79.22’
‘BRAZIL’.
Tetrapedia nasuta Smith; holotype male (BMNH);
‘Type\ H.T.’, B.M. Type\ Hym\ 17B.896’ Tetrapedia\
nasuta Sm.’ ‘Type F Sm. Coll\ 79-22’ ‘Brazil’.
Additional material: See Appendix.
PARATETRAPEDIA BIFRONS SP. NOV.
(FIGS 45, 71, 92, 123, 159, 160)
Comments and diagnosis: Paratetrapedia bifrons is
easily identified by its conspicuously biconvex frons,
whose mid line is strongly sulcated (Fig. 45). Paratet-
rapedia bifrons has some characters of the bicolor
group: lamella of pronotal collar thinly acute; male
S2–S3 with row of continuous setae adjacent to
margin; male T6 with margins entirely glabrous;
male hind basitarsus without tooth on anterior
margin; male S6 without conspicuous rows of stout
plumose hairs on margin laterally. It differs from the
remaining species in the group mainly by the bicon-
vex frons (Fig. 45), mesoscutum completely black,
without yellow stripes, and female metapostnotum
with fine punctures intermingled amongst sparse
minute punctures (> 2 pd).
Distribution: Costa Rica; Mexico (Fig. 71).
382 A. J. C. AGUIAR and G. A. R. MELO
Figure 71. Geographical distribution of Paratetrapedia bifrons sp. nov., Paratetrapedia flavescens sp. nov., and
Paratetrapedia mexicana sp. nov.
Description
Holotype male: Body length: 9.1; maximum head
width: 2.8; fore wing length (including tegula): 8.6.
Colour. Integument of head, mesosoma and legs
mostly black; terga and sterna pale brown. Mandible
yellow with apex black; labrum, clypeus and supra-
clypeal area yellow; paraocular area with wide yellow
stripe; disc of frons with one thin yellow stripe along
mid line; scape yellow, pedicel and flagellomeres
reddish brown (Fig. 45); wide yellow stripe occupying
about the lower three quarters of genal area (about
1.2 ¥ F2 diameter). Mediotarsus and distitarsus
orange yellow; tibial spurs white. Wing membrane
yellow infumate with orange yellow microtrichia,
veins and pterostigma orange yellow. Pubescence.
Mostly pale yellow, except tibia and basitarsus with
pale brown pubescence. Face mostly glabrous, with
sparse short plumose hairs. Scape with simple yellow
hairs on inner surface, about 0.2 mm in length.
Mesoscutum and scutellum with dense short plumose
pale yellow pubescence; metapostnotum with short
sparse plumose hairs, about 0.07 mm in length; T4
with marginal hair band occupying almost whole
margin; T5 with marginal hair band throughout its
margin; T6 with margin completely glabrous; S2–S3
with continuous row of stout erect simple setae adja-
cent to margin; S6 with simple plumose hairs sparse
on margin laterally and apex. Integument sculpture.
Clypeus and supraclypeal area mostly smooth, with
sparse fine punctures (> 2 pd); frons with sparse
© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
coarse punctures (> 2 pd). Mesoscutum with dense
fine punctures (< 1 pd), intermingled amongst sparse
coarse punctures (> 2 pd); metapostnotum with fine
punctures, mostly sparse (1–3 pd); lateral margins of
metapostnotum with smooth areas; mesepisternum
with sparse fine punctures (3 pd), with some coarser
punctures on upper portion (1 pd). Structure. Epis-
tomal suture convex, above upper margin of clypeus;
frons biconvex with mid line strongly sulcate. Lamella
of pronotal collar acute throughout its length. Scutel-
lum disc strongly convex. Head about 1.2¥ broader
than long (2.8 : 2.3); ratio of lower to upper interocu-
lar distance 0.78 (1.1 : 1.4); clypeus about 1.7¥
broader than long (1.15 : 0.67); scape: length 0.7,
maximum width 0.2; length of F1–3: 0.22, 0.17, 0.25;
diameter of F2: 0.22.
Paratype female: Body length: 8.5; maximum head
width: 3.0; fore wing length (including tegula): 8.5.
Colour. Similar to male, except that mandible,
labrum, and clypeus completely dark brown; disc of
frons with one thin short yellow stripe along mid line.
Pubescence. Similar to male, mostly pale yellow and
legs with brown pubescence; mesoscutum with dense
plumose pale yellow pubescence, intermingled
amongst sparse simple erect setae (about two times
longer than short plumose hairs); scutellum with
pubescence similar to mesoscutum except for simple
erect long setae (about 0.3 mm in length); metapost-
notum with sparse short plumose hairs (about
 SYSTEMATICS OF THE GENUS PARATETRAPEDIA
0.1 mm in length). T4–T6 with marginal hair band
present throughout margin. Integument sculpture.
Clypeus with dense coarse punctures on lateral por-
tions and sparse punctures on disc (0.5–2 pd), inter-
mingled amongst sparse minute punctures (2 pd); disc
of frons with sparse coarse punctures (> 2 pd); anten-
nal scrobe with dense fine punctures (1 pd). Mesoscu-
tum and scutellum with dense minute punctures
(0.5 pd), intermingled amongst sparse coarser punc-
tures (2 pd); metapostnotum with sparse minute
punctures (2 pd), intermingled amongst sparse coarse
punctures on disc (3 pd); lateral margins of metapost-
notum and propodeum with small smooth areas;
mesepisternum with coarse punctures, mostly sparse
(2 pd). Structure. Similar to male. Lamella of pronotal
collar acute throughout its length. Scutellum disc
convex, weakly concave on mid line. Head about 1.3¥
broader than long (3.0 : 2.3); ratio of lower to upper
interocular distance: 0.8 (1.3 : 1.6); clypeus about 1.9¥
broader than long (1.25 : 0.65); scape: length 0.8,
maximum width 0.2; length of F1–F3: 0.24, 0.1, 0.18;
diameter of F2: 0.2.
Variation: The colour of the terga varies from black to
pale brown. One of the specimens has the marginal
hair band occupying about one-quarter of the margin
laterally.
Etymology: The species name refers to its biconvex
frons.
Type material: Holotype male (SEMC), ‘Mexico: San\
Luis Potosi, Xilitla\ 400 m, 7 July 1990\ R.L. Minck-
ley\ ex., on Malvaceae’; paratypes: COSTA RICA: 2
females (BLCU), ‘Costa Rica Alaj.\ Bijagua, 20 km S\
Upala, 10-29 May\ 1991 FD Parker’ ‘USDA-ARS Bee-
Biol.\ & Syst. Lab. Logan\ Utah, FaunalSurvey\ no.
000 025 903’, 1 female (BLCU), idem except ‘000 008
184’; MEXICO: 1 male (MNRJ), ‘Mexico\ Jacubaya\
1900\ Barret’ ‘Tetrapedia\ calcarata\ 1900 Friese
det. Cr.’ ‘N.o 15/850\ Proc.’; 1 male (BLCU), ‘Mexico:
State of\ Vera Cruz, Fortin\ de las Flores-\ Sumi-
dero’ ‘Planta de la\ cerveceria, Ing.\ Daniel Rábago
Res.\ Elev. 2500–3000 ft.’ ‘H.V. Weens Jr.\ Coll.
17-V-65’ ‘NativeBeeSurvey\ USDA, Logan,Utah\
BBSL517253’ ‘Paratetrapedia\ (Paratetrapedia)\ 씹
sp.8\ T Griswold det. 90’; 1 male, ‘DZUP\ 026620’
‘Mex.’ ‘Lophopedia\ abdominalis 씹\ (Cresson,
1878)\ Det. J.S. Moure, 1992’.
PARATETRAPEDIA CALCARATA (CRESSON, 1878)
(FIGS 46, 68, 97, 125, 165, 166)
Tetrapedia calcarata Cresson, 1878: 136; lectotype
female, designated by Cresson (1916): 113, Mexico
(ANSP; examined).
© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
383
Tetrapedia antennata Friese, 1899: 297; lectotype
male, designated by Aguiar (2007), Mexico: Ver-
acruz, Orizaba (ZMB, examined).
Tetrapedia calcarata; Cresson, 1879: 229. Dalla Torre,
1896: 299. Cockerell, 1899: 16. Friese, 1899: 296.
Cockerell, 1906: 98. Ducke, 1910b: 369. Cockerell,
1912b: 31. Cockerell, 1919: 211. Schwarz, 1934: 13.
Tetrapedia calcarata Friese [sic]; Lutz & Cockerell,
1920: 562.
Tetrapedia antennata; Cockerell, 1906: 98. Ducke,
1910b: 369. Friese, 1916: 334. Lutz & Cockerell,
1920: 568. Friese, 1921: 79.
Chalepogenus calcarata; Cockerell, 1914: 320.
Chalepogenus calcaratus; Cockerell, 1917: 302. Lutz
& Cockerell, 1920: 562.
Chalepogenoides calcaratus; Michener, 1942: 281.
Paratetrapedia calcarata; Michener, 1954: 114,
figs 65–67 (drawings of S7–S8 and genital capsule
of male). Vogel, 1974: 247 [floral record: Ornitho-
cephalus sp. (Orchidaceae)]. Cane, 1979: 128,
fig. 128 (drawing of tibial-femur articulation of hind
leg). Heithaus, 1979: 195. Buchmann & Buchmann,
1981: 17, fig. 9 [morphology of oil collector appara-
tus of fore basitarsus; floral record: Mouriri myrtili-
oides (Melastomataceae)]. Cocucci, Sersic & Roig-
Alsina, 2000: 69. Michener, 2000: 667, figs 106–2.
Smith-Pardo, 2003: 338. Aguiar, 2007: 621. Davies,
2009: 175.
Paratetrapedia calcarata Cresson [sic]; Steiner, 1985:
1540 (floral record: Spachea membranacea (Mal-
pighiaceae).
Paratetrapedia (Paratetrapedia) calcarata; Michener
& Moure, 1957: 415. Ayala et al., 1996: 461.
Comments and diagnosis: Paratetrapedia calcarata is
very similar to P. bicolor, differing only by omaular
angle slightly acute on upper third with adjacent
surface of mesepisternum weakly concave, mesoscu-
tum disc with thin yellow stripes; axilla yellow
dorsally; female with terga mostly smooth (few speci-
mens with some thin reticulate microsculpture on
disc of T3–T4); gena of male with yellow stripe along
most of eye margin; paraocular area of male with
yellow stripe occupying more than the lower half.
Variation: The colour of terga varies between dark
brown to brownish yellow. Some specimens lack
yellow marks on the frons. Lamella of pronotal collar
in some specimens is slightly low on lateral portions,
whereas in others it is higher and diverges from the
margins of mesoscutum, but in all specimens it is
very thinly lamellate and acute. Some females have
microsculpture of thin reticulate stripes on disc of T3.
The vertical surface of T1 varies from completely
smooth to sculptured with dense minute punctures.
One specimen from Costa Rica has the scutellum and
384 A. J. C. AGUIAR and G. A. R. MELO
metanotum completely yellow. Most of specimens
have wing membrane brown infumate but the
specimens from Venezuela have it orange yellow
infumate.
Male. Body length: 7.0–7.8; maximum head width:
2.25–2.35; fore wing length (including tegula): 6.7–
7.8. Female. Body length: 6.8–7.6; maximum head
width: 2.6–2.9; fore wing length (including tegula):
7.2–7.9.
Distribution: Colombia; Costa Rica; Ecuador; Guate-
mala; Honduras; Mexico; Nicaragua; Panama; Ven-
ezuela (Fig. 68).
Type material: Tetrapedia calcarata Cresson; lecto-
type female (ANSP) designated by Cresson (1916):
‘Mex.’ ‘Lectotype\ 2380’; paralectotype male (ANSP):
‘Mex’ ‘Allotype\ 2380’ ‘Calcarata\ Cress.’; paralecto-
type male: ‘Mex’ ‘Allotype\ 2380’ ‘Calcarata\ Cress.’.
Tetrapedia antennata Friese; lectotype male (ZMB):
‘Mexico\ Ehrenbg. S.’ ‘11700’ ‘Tetrapedia 씹\ anten-
nata\ det. Friese 1898\ n.sp.’ ‘co.\ Type’.
Additional material: See Appendix.
PARATETRAPEDIA TESTACEA (SMITH, 1854)
(FIGS 9, 21, 25, 37, 68, 96, 126, 167, 168)
Tetrapedia testacea Smith, 1854: 366; holotype
female, Brazil: Pará (BMNH, examined).
Tetrapedia obtusa Vachal, 1909: 27; lectotype female
designated by Aguiar (2007), French Guiana:
Kourou (MNHP, examined).
Tetrapedia testacea; Dalla Torre, 1896: 300. Friese,
1899: 297. Ducke, 1901: 55 [floral record: Stachy-
tarpheta sp. (Verbenacaea)]. Ducke, 1902: 323
[floral record: Psychotria sp. (Rubiaceae), Pavonia
typhalaea (Malvaceae)]. Friese, 1906: 91 (misiden-
tification, suggested by geographical distribution).
Ducke, 1908: 84. Ducke, 1910a: 364. Ducke, 1910b:
98. Friese, 1923: 3. Popov, 1939: 164, figs D–G
(drawings of S7–S8 and genital capsule of male).
Tetrapaedia [sic] testacea; Schrottky, 1902: 554.
Tetrapaedio [sic] testacea; Schrottky, 1902: 554
Tetrapedia obtusa; Ducke, 1910b: 98. Cockerell, 1931:
413.
Chalepogenus testaceus; Cockerell, 1923a: 449.
Chalepogenoides testaceus; Michener, 1942: 281.
Paratetrapedia (Paratetrapedia) testacea; Michener &
Moure, 1957: 416. Albuquerque & Rego, 1989: 168;
Rêgo & Albuquerque, 1989: 185 [floral record: Byr-
sonima crassifolia (Malpighiaceae)]. Silveira et al.,
2002: 136.
Paratetrapedia testacea; Moure, 1941: 518. Vogel,
1974: 219 [floral record: Ornithocephalus sp.
© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
(Orchidaceae)]. Shanks, 1986: 2. Cocucci et al.,
2000: 69. Rebêlo, Rêgo & Albuquerque, 2003: 273.
Aguiar, 2007: 625. Davies, 2009: 175.
Comments and diagnosis: Paratetrapedia testacea is
very similar to P. bicolor, being distinct mainly by
orange yellow integument and allopatric distribution.
The female of P. testacea has T1 with dense minute
punctures on vertical surface and microsculpture of
fine reticulate lines on terga. Males have the scape
conspicuously dilated on lateral surface, margins of
T5–T6 with margins completely glabrous, and S6
with setae only on apex, without rows of plumose
hairs on lateral margins.
Variation: Some Brazilian specimens from Maranhão,
Goiás, and Amazonas lack the tergal microsculpture
of fine reticulated lines. Some specimens have micros-
culpture only on T4 or only on disc of T3. A few
specimens have disc of mesoscutum and central
portion of frons partially pale brown.
Male. Body length: 6.5–8.0; maximum head width:
2.2–2.5; fore wing length (including tegula): 7.1–8.1.
Female. Body length: 6.0–8.1; maximum head width:
2.3–2.6; fore wing length (including tegula): 7.5–8.0.
Distribution: Brazil: Acre, Amapá, Amazonas, Goiás,
Maranhão, Pará, Roraima, Tocantins; Guyana;
French Guiana; Peru; Suriname (Fig. 68).
Type material: Tetrapedia testacea Smith; holotype
female (BMNH): ‘Type\ H.T.’ ‘B.M. Type\ Hym.\
17.B.895’ ‘Tetrapedia\ testacea Sm.’ ‘81’ ‘Type F.Sm.
Coll.\ 79.22’.
Tetrapedia obtusa Vachal, lectotype female
(MNHN): ‘Juin’ ‘Guyane Française\ Roches de
Kourou\ Coll. Le Moult’ ‘Tetrap. 씸\ obtusa\ Vach.’
‘Museum Paris\ Coll. J. Vachal 1911’ ‘Holotype’.
Additional material: See Appendix.
PARATETRAPEDIA VOGELI SP. NOV.
(FIGS 68, 81, 94, 127, 199, 200)
Comments and diagnosis: Paratetrapedia vogeli is
very similar to species with brown integument of the
bicolor group. Males of P. vogeli are distinct from P.
bicolor and P. calcarata by S6 with two conspicuous
rows of hairs on lateral margins (Fig. 81). Paratetra-
pedia vogeli differs from P. basilaris by antennal
scape of male dilated laterally on distal portion and
terga with marginal zone completely brown. Females
of P. vogeli can be confused with those of P. andina
because of their sympatric distribution, and similar
 SYSTEMATICS OF THE GENUS PARATETRAPEDIA
colour and morphology. Paratetrapedia vogeli is dis-
tinguished by pronotal collar being acutely lamellate
throughout its length.
Description
Holotype male: Body length: 7.2; maximum head
width: 2.4; fore wing length (including tegula): 7.2.
Colour. Integument mostly dark brown; mandible
with basal half pale yellow and apical half black;
labrum pale yellow; clypeus with two yellow spots on
lateral margins; paraocular area with yellow stripe
occupying lower half; frons with thin yellow stripe
along the mid line (c. 0.2 ¥ F2 diameter); scape
weakly yellow laterally; mesoscutum black with two
faint yellow stripes on disc; T2 and S2 with disc pale
yellow, almost white. Fore wing membrane brown
infumate, with dark brown microtrichia; veins and
pterostigma orange yellow. Pubescence. Paraocular
area, frons, vertex, genal area, mesepisternum,
metepisternum, propodeum, and sternal margins
mostly with whitish pubescence; mesoscutum, scutel-
lum, tergal margins, and legs with pubescence mostly
brown. Scape with long setae on inner face, c.
0.08 mm in length; mesoscutum and scutellum with
short plumose pale brown pubescence; metapostno-
tum with short plumose setae, c. 0.2 mm in length.
Margins of T4–T6 mostly glabrous; margin of S2–S3
with continuous rows of simple setae; S4 and S5
typical of genus; S6 with two rows of plumose hairs on
margin laterally, and plumose hairs on margin of
apex. Integument sculpture. Clypeus with coarse
shallow punctures (1 pd), intermingled amongst
sparse minute punctures (2 pd); supraclypeal area
with coarse punctures (0.5–2 pd); frons with dense
coarse punctures on disc (0.5 pd), and finer dense
punctures on lateral portions and antennal scrobe
(< 0.5 pd); antennal scrobe with small smooth area.
Mesoscutum with dense fine punctures (< 0.5 pd)
intermingled amongst sparse coarse punctures
(> 2 pd); metapostnotum with dense fine punctures
(< 0.5 pd), with finer punctures on lateral margins;
mesepisternum mostly smooth, with sparse fine punc-
tures (> 2 pd). Vertical surface of T1 with dense
minute punctures (1 pd). Structure. Scape with apical
portion dilated laterally; pronotal collar with lamella
acute throughout its length; scutellum biconvex.
Head about 1.2¥ broader than long (2.4 : 2.0); ratio
between lower interocular distance to upper interocu-
lar distance: 0.84 (1.1 : 1.33); clypeus about 2.1¥
broader than long (1.07 : 0.57); scape: length 0.65,
maximum width 0.2; length of F1–F3: 0.2, 0.12, 0.22;
diameter of F2: 0.2.
Paratype female: Body length: 7.6; fore wing length
(including tegula): 7.1; maximum head width: 2.6.
Colour. Similar to male, except for the yellow marks
© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
385
on face faint; mandible with basal half pale yellow;
labrum completely brown; clypeus with two small
yellow faint spots on lateral portions; frons with
yellow stripe on mid line (c. 0.2 ¥ F2 diameter);
mesoscutum with two faint yellow stripes on disc;
axilla pale yellow dorsally. Disc of T2 pale yellow,
almost white; S2–S3 pale yellow, almost white. Pubes-
cence. Similar to male; scutellum with simple sparse
erect setae, c. 0.3 mm in length. Marginal bands on
terga similar to male, except for marginal bands
complete on T5–T6. Sterna with row of setae on
margin, weakly oval sinuated on mid portion. Integu-
ment sculpture. Clypeus with dense coarse punctures
(< 0.5 pd); frons with dense coarse punctures on disc
(< 0.5 pd), and dense minute punctures on lateral
portions (< 0.5 pd). Vertical surface of T1 with dense
minute punctures (0.5 pd). Structure. Similar to male;
scutellum with profile convex, mid line slightly
concave. Head c. 1.2¥ broader than long (2.6 : 2.1);
ratio of lower interocular distance to upper interocu-
lar distance: 0.85 (1.2 : 1.4); clypeus c. 1.95¥ broader
than long (1.17 : 0.6); scape: length 0.65, maximum
width 0.17; length of F1–F3: 0.22; 0.12; 0.17; diameter
of F2: 0.2.
Variation: The yellow marks on clypeus, paraocular
area, frons, mesoscutum, and scutellum can be faint
or completely absent.
Etymology: The species name is dedicated to Dr
Stefan Vogel.
Distribution: Brazil: Acre, Mato Grosso; Peru
(Fig. 68).
Type material: Holotype male, ‘DZUP\ 022025’
‘Quincemil-Cuzco\ PERU 20 X-61\ LUIS E PENA’.
Paratypes: BRAZIL: Acre: 1 male (RPSP), ‘Brasil-
Acre\ RBr-UFAC/PZ.\ Data 29/11/1994’ ‘A.H.
Machado\ E.M. Santos’ ‘No 320’, 4 males and 1
female, idem except ‘No 317’, ‘Data 04/05/1995’ ‘904’,
‘Data 28/02/1995’ ‘No 495’, ‘Data 08/06/1995’ ‘1045’,
‘Data 08/06/1995’ ‘1025’; 1 male, ‘DZUP\ 022009’
‘CRUZEIRO DO SUL\ ACRE-Brasil II-63\ M. Alva-
renga’; 1 female (INPA), ‘BRASIL-ACRE\ R. Branco-
UFAC\ 29\04\1993\ M.L. Oliveira’; Mato Grosso: 1
male, ‘DZUP\ 022689’ ‘Chapada dos\ Guimarães,
MT\ 8-1-1987\ C. Elias leg.’; PERU: 1 female
(CLAUS),’PERU, JU, Valle Chan-\ chamayo, Picha-
naki\ Puente Ipoki, 658 masl\ S11 00,840 W74
44,747\ 27x01 C. Rasmussen leg.’; 2 males (CLAUS),
‘PERU, M, Tambopata\ Jungle Lodge, 225 m.a.s.l.\
S12 49,456 W69 24,163\ 9-20x01, C Rasmussen leg.’;
1 male (AMNH), ‘Peru, Quincemil,\ on branch R.
Manu\ Madre de Dios Prov.\ X-16-31-1962’; 1 female,
‘DZUP\ 022602’ ‘Quincemil-Cuzco\ PERU 9 XI 62\
386 A. J. C. AGUIAR and G. A. R. MELO
LUIS E PENA’, 2 males and 1 female (DZUP), idem
except ‘022654’, ‘20 X 61’ ‘022212’, ‘20 X 61’ ‘022011’.
PARATETRAPEDIA BASILARIS SP. NOV.
(FIGS 63, 68, 93, 128, 163, 164)
Comments and diagnosis: Paratetrapedia basilaris
can be recognized by the translucent hyaline margins
of its terga. The male is easily distinguished from the
other species of the bicolor group by its unmodified
scape, not dilated apically. Additionally, as in P.
vogeli, the male S6 has two rows of plumose setae on
the lateral margins. Paratetrapedia basilaris differs
from P. vogeli by the almost right angled omaulus.
Distribution: Brazil: Amapá, Amazonas, Pará;
Guyana; French Guiana (Fig. 68).
Description
Holotype male: Body length: 7.0; maximum head
width: 2.5; fore wing length (including tegula): 7.0.
Colour. Integument mostly dark brown. Mandible
with basal half pale yellow and apical half black;
labrum pale yellow, almost white; clypeus with two
yellow stripes on lateral margins, and a thin yellow
stripe on lower margin; paraocular stripe very thin (c.
0.8 ¥ F2 diameter), occupying more than half of lower
paraocular area; supraclypeal area with yellow spot
on disc; frons with thin yellow stripe on mid line;
genal area with thin and short yellow stripe on lower
portion of ocular margin; scape pale brown, weakly
yellow on outer lateral face; pedicel and antennal
flagellomeres pale brown. Mesoscutum with two thin
pale yellow stripes on disc and lateral margins; disc of
scutellum with two yellow spots on lateral portions;
axilla pale yellow dorsally. Terga and sterna pale
brown; margins of sterna translucent hyaline, with
silvery reflections. Wing membrane brown infumate;
veins and pterostigma orange yellow; microtrichia
dark brown. Tibial spurs white. Pubescence. Pubes-
cence mostly white on paraocular area, mesepister-
num and propodeum; remaining body with brownish
pubescence. Scape with brown setae on inner face, c.
0.12 mm in length. Mesoscutum and scutellum with
very short plumose pubescence, velvet-like. Terga
with margins mostly glabrous; T4 mostly glabrous,
with few plumose hairs on margin laterally; T5–T6
with margins glabrous; margin of S2 with row of
continuous stout simple brown setae; margin of S3
with row of continuous stout simple pale yellow setae;
S4 and S5 with pubescence typical of the genus; S6
with row of stout plumose setae on margins laterally,
and short plumose hairs on the margin of apex.
Integument sculpture. Clypeus and supraclypeal area
with dense coarse punctures (< 1 pd), intermingled
amongst sparse minute punctures (> 3 pd); disc of
© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
frons with sparse coarse punctures (3 pd); antennal
scrobe with dense fine punctures (1 pd), with one
small smooth area. Mesoscutum and scutellum with
fine dense punctures (0.5 pd), intermingled amongst
coarse and sparser punctures (2 pd); metapostnotum
with sparse fine punctures (2–3 pd); mesepisternum
with sparse coarse punctures (3 pd), with smooth
shiny integument between the punctures. Structure.
Lamella of pronotal collar acute throughout its
length. Disc of scutellum convex. Upper third of
omaulus in acute angle, almost carinate. Head about
1.2¥ broader than long (2.5 : 2.0); ratio of lower
interocular distance to upper interocular distance:
0.76 (1.0 : 1.3); clypeus c. 1.5¥ broader than long
(1.0 : 0.65); scape: length 0.8, maximum width 0.19;
length of F1–3: 0.17, 0.12, 0.19; diameter of F2: 0.17.
Paratype female: Body length: 7.1; maximum head
width: 2.6; fore wing length (including tegula): 7.2.
Colour. Similar to male, except for yellow stripe on
paraocular area slightly faint; genal area completely
dark brown. Pubescence. Similar to male, except
T4–T6 with marginal hair band of brown hairs
throughout its margin. Integument sculpture. Similar
to male, except that punctures denser and finer;
clypeus and supraclypeal area with dense coarse
punctures (< 1 pd); disc of frons with dense coarse
punctures (< 0.5–2 pd), and dense fine on lateral por-
tions (< 1 pd). Structure. Similar to male; scutellum
weakly convex, almost flat dorsally. Head about 1.2¥
broader than long (2.5 : 2.0); ratio of lower interocular
distance to upper interocular distance: 0.78 (1.1 : 1.4);
clypeus about 1.7¥ broader than long (1.1 : 0.65);
scape: length 0.7, maximum width 0.19; length of
F1–F3: 0.16, 0.08, 0.10; diameter of F2: 0.16.
Variation: Two specimens from Manaus (Brazil, Ama-
zonas) have very faint yellow marks on paraocular
area and frons. The colour of these specimens,
however, has probably been altered by deterioration.
Most examined female specimens have T4 with
margins mostly glabrous, with few setae on lateral
third, but other apparently better preserved speci-
mens have a complete marginal band.
Etymology: The species name refers to the plesiomor-
phic characters of this species related to the other
species of the bicolour group.
Type material: Holotype male, ‘DZUP\ 022110’
‘MANAUS-AM\ Brasil-VII 62\ F.M. OLIVEIRA’.
Paratypes: BRAZIL: Amapá: 1 female, ‘DZUP\
022789’ ‘OIAPOQUE Amapá\ Brasil-V-1959\ M.
Alvarenga’, 2 females (DZUP), idem except ‘022985’,
‘022986’; Amazonas: 1 male, ‘DZUP\ 022107’
‘MANAUS-AM\ Brasil-VII 62\ F.M. OLIVEIRA’, 39
 SYSTEMATICS OF THE GENUS PARATETRAPEDIA
males (DZUP), idem except ‘022110’, ‘022111’,
‘022704’, ‘022705’, ‘022712’ to ‘022715’, ‘022717’ to
‘022721’, ‘022723’ to ‘022734’, ‘022736’ to ‘022740’,
‘022742’, ‘022743’, ‘022786’, ‘022788’; ‘022790’,
‘022791’, ‘022987’, ‘022991’, ‘023542’; 1 female
(DZUP), ‘DZUP\ 022792’ ‘MANAUS-AM\ Brasil-VII
62\ F.M. OLIVEIRA’, 2 females (DZUP), idem, except
‘022793’, ‘022990’; 1 male, ‘DZUP\ 22698’ ‘MANAUS-
AM\ Brasil-I-61\ C.ELIAS leg’, 1 male and 1 female
(DZUP), idem except ‘022983’, ‘022988’; 1 female,
‘DZUP\ 22984’ ‘MANAUS-AM\ Brasil-VII 58\
C.ELIAS leg’, 1 female (DZUP), idem except ‘022989’;
4 females (INPA), ‘Sauv. erect.\ Ducke\ ‘buzz.’\
12.5.91’; 1 female (INPA), ‘Sauv. erect.\ Res. Ducke\
‘buzz.’\ 12.5.91’; 1 female (INPA), ‘91/18\ Sauv.
erect.\ buzzen\ 16.6.91’; 1 female (INPA), ‘BRASIL:
Amazonas\ 26 km NE Manaus\ Reserva Ducke\
VIII- 1981’ ‘J.A.Rafael\ Ar: Malaise’; 1 female (INPA),
‘AM-Manaus\ 5-V-81\ Reviver 69’; 1 female (INPA),
‘Brasil-AM\ Manaus\ PDBFF\ 28 à 30-V-2003’ ‘M.L.
Oliveira &\ J.A. Cunha leg.\ Cap. P. Alegre\ Em
Melastomat.’; 1 male (NHRS), ‘Maná-\ os’ ‘Amazon\
Roman’; 1 female (NHRS), ‘Maná\ -os’ ‘T. albipennis\
1920 Friese det. Fr.’ ‘Tetrapedia F\ sp.?’; 1 male
(NHRS), idem except ‘M’; Pará: 1 male (MZSP), ‘Alter
do Chão\ Pará-Brasil\ 19.IX.1969\ Exp. Perm.
Amaz.’; FRENCH GUIANA: 1 female (SEMC),
‘FRENCH GUIANA\ Kourou, km. 16 SW.\ 8 April
1977\ D. Roubik, no. 125’, 1 female (SEMC), idem
except ‘Paratetrapedia\ det. R.W. Brooks 1988’; 1
female (SEMC), ‘FRENCH GUIANA\ Kourou, km. 17
SW.\ 22 March 1977\ D. Roubik, no. 115’; 1 female
(SEMC), ‘FRENCH GUIANA\ Matoury, 81 km S
and\ 5 km W on Belizon Rd., 10 m\ 4o21′35′N,
52o19′41′W\ 27 MAY 1997; J. Ashe, R. Brooks\
FG1AB97 047’; GUYANA: 1 female (AMNH), ‘British
Guiana\ Kartabo, Bartica\ Dist. 1920’ ‘Trop.
Research Station\ New York Zool. Society\ No.
20860’ ‘Gift of New York\ Zoo.Soc. Dept.\ Tropical
Research\ Willian Beebe, Dir.’; 1 female (SEMC),
‘Kartabo\ British Guiana\ 25.VII.1924’ ‘Collected by\
Jay F. W. Pearson’ ‘Gift of New York\ Zoo.Soc. Dept.\
Tropical Research\ Willian Beebe, Dir.’.
THE LINEATA SPECIES GROUP
Species of the lineata group are very similar to
species of the flavipennis group, with male S3 deeply
concave and ‘U’-shaped, covered by dense short
plumose pubescence (Figs 17, 27, 28; character 47:3;
except for P. punctata, P. flavipennis, P. duckei, P.
romani). Species of the lineata group area distinct
from those of the flavipennis group mainly by the
male hind basitarsus with tooth on anterior margin
(Figs 32, 33; character 30:1; P. flavifrons and P. punc-
tata have a short carina in the corresponding position
© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
387
of the tooth); lamella of pronotal collar acute through-
out its length (except P. tocantinensis, which is
slightly obtuse); row of setae on margin of male S2
interrupted by two gaps, isolating a short row of setae
in the mid portion (characters 45:1, 46:1); female
mandible without any indication of a vestigial second
preapical tooth; row of setae on female S2–S3 with
weak sinuosity, ‘U’-shaped on mid portion (Fig. 30);
mandible, lower portion of gena, coxa, trochanteres,
and mesepisternum with long fine hairs ventrally,
hairs longer than width of mandibular base (charac-
ter 36:1; except P. fervida and P. punctata); premen-
tum of female with numerous stout curved setae;
epistomal suture on upper margin of clypeus con-
spicuously convex; supraclypeal area with central
portion mostly smooth and conspicuously convex
(except P. punctata). Most of species of the lineata
group have a bicoloured integument and are found in
the Amazonian basin. The only species with dark
brown integument in the group are P. fervida and P.
punctata, although some specimens from lowland
areas have paler integument and faint pale yellow
lines on mesoscutum. This pattern of species with
dark integuments on highlands, mainly the Atlantic
Forest, and species with pale integuments on lowland
areas of the Amazon basin is also observed in other
groups of bees (e.g. Partamona; Pedro & Camargo,
2003).
PARATETRAPEDIA LINEATA (SPINOLA, 1853)
(FIGS 10, 19, 22, 27, 30, 49, 72, 80, 104, 137, 184)
Ancyloscelis lineata Spinola, 1853: 87; holotype
female, Brazil: Pará (MRSN, examined).
Tetrapedia lacteipennis Vachal, 1909: 26; lectotype
female designated by Aguiar (2007), Peru:
Huanúco, Paquitea (MNHP, examined).
Ancyloscelis lineatus; Smith, 1854: 367.
Eucera lineata; Dalla Torre, 1896: 236.
Macrocera lineata; Schrottky, 1902: 522.
Tetrapedia lineata; Ducke, 1910a: 368; Ducke, 1910b:
98.
Paratetrapedia lineata; Moure, 1941: 517.
Gottsberger, 1986: 33 (misidentification, based on
geographical distribution). Santos et al., 2004: 323.
Melo & Zanella, 2003: 2919 (misidentification of
P. flaveola).
Tetrapedia lacteipennis; Moure, 1944a: 74.
Paratetrapedia (Paratetrapedia) lineata; Michener &
Moure, 1957: 415, figs 19–21 (drawings of S7–8 and
genital capsule of male). Silveira & Campos, 1995:
375; Pedro, 1996: 251 (misidentification, based on
geographical distribution). Pedro & Camargo, 1999:
202. Silveira et al., 2002: 136.
Ancyloscelis lineata; Casolari & Moreno, 1980:
143.
388 A. J. C. AGUIAR and G. A. R. MELO
Paratetrapedia aff. lineata; Smith-Pardo, 2003: 338.
Paratetrapedia lineata; Aguiar, 2007: 620.
Comments and diagnosis: Paratetrapedia lineata has
been frequently confused with other species of
Paratetrapedia and related genera (e.g. Tropidopedia
flavolineata Aguiar & Melo, 2007a) exhibiting a
similar pattern of bicoloured integument. This colour
pattern, black to reddish brown head and thorax,
with many yellow marks, combined with a mostly
yellow and elongated metasoma, is found in many
species of the lineata and flavipennis groups and
some stingless bee genera (subtribe Meliponina;
Camargoia, Ptilotrigona, Tetragona) in the Amazon
Forest.
Paratetrapedia lineata can be distinguished mainly
by the pale hyaline wing membrane with pale yellow
microtrichia, and a few dark microtrichia on the
margin of the forewing. Only two other species, P.
flaveola and P. tocantinensis, have a bicoloured
integument and pale hyaline wing membrane as in P.
lineata. Paratetrapedia lineata differs from these two
species by the metapostnotum dull with fine reticu-
late sulcus between the punctures (Fig. 22), pronotal
collar with an acute lamella throughout its length;
genal area completely yellow or with at least a wide
yellow stripe (c. two times F2 diameter); supraclypeal
area with surface strongly convex and mostly smooth;
disc of supraclypeal area with sparse punctures (c.
2 pd).
The holotype of P. lineata has the metasoma glued
to the mesosoma and the glue hides the vertical
surface of T1 and propodeum. Ducke (1910b) sug-
gested that Tetrapedia elongata Friese, 1899 could be
a synonymy of P. lineata (Spinola). Although the holo-
type of P. elongata has not been located, P. lineata is
not known from south-eastern Brazil (the type local-
ity of P. elongata is Santa Leopoldina, in Espírito
Santo). In the ZMB collection, there are 15 specimens
identified by Friese as T. elongata, from far apart
localities (Pará, Bolivia, Maranhão).
Variation: The yellow mark on the disc of metapost-
notum can be very large, covering the metapostnotum
entirely, or it can be very small and restricted to
metanotum margin. Two specimens have the yellow
stripe on paraocular area with the upper portion
acutely sinuous, similar to the condition in P. flaveola.
The colour of genal area can vary between entirely
yellow or reddish brown to a wide yellow stripe (c. two
times F2 diameter). One specimen from Yungas
(Bolivia) possesses the wing membrane weakly pale
yellow infumate, metapostnotum with fine dense
microsculpture and frons with minute subcontiguous
punctures (< 0.5 pd). Some specimens from Peru,
© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
Ecuador, and Óbidos (Brazil, Pará) have T1 with
minute punctures and disc of terga with microsculp-
ture of fine reticulated lines.
Male. Body length: 6.7–8.4; maximum head width:
2.5–2.7; wing length (including tegula): 6.8–7.2.
Female. Body length: 7.1–7.8; maximum head width:
2.3–2.9; fore wing length (including tegula): 7.6–7.8.
Distribution: Bolivia; Brazil: Acre, Amazonas, Mato
Grosso, Pará, Rondônia; Colombia, Ecuador, Peru
(Fig. 72).
Type material: Ancyloscelis lineata Spinola, holotype
female (MRSN), ‘Holotype\ lineata\ Spinola\ JS.
Moure 1958’, information associated to the specimen
in the drawer (Guido Pagliano, pers. comm., 2003):
‘Ancyloscelis lineata mihi, femmina – D. Ghiliani –
Para 1846’. This information agrees with the data
presented by Casolari & Moreno (1980) about Spino-
la’s type material.
Tetrapedia lacteipennis Vachal; lectotype female
(MNHP): ‘Pérou\ Paquitea’ ‘Museum Paris\ Coll. J.
Vachal 1911’.
Additional material: See Appendix.
PARATETRAPEDIA ALSINAI SP. NOV. (FIG. 73)
Comments and diagnosis: Paratetrapedia alsinai is
very similar to P. flavifrons in the yellow infumate
wing membrane and to P. tocantinensis and P. romani
in the long acute tooth on anterior margin of hind
basitarsus of male. The female of P. alsinai differs
from P. romani by T4 with a complete marginal hair
band, occupying the entire margin, and smaller body
size. The male of P. alsinai differs from P. flavifrons by
hind basitarsus with one acute tooth on anterior
margin. Paratetrapedia alsinai is distinct from P.
flavifrons and P. romani by the disc of mesoscutum
without orange yellow lateral portions. The male
genitalia of P. alsinai were not dissected in order to
preserve the only male specimen known, here selected
as the holotype.
Description
Holotype male: Body length: 8.0; maximum head
width: 2.65; fore wing length (including tegula): 8.0.
Colour. Integument mostly orange yellow. Mandible
yellow with apex black; disc of frons, upper paraocu-
lar area, and vertex black; frons with one thin yellow
stripe on mid line, not contiguous with supraclypeal
area and contacting the mid ocellus; genal area com-
pletely yellow; scape yellow, pedicel and flagellomeres
reddish orange. Mesoscutum black with two thin
yellow stripes on disc and lateral margins; mid
portion of lateral yellow stripe of mesoscutum weakly
 SYSTEMATICS OF THE GENUS PARATETRAPEDIA 389

Figure 72. Geographical distribution of Paratetrapedia lineata.

Figure 73. Geographical distribution of Paratetrapedia alsinai sp. nov., Paratetrapedia tocantinensis sp. nov.,
Paratetrapedia romani, and Paratetrapedia flavifrons sp. nov.

enlarged; scutellum yellow; axillar fossa black. Terga
orange yellow; marginal zone slightly brown. Wing
membrane yellow infumate; microtrichia mostly
yellow; apical third of fore wing dark because of
presence of dark brown microtrichia; veins and
pterostigma orange yellow; tibial spurs white. Pubes-
cence. Mostly pale yellow, except hind basitarsus with
pubescence pale brown. Mandible, lower portion of
mesepisternum, coxa, trochanter, and lower mesepis-
ternum with long fine hairs, length about twice the

© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
390 A. J. C. AGUIAR and G. A. R. MELO
width of mandibular base. Scape with long pale
yellow setae on inner face, about 0.18 mm in length.
Mesoscutum and scutellum with sparse short
plumose yellow pubescence; T4–T6 with dense mar-
ginal band of yellow hairs occupying whole margin;
posterior margin of S2 with short row of simple setae
isolated on mid portion; posterior margin of S3 of
male deeply concave in ‘U’ shape covered by dense
short plumose pubescence; S4 and S5 typical of genus;
S6 with two rows of stout plumose setae on lateral
margin and short plumose hairs on margin of apex.
Integument sculpture. Clypeus with sparse shallow
coarse punctures (1–2 pd), intermingled amongst
sparse minute punctures (> 2 pd); disc of frons with
sparse coarse punctures (1–2 pd); antennal scrobe
with dense fine punctures (1 pd). Mesoscutum and
scutellum with dense minute punctures (< 0.5 pd),
intermingled amongst sparse coarser punctures
(> 2 pd); metapostnotum with sparse fine punctures
(> 3 pd); lateral mesepisternum mainly with sparse
coarse punctures (> 2 pd). Structure. Pronotal collar
with lamella thin acute throughout its length. Scutel-
lum profile weakly convex. Hind basitarsus with
acute tooth on anterior margin. Head about 1.2¥
broader than long (2.65 : 2.15); ratio of lower interocu-
lar distance and upper interocular distance: 0.82
(1.17 : 1.42); clypeus about 1.9¥ broader than long
(1.1 : 0.6); scape: length 0.67, maximum width 0.22;
length of F1–F3: 0.2, 0.2, 0.25; diameter of F2: 0.2.
Paratype female: Body length: 8.0; maximum head
width: 2.7; fore wing length (including tegula): 7.8.
Colour. Similar to male. Pubescence. Similar to male,
except for pubescence on margin of mandible, coxa,
trochanter, and mesepisternum with only short hairs;
T4–T6 with marginal band of golden yellow hairs
occupying whole margin. Integument sculpture.
Similar to male, except for not having minute punc-
tures on clypeus; frons with coarser punctures; frons
with dense coarse punctures (1 pd) on disc and fine
punctures on lateral portions (0.5 pd); metapostnotum
with dense minute punctures. Structure. Similar to
male, except that scutellum disc weakly convex,
almost flat. Head about 1.2¥ broader than long (2.7;
2.25); ratio of lower interocular distance to upper
interocular distance: 0.86 (1.3 : 1.5); clypeus about
two times broader than long (1.25 : 0.6); scape: length
0.77, maximum width 0.17; length of F1–F3: 0.22, 0.1,
0.2; F2 diameter: 0.22.
Etymology: The species name is dedicated to Dr
Arturo Roig-Alsina.
Distribution: Brazil: Amazonas (Fig. 73).
© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
Type material: Holotype male, ‘DZUP\ 022973’
‘MANAUS-AM\ BRAZIL VII-52\ F.M. Oliveira’.
Paratypes: 1 female (INPA), ‘BRAZIL: AM\
MANAUS\ C. Univers.\ 11-VI-1982’ ‘J.A. Rafael\
Arm. Malaise’.
PARATETRAPEDIA FERVIDA (SMITH, 1879)
(FIGS 2, 17, 33, 41, 70, 99, 131, 171, 172)
Tetrapedia fervida Smith, 1879; holotype male,
Brazil: Rio de Janeiro, Teresópolis [‘Constancia’]
(BMNH, examined).
Tetrapedia bunchosiae Friese, 1899: 286; lectotype
male, designated by Aguiar (2007), Brazil: Santa
Catarina, Blumenau (ZMB, examined).
Tetrapedia flaviventris Friese, 1899: 292; lectotype
female, Brazil: Santa Catarina, Blumenau (ZMB,
examined), designated by Moure (1996: 927) (rede-
scription of male and female).
Tetrapaedia [sic] obsoleta Schrottky, 1902: 547; holo-
type male, Brazil: São Paulo, Jundiaí (MZSP, exam-
ined).
Tetrapedia monacha Strand, 1910: 518; lectotype
female, designated by Aguiar (2007), Paraguay:
Assuncion (ZMB, examined).
Tetrapedia velutina Friese, 1910: 459; lectotype male,
here designated, Paraguay: La Cordillera, San Ber-
nardino (ZMB, examined).
Tetrapedia fervida; Friese, 1899: 302. Schrottky, 1902:
557.
Tetrapaedia [sic] bunchosiae; Schrottky, 1901: 212.
Schrottky, 1902: 546.
Tetrapaedia [sic] flaviventris; Schrottky, 1902:
551.
Tetrapedia velutina; Friese (in Strand, 1909: 234),
nomen nudum.
Tetrapedia bunchosiae; Ducke, 1910b: 369. Cockerell,
1912b: 31. Schrottky, 1913: 260.
Chalepogenus bunchosiae; Cockerell, 1923a: 450.
Pratetrapedia [sic] bunschosiae [sic]; Moure, 1941:
518.
Paratetrapedia velutina; Moure, 1941: 518. Michener,
1954: 115.
Chalepogenoides bunchosiae; Michener, 1942: 281.
Paratetrapedia (Paratetrapedia) bunchosiae;
Michener & Moure, 1957: 415. Silveira et al., 2002:
136.
Paratetrapedia (Paratetrapedia) obsoleta; Michener &
Moure, 1957: 416. Silveira et al., 2002: 136.
Paratetrapedia bunchosiae; F. Müller in Vogel (1974:
179) [floral record: Bunchosia gaudichaudiana
(Malpighiaceae)]. Cocucci et al., 2000: 69. Silveira
et al., 2002: 136.
Paratetrapedia (Paratetrapedia) velutina; Michener
& Moure, 1957: 416. Pedro, 1996: 251. Pedro &
Camargo, 1999: 202. Sigrist & Sazima, 2004: 38
 SYSTEMATICS OF THE GENUS PARATETRAPEDIA
[floral records: Stigmaphyllon lalandianum, Tetrap-
terys guilleminiana (Malpighiaceae)]. Silveira &
Campos, 1995: 375.
Paratetrapedia flaviventris; Bortoli & Laroca, 1990:
20 [floral record: Senecio oleosus (Asteraceae)].
Paratetrapedia (Paratetrapedia) flaviventris; Silveira
et al., 2002: 136.
Paratetrapedia aff. Velutina; San Martin-Gajardo &
Sazima, 2004: 510 [floral record: Sinningia eumor-
pha (Gesneriaceae); Brazil: São Paulo].
Paratetrapedia (Paratetrapedia) fervida; Gonçalves &
Melo, 2005: 564. Aguiar, 2007: 622.
Paratetrapedia fervida; Mickeliunas et al., 2006: 253
[Figs 1–4: flower records: Grobya amherstiae
(Orchidaceae)].
Comments and diagnosis: Paratetrapedia fervida can
be distinguished mainly by the first labial palpus and
prementum of female with numerous long stout
curved setae, epistomal suture curved above upper
margin of clypeus, male S2 with one isolated row of
setae on mid portion (Fig. 17), S3 wide concave in ‘U’
shape, covered by dense short plumose hairs (Fig. 17),
female T4–T5 with marginal hair band occupying
entire margin, and male hind basitarsus with acute
tooth on proximal third of anterior margin. Moure
(1996) presents a very detailed redescription of the
female lectotype of Tetrapedia flaviventris Friese.
Variation: Paratetrapedia fervida has a frons with
coarse punctures whose density varies from sparse
(2 pd) to dense (0.5 pd). The colours of integument,
pubescence, tibial spurs, and wing membrane vary
amongst specimens. Specimens from the northern
portion of the distribution (Bahia, Espírito Santo,
Minas Gerais, Rio de Janeiro, São Paulo) have terga
pale brown to yellow, sterna usually yellow, and wing
membrane pale brown infumate, whereas those from
the southern portion (Paraná, Santa Catarina) have
terga and sterna completely dark brown and wing
membrane dark brown infumate with black microtri-
chia. One specimen from Viçosa (Minas Gerais) pre-
sents two pale yellow stripes on disc of mesoscutum.
One specimen from Ritápolis (Minas Gerais) and
other from Ipanema (Minas Gerais) have sterna com-
pletely yellow and lateral portions of T3–T6 yellow.
The pubescence of hind leg can vary from completely
dark brown, almost black, to orange yellow. The
yellow marks on paraocular area, clypeus, and
labrum can vary from very conspicuous, slightly faint,
or can be completely absent. A few male specimens
from Espírito Santo have the tooth on hind basitarsus
very reduced.
Male. Body length: 8.5–11.3; maximum head width:
2.8–3.5; fore wing length (including tegula): 8.4–11.0.
© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
391
Female. Body length: 9.5–9.6; maximum head width:
3.1–3.6; fore wing length (including tegula): 8.3–8.7.
Distribution: Argentina: Misiones; Brazil: Bahia,
Espírito Santo, Mato Grosso, Minas Gerais, Paraná,
Pernambuco, Santa Catarina, Rio de Janeiro, São
Paulo; Paraguay (Fig. 70).
Type material: Tetrapedia fervida Smith; holotype
male (BMNH), ‘Type\ H.T.’ ‘B.M. Type\ Hym.\
17B.882’ ‘Tetrapedia fervida Sm.\ (Type)’ ‘Constan-
cia\ January 1857\ J. Gray\\ 5757’.
Tetrapedia velutina Strand; lectotype male (ZMB):
‘Paraguay\ (San Bernardino)\ K.Fiebrig S.V.’ ‘Type’
‘Tetrapedia\ velutina 씹 1907 Friese det.\ var. Fr.’
‘Tetr. velutina\ (Fr.i.l.) 씹\ Strand det.’ ‘Zool. Mus.\
Berlin’ ‘sem dobra episternal\ basitarso armado’.
Tetrapedia bunchosiae Friese; lectotype male
(ZMB): ‘Brazil\ Blumenau\ 1897’ ‘Tetrapedia 씹\
bunchosiae\ 씹det. Friese 1898\ Fr.’ ‘Type’ ‘Zool.
Mus.\ Berlin’.
Tetrapedia flaviventris Friese; lectotype female
(ZMB): ‘Brazil\ Blumenau\ 1897\ Virgil’ ‘Tetrap. 씸\
flaviventris\ det. Friese 1898\ n.sp.’ ‘Typus’ ‘Zool.
Mus.\ Berlin’ ‘Paratetrapedia\ flaviventris 씸\
(Friese, 1899)\ det. J.S.Moure 1992’.
Tetrapedia obsoleta Schrottky, holotype male
(MZSP), ‘17.838’ ‘씹’ ‘obsoleta 96918’ ‘Paratetrapedia\
obsoleta 씹\ (Schrottky, 1902)\ Holotype\ Pe.J.S.
Moure det. 1987’, information associated to number
17.838 in the registry book of MZSP: ‘familia Anthop.;
det. Ducke 13, coll. C. Schrottky, Jundiahy (S.P.),
Type!’.
Tetrapedia monacha Strand; lectotype female
(ZMB): ‘J.B. Anistis\ Assuncion\ Paraguay\ Sapu-
cay\ 12.I.05’ ‘Type’ ‘T. sapucay\ ensis Schr.?\
(Monacha\ Strand det. m.)’; paralectotype female
(ZMB): ‘J.B. Anistis\ Asuncion\ Paraguay\ Sapucay
씸\ 12.I.05’ ‘Co-Type’ ‘T. sapucayensis\ Schr.?\
(Monacha\ Strand. det. m.’.
Additional material: See Appendix.
PARATETRAPEDIA PUNCTATA SP. NOV.
(FIGS 8, 26, 48, 70, 98, 132, 173, 174)
Comments and diagnosis: Paratetrapedia punctata
can be distinguished by its protuberant supraclypeal
area (Figs 8, 48), frons with very coarse punctures,
terga with fine dense punctures (1 pd), and posterior
margin of male S3 mostly glabrous (in some species
with fine long appressed hairs).
The distribution of P. punctata is closely associated
to the savannahs of central Brazil (Cerrado), includ-
ing marginal areas of Cerrado in southern and north-
ern Brazil (respectively, Paraná’s grasslands and
392 A. J. C. AGUIAR and G. A. R. MELO
lowland areas in Amapá state). Two nests of this
species have been found: one in a rotten branch of
Caryocar brasiliensis, a common Cerrado tree (A. J.
C. A., pers. observ.), and another in an exposed piece
of dead root in an area covered by open grasslands in
the Vila Velha State Park, Paraná (G. A. R. M. & R.
B. Gonçalves, pers. observ.). This seems to indicate
that this species is well adapted to nonforested
habitats.
Description
Holotype male: Body length: 8.5; maximum head
width: 3.1; fore wing length (including tegula): 9.0.
Colour. Integument mostly black. Mandible yellow
with apex black; labrum pale yellow; clypeus black
with yellow stripes on lateral margins and lower
margin; supraclypeal area with small yellow spot on
disc; paraocular area with thin long yellow stripe;
scape mostly yellow, pedicel and flagellomeres black
(Fig. 48). Wing membrane black infumate, with
dense black microtrichia; veins and pterostigma
orange yellow. Tibial spurs mostly white. Pubes-
cence. Mostly white, except vertex, mid and hind
legs, margins of T5–T6 with pubescence mostly
black; mesoscutum and scutellum with pale brown
plumose pubescence. Mandible with long pale hairs,
about 1.2¥ mandible width on base. Paraocular area
and genal area with dense white short plumose
pubescence. Scape with long setae on inner surface,
about 0.3 mm in length. Mesoscutum and scutellum
with short plumose pale brown pubescence; meta-
postnotum with short stout plumose hairs (about
0.1 mm in length); mesepisternum ventrally with
dense white plumose pubescence. T4 with marginal
hair band occupying about one third of margin lat-
erally; T5–T6 with dense marginal hair band of
plumose hairs throughout its margin; S2 with short
row of simple setae on mid portion (Fig. 26); S3 with
two small rows of decumbent thin simple hairs on
mid portion and long plumose hairs laterally; S4 and
S5 typical of genus; S6 with two rows of stout
plumose hairs on margin laterally and short plumose
hairs on margin of apex. Hind leg with pubescence
mostly black; scopa of tibia and basitarsus of hind
leg with dense plumose white hairs on apical
portion. Integument sculpture. Clypeus and supra-
clypeal area with dense and contiguous coarse punc-
tures (< 0.5 pd), intermingled amongst sparse minute
punctures (> 2 pd); frons with dense and contiguous
very coarse punctures (< 0.5–1 pd) (Fig. 8); mesoscu-
tum with dense fine punctures (0.5 pd), intermingled
amongst sparse coarse punctures (2 pd); scutellum
with dense fine punctures (1 pd), intermingled
amongst coarse punctures (< 0.5–3 pd); metapostno-
tum with dense minute punctures (< 0.5–1 pd), inter-
mingled amongst sparse coarse punctures on disc
© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
(1–2 pd); mesepisternum laterally with dense coarse
punctures (< 0.5 pd). Terga with dense fine punctures
(1 pd); terga with thin portion of margin smooth.
Structure. Upper portion of supraclypeal area pro-
jected, with trapezoidal aspect (Figs 8, 48). Lamella
of pronotal collar acute throughout its length. Scutel-
lum disc weakly biconvex; mid line sulcate on pos-
terior margin. Hind basitarsus with short carina,
weakly conspicuous, on proximal third of anterior
margin. Head about 1.2¥ broader than long
(2.4 : 3.1); ratio of lower to upper interocular dis-
tance: 0.76 (1.3 : 1.7); clypeus about 1.8¥ broader
than long (1.3 : 0.7); scape: length 0.8; maximum
width 0.25; length of F1–F3: 0.27, 0.17, 0.22; diam-
eter of F2: 0.22.
Paratype female: Body length: 9.0; maximum head
width: 3.0; fore wing length (including tegula): 8.0.
Colour. Similar to male, except for face without yellow
marks; mandible mostly black. Pubescence. Similar to
male, except for mesoscutum with sparse simple short
setae; T4 with marginal hair band incomplete, with
one small band on mid portion of margin and two
other small bands laterally; T5–T6 with marginal
band of short plumose hairs occupying whole
margins. Integument sculpture. Similar to male,
except for punctures denser; terga with dense fine
punctures on disc (< 1–2 pd), sparse punctures on
lateral portions (> 1 pd) and marginal zone smooth.
Structure. Similar to male, with supraclypeal area
projected. Scutellum disc weakly convex, almost flat;
profile weakly truncate. Head about 1.3¥ broader
than long (3.0 : 2.3); ratio of lower to upper interocu-
lar distance: 0.75 (1.35 : 1.75); clypeus about 2.0¥
broader than long (0.67 : 0.32); scape: length 0.8,
maximum width 0.2; length of F1–F3: 0.25, 0.12, 0.17;
diameter of F2: 0.25.
Variation: The yellow marks on head can be larger or
completely absent. The pubescence of tibia and basi-
tarsus of hind leg can vary between completely
orange yellow to mostly black, usually with apical
portion white. The scutellum can vary from almost
flat, convex, to biconvex with mid line strongly
sulcate. The colour of the tibial spurs and of the
pubescence of terga and sterna can vary between
black and white. The decumbent hairs on mid portion
of S3 can be entirely absent or can form two decum-
bent rows of fine hairs. The marginal band on T4 of
females can be complete or almost absent. The speci-
mens from Maranhão and Amapá have smaller body
size and metasoma pale brown, almost yellow.
Etymology: The species name refers to the densely
punctate integument.
 SYSTEMATICS OF THE GENUS PARATETRAPEDIA
Distribution: Brazil: Amapá, Bahia, Distrito Federal,
Goiás, Mato Grosso, Maranhão, Minas Gerais,
Paraná, Rio de Janeiro, Rondônia, São Paulo
(Fig. 70).
Type material: Holotype male, ‘DZUP\ 023496’
‘Brazil, Minas Gerais,\ Serra do Salitre,\ RPPN
Cachoeira\ do Campo; 30.xii.2004\ AJCAguiar’.
Paratypes: BRAZIL: Amapá: 1 male (IEPA), ‘Brazil-
AP\ Ressaca do Curiaú\ N00:08:45.0/W 51:02:28.6\
06/V/2002\ Charton’; 1 male (IEPA), ‘Brazil-Amapá\
Dist. de Fazendinha\ Campus do INPA\ Ambiente de
mata\ 2003\ J. Madson’; 1 female (IEPA), ‘Brazil-
AP\ Ressaca do Curralinho\ N 00:07:55,1/W
51:06:48,6\ 08/V/2002\ C.Henrique\ J. Chaves’; 1
female (IEPA), ‘Brazil-AP\ Ressaca do Coração\
Escola Agrícola\ 10/V/2002\ (10:30 às 12 hs)\
Charton’; 1 female (IEPA), ‘Brazil-Amapá\ Município
de Macapá\ Lagoa dos Índios\ Coleta com Rede\
23/VIII/2002\ J.Chaves’; 2 females (IEPA), Ressaca
Jardim\ 15/XI/01\ sobre flores\ R. Frazão’; Bahia: 1
female (UEFS), ‘Brazil, Bahia, 12 km\ a NE de
Palmeiras,\ base do Morro do\ Pai Inácio, 4.v.2000,\
Gabriel A.R. Melo’; 1 female, ‘DZUP\ 022309’ ‘Brazil,
Bahia, 12 km\ a NE de Palmeiras,\ base do Morro
do\ Pai Inácio, 4.v.2000,\ Gabriel A.R. Melo’; 1 female
(UEFS), ‘BRAZIL, BA, Lençóis\ 28-29, iv-1999\ Lg.
Gisele’; 1 female, ‘DZUP\ 022252’ ‘Lençois\ Bahia\
Brazil 18.II.1988’ ‘Col: P296 8:40’; 1 male, ‘DZUP\
022263’ ‘Lençois-BA\ 23-III-88\ Celso F. Martins’; 1
female, ‘DZUP\ 023268’ ‘Rio de Contas\ Bahia
Brazil\ 21.03.1980\ F.P. Benton\ perto barragem’; 1
female, ‘DZUP\ 022260’ ‘Rio de Contas-BA\
24.IV.1976\ Enoque&C.Elias col’; Distrito Federal: 1
male (DBAI), ‘1025’ ‘BRAZIL DF\ Reserva do IBGE\
RIP Freitas & GS Freitas\ 10/XII/1996’ ‘Paratetrape-
dia sp. AR 5\ det. RIP Freitas’, 4 males and 6 females
(DBAI), idem except ‘1028’, ‘3848’, ‘3824’, ‘3825’,
‘3879’, ‘3889’, ‘3833’, ‘3866’, ‘3850’; 1 male (DBAI),
‘2220’ ‘BRAZIL DF\ Reserva do IBGE\ RIP Freitas &
GS Freitas\ 27/IX/1996’, 2 females (DBAI), idem
except ‘2240’, ‘2120’; 16 males and 6 females (DBAI),
idem except ‘3958’ ‘16/XII/1996’, ‘3959’, ‘3946’, ‘2796’
‘12/X/1996’; ‘1035’ ‘29/VIII/1996’, ‘1028’ ‘29/VIII/1996’,
‘2524’ ‘04/X/1996’, ‘2484’ ‘04/X/1996’, ‘2434’ ‘04/X/1996’,
‘124’ ‘02/VIII/1996’, ‘2672’ ‘08/X/1996’, ‘3649’ ‘25/XI/
1996’, ‘3598’ ‘14/XI/1996’, ‘1592’ ‘13/IX/1996’; ‘2302’
‘30/IX/1996’, ‘565’ ‘22/VIII/1996’, ‘561’ ‘22/VIII/1996’;
‘2902’ ‘22/X/1996’, ‘2896’ ‘22/X/1996’, ‘2895’ ‘22/X/1996’,
‘2892’ ‘22/X/1996’; ‘19/X/1996’ ‘2868’, ‘19/X/1996’ ‘2816’;
1 female (DBAI), ‘BRAZIL DF\ Reserva do IBGE\
RIP Freitas & GS Freitas\ 29/X/1996’ ‘3195’ ‘Paratet-
rapedia sp. AR 5\ det. RIP Freitas’, 1 female (DBAI),
idem except ‘3222’; 1 female (DBAI), ‘BRAZIL DF\
Reserva do IBGE\ RIP Freitas & GS Freitas\ 12/IX/
1996’ ‘1543’ ‘Paratetrapedia sp. AR 5\ det. RIP
© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
393
Freitas’; 6 females (DBAI), idem except ‘08/XI/1996’
‘3471’, ‘14/VII/1996’ ‘396’, ‘12/IX/1996’ ‘3110’ ‘12/IX/
1996’ ‘3706’, ‘19/X/1996’ ‘2834’, ‘02/XII/1996’ ‘3717’; 1
female (DBAI), ‘BRAZIL DF\ Pq Nacional de
Brasília\ RIP Freitas & GS Freitas\ 25/IX/1996’
‘2114’ ‘Paratetrapedia sp. AR 5\ det. RIP Freitas’; 2
females (DZUP), ‘BRAZILIA\ Água Limpa\ 6-5-
1978\ coll. A.Raw’ ‘Paratetrapedia\ sp.5’, 1 female
(DZUP), idem except ‘11-11-1977’; 2 females (DBAI),
‘Faz Água Limpa\ UNB/Brasília-DF\ 03/12/1988\
col. A.Raw’; 2 females (DZUP), ‘BRAZILIA\ Lago
Sul\ 15.7.1977\ coll. A. Raw\ MUDB’ ‘Peixotoa\ Mal-
pighiaceae’ ‘Paratetrapedia\ sp.5’; 1 female (DZUP),
‘BRAZILIA\ Lago Sul\ 20.3.1977\ coll. A. Raw\
MUDB’ ‘Paratetrapedia\ sp.5’; 1 female (DZUP),
‘Brasília\ Cab. de Veado\ 23.10.82\ A.Raw 19’ ‘Tet-
rapterys\ ambigna’ ‘Paratetrapedia\ sp.5’; 1 female
(DZUP), ‘Brasília\ Cab. de Veado\ 9.10.82\ A.Raw
19’ ‘Tetrapterys\ ambigna’ ‘Banisteriopsis’ ‘Paratetra-
pedia\ sp.5’; 1 male (DZUP), ‘Brasília\ A. Limpa\
25-II-79’ ‘estava dormindo qdo coletada’ ‘Paratetrape-
dia\ sp.5’; 1 male (DZUP), ‘Brasília\ Lago Norte\
5.4.83 A.Raw’; 1 female (DZUP), ‘BRAZILIA\ P.
Norte\ 10.II.1977\ coll. A. Raw’ ‘Paratetrapedia\
sp.5’; 1 female (DZUP), ‘BRAZILIA\ P. Norte\
10.11.1977\ coll. A. Raw\ Pena Norte’ ‘Paratetrape-
dia\ sp.5’; 1 male and 3 females (DZUP), ‘DZUP\
023497’ ‘Brazil, Distrito Federal,\ Brasília, MSPW,\
24.xii.2004 AJCAguiar’, 3 females (DZUP), idem
except ‘023459’, ‘023466’, ‘023468’; Goiás: 1 female,
‘DZUP\ 022838’ ‘15-17-II-2004, 7 Km N\ Alto
Paraíso, Goiás\ 1300-1550 m, Mielke\ & Casagrande
leg.’, 1 female (DZUP), idem except ‘022903’; 1 male
(DZUP), ‘Brazil:GO\ Ch. Veadeiros\ 18.04.97\ Raw,A’
‘Cerrado\ Alto Paraíso’; 1 male, ‘DZUP\ 023684’
‘Dianópolis-GO\ Brazil 12-III-62\ FM Oliveira leg’, 4
females (DZUP), idem except ‘023668’, ‘023671’,
‘023677’, ‘023681’; 1 female (RPSP), ‘Itaberaí do\ Sul-
GO. BRAZIL\ 11-I-1971\ Col. Y.Terada’; 1 female,
‘DZUP\ 023675’ ‘Jataí GO\ Brazil XI 63\ M. Alva-
renga’, 1 female (DZUP), idem except ‘023688’; 1
male, ‘DZUP\ 022280’ ‘Brazil, Goiás, 11 km SE de\
Campos Belos, 13°07′32′S\ 46°44′29′ W, 650 m,
04.iv.2003,\ Melo, Aguiar, Marchi e\ Gonçalves, em
cerradão sobre\ massapé, em Hyptis’; 1 female,
‘DZUP\ 023249’ ‘Brazil, Goiás,\ 12 Km N de Caval-
cante,\ 13°41′32′S47°28′08′W,\ 1130 m, 03.iv.2003,
Melo,\ Aguiar, Marchi e Gonçalves,\ em Malphigui-
aceae 5’, 1 female (DZUP), idem except ‘022273’; 1
female, ‘DZUP\ 023258’ ‘Brazil, Goiás, 2 Km W de\
Teresina de Goiás,\ Fazenda Santa Tereza, 13°47′43′
S 47°17′39′W,\ 800 m, 02.iv.2003, Melo,\ Aguiar,
Marchi e Gonçalves’; 1 male (DZUP), ‘Brazil:
GO\ Ch. Veadeiros\ 0507.12.1996\ Boaventura,\
Freitas&Freitas’ ‘8’ ‘Paratetrapedia\ sp1’; 1 male,
‘DZUP\ 022274’ ‘Brazil, Goiás, Chapada dos\ Vead-
394 A. J. C. AGUIAR and G. A. R. MELO
eiros, Vale Dourado,\ 14°11′S 47°37′W,1200 m,\
31.iii.2003, Melo, Aguiar,\ Marchi e Gonçalves’, 2
males and 1 female (DZUP), idem except ‘023266’,
‘023267’, ‘023256’; 1 male, ‘DZUP\ 023253’ ‘Brazil,
Goiás, Chapada dos\ Veadeiros, Vale Dourado,\
14°12′S\ 47°37′ W, 1100 m, 30.iii.2003, Melo, Aguiar,\
Marchi e Gonçalves’, 5 females (DZUP), idem except
‘023250’ to ‘023252’, ‘023254’, ‘022281’; 1 male,
‘DZUP\ 023264’ ‘Brazil, Goiás, Chapada dos Vead-
eiros\ Fazenda Templo Terra,\ Vale Dourado,\
14°10′38′S 47°38′33′W,\ 1200 m, 01.iv.2003, Melo,\
Aguiar, Marchi e Gonçalves’, 2 females (DZUP),
idem except ‘023265’, ‘023257’; 1 male, ‘DZUP\
023261’ ‘Brazil, Goiás, Chapada dos\ Veadeiros,
Alpes Goianos,\ 13°53′59′S 47°23′49′W,\ 1300 m,
05.iv.2003, Melo,\ Aguiar, Marchi e Gonçalves’, 2
females (DZUP), idem except ‘023263’, ‘023262’; 1
female, ‘DZUP\ 022273’ ‘Brazil, Goiás,\ 12 Km N de
Cavalcante,\ 13°41′32′S47°28′08′W,\ 1130,
03.iv.2003, Melo,\ Aguiar, Marchi e Gonçalves\ em
Malpighiaceae 5’, 1 female (DZUP), idem except
‘023249’; 1 female, ‘DZUP\ 023248’ ‘Brazil, Goiás,\
15 Km N de Cavalcante,\ 13°39′17′S47°28′31′W,\
1140 m, 03.iv.2003, Melo,\ Aguiar, Marchi e
Gonçalves’; 1 male, ‘DZUP\ 023259’ ‘Brazil, Goiás,\
20 Km N de Cavalcante,\ 13°37′40′S47°29′20′W,\
1220 m, 03.iv.2003, Melo,\ Aguiar, Marchi e
Gonçalves’, 1 female (DZUP), idem except ‘023260’; 1
male (MPEG), ‘Brazil GO\ São Domingos\ Fazenda
CIPASA\ 18-IX a 2-X-93’ ‘Dionisio Pimentel’ ‘Malaise’
‘Hymenoptera: Apocrita\ Aculeata:\ Apoidea:\
Apidae:\ Incorp: 17/VII/2002’; 1 female, ‘DZUP\
023255’ ‘Brazil, Goiás, 24 Km NE de\ Teresina de
Goiás,\ 13°34′53′S47°11′07′W,\ 450 m, 05.iv.2003,
Melo\ Aguiar, Marchi e Gonçalves’; Maranhão: 1
female (LEA), ‘Chapadinha-MA\ Brazil 23/V/94\
Brito & Rego Leg\ 0492’ ‘Pl. no. 027 F\ Hr. 10-12’,
‘Paratetrapedia sp2’; 1 male, ‘DZUP\ 022266’
‘Imperatriz-MA\ Brazil-20-II-62\ F.M. Oliveira’, 1
female (DZUP), idem except ‘022261’; Mato Grosso: 2
males (DZUP), ‘MATO GROSSO\ Barra das Garças\
19-4-1978\ coll. A. Raw’; 1 male, ‘DZUP\ 022310’
‘Cáceres, MT\ 11.XII.1984\ C.Elias leg.\ Polono-
roeste’; 2 males and 4 females (DZUP), idem except
‘022316’ ‘12.18.XI.1984’, ‘022323’ ‘24.XII.1984’,
‘022318’ ‘27.III.1985’, ‘022305’ ‘3.III.1985’, ‘022312’
‘7.II.1985’, ‘022320’ ‘7.II.1985’, ‘022241’ ‘9.I.1985’,
‘022321’ ‘9.I.1985’; 1 male, ‘DZUP\ 022314’ ‘Chapada
MT\ Brazil 27-X-61\ F.M. Oliveira leg’, 4 females
(DZUP), idem except ‘022306’, ‘022308’, ‘022317’,
‘022319’; 1 male, ‘DZUP\ 022322’ ‘Chapada MT\
Brazil XI-63\ M. Alvarenga’; 1 female (IBAI), ‘Ch.
Guimarães-MT\ Manso\ I.R.D. Rocha\ 3.12.88’; 1
female (DBAI), idem except ‘10.12.88’; 1 female,
‘DZUP\ 022307’ ‘Chap. Guimarães-MT\ 28.III-
IV.1983\ Exc. Dep. Zool.-UFPR\ (Polonoroeste)’; 1
© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
male, ‘DZUP\ 022311’ ‘Chapada dos\ Guimarães,
MT\ 8-I-1987\ C. Elias, leg.’; 1 male, ‘DZUP\ 022300’
‘Chapada de Guimarães\ MT-Brazil 1-4-II-65\
Sebastião Laroca leg’, 1 female (DZUP), idem except
‘022299’; 1 male, ‘DZUP\ 022303’ ‘Rosário Oeste\ MT
Brazil XI-63\ M. Alvarenga leg’; 1 male, ‘DZUP\
022304’ ‘São Félix M.Grosso\ Brazil 21-28/VII/68\ C.
Elias leg’; 1 male (RPSP), ‘Serra das Araras-MT\
Brazil 19.27-IX-1987’ ‘SD 21,57°14′W, 15°27′S\
Gimenes leg. 871130’; 1 male, ‘DZUP\ 022242’ ‘Serra
do Roncador\ R.S. Base Camp.-MT\ Brazil-17/7/
1968\ Claudionor Elias’; 1 female (RPSP), ‘Bra-
zil.MT.BR 364, Km7\ Serra de São Vicente\ 15o30′S/
55o12′W\ 5,8.II.1997-M.L. Oliveira’; 1 male, ‘DZUP\
022315’ ‘Serra Roncador\ MT-R.S. Base Camp\
Brazil-13/7/1968\ Laroca & Azevedo’; Mato Grosso do
Sul: 1 male and 1 female (SEMC), ‘Rio Caraguata\
Matto Grosso\ III-53 Brazil\ F. Plaumann’; Minas
Gerais: 1 female, ‘DZUP\ 022253’ ‘Araxá-MG-Brazil\
20-V-1965\ C. &T.Elias leg’; 1 female, ‘DZUP\
022295’ ‘ARAXÁ-MG-BRAZIL\ 5-IX-1965\ C. & T.
Elias leg’; 1 male, ‘DZUP\ 023686’ ‘Barbacena-MG\
Brazil 14-16-II-62\ M. Alvarenga leg.’; 1 female,
‘DZUP\ 22286’ ‘Coleção\ Campos Seabra’ ‘Belo Hori-
zonte\ M.Gerais Brazil\ VIII-1961\ F.M.Oliveira’, 1
female (DZUP), idem except ‘022292’; 1 female,
‘DZUP\ 022283’ ‘Brazil, Minas Gerais,\ Brumadinho,
Serra da\ Moeda, 18.iv.1998,\ 20°05′S 43°59′W,\
1400 m G.A.R.Melo’; 1 female, ‘DZUP\ 022244’
‘Campos Altos-MG\ BRAZIL 28/8/965\ C. Elias leg’; 1
female (FCVZ), ‘BRAZIL, MG, Capitólio,\ Rio Turvu,
15.V.1999\ 20°38′S,46°13′W.\ 950 m. F. Zanella leg.’;
1 male, ‘DZUP\ 022096’ ‘Cássia-MG\ Brazil- 13-II-
63\ Claudionor Elias’, 1 male (DZUP), idem except
‘022112’; 1 male (MNRJ), ‘M. Gerais, Brazil\ C. Rio
Claro-47\ Carvalho col.’; 1 female, ‘DZUP\ 022094’
‘Brazil, Minas Gerais,\ Corinto, 16-31.viii.\ 1979, C.
Elias leg.’; 1 female, ‘DZUP\ 022251’ ‘Curvelo-MG.\
IV/1980.\ col. C.Elias’; 1 male, ‘DZUP\ 022248’ ‘Ibiá-
Brazil\ MG-10/12/1965\ C.Elias leg.’, 1 male (DZUP),
idem except ‘022095’; 1 male, ‘DZUP\ 022098’ ‘Ibiraci-
MG\ Brazil-X-61\ C. Elias leg.’, 5 females (DZUP),
idem except ‘022258’, ‘022109’, ‘022104’, ‘022101’,
‘022099’, ‘022098’; 1 male, ‘DZUP\ 022093’ ‘Jacui-
MG\ Brazil 27-XI-63\ C. Elias, leg’; 1 male, ‘DZUP\
022268’, ‘Paraopeba, MG, Brazil\ Data 06/05/1987\
F.A. Silveira’ ‘303/785’ ‘Paratetrapedia\ no16’; 1 male,
‘DZUP\ 022269’ ‘Paraopeba, MG, Brazil\ Data 07/06/
1987\ F.A. Silveira’ ‘343/886’ ‘Paratetrapedia\ no3’; 1
female, ‘DZUP\ 022267’ ‘147/407’ ‘Paraopeba, MG,
Brazil\ Data 04/12/1986\ F.A. Silveira’, ‘Paratetr.\
no3’; 1 female, ‘DZUP\ 022264’ ‘277/690’ ‘Paraopeba,
MG, Brazil\ Data 08/04/1987\ F.A. Silveira’
‘Paratetr.\ no7’; 1 male, ‘DZUP\ 023667’ ‘Passos MG\
Brazil 4-10-X 62\ Claudionor Elias’; 1 female,
‘DZUP\ 041306’ ‘Passos MG\ Brazil 12 XII 63\ C.
 SYSTEMATICS OF THE GENUS PARATETRAPEDIA
Elias leg’; 1 female, ‘DZUP\ 022255’ ‘Passos MG\
Brazil 13-18 XI-61\ Claudionor Elias’; 1 female,
‘DZUP\ 022102’ ‘Passos MG\ Brazil 20-25 XI 61\ C.
Elias leg.’; 1 female, ‘DZUP\ 023465’ ‘Passos MG\
Brazil 21-28 III-62\ C. Elias leg’; 1 male, ‘DZUP\
022100’ ‘Passos MG\ Brazil IX 1961\ C. Elias leg.’, 1
male and 3 females (DZUP), idem except ‘022240’,
‘023469’, ‘022108’, ‘023660’; 1 female, ‘DZUP\ 023464’
‘Passos MG\ Brazil VI-1961\ C. Elias leg’; 1 male,
‘DZUP\ 022106’ ‘Passos MG Brazil\ 5-10-XI 1961\ C.
Elias leg.’, 1 male and 2 females (DZUP), idem except
‘022097’, ‘023648’, ‘022103’; 1 female, ‘DZUP\ 022105’
‘Passos-Brazil\ M.Gerais-6/1963\ C.Elias legit’; 1
male, ‘DZUP\ 023655’ ‘Passos-Brazil\ MG 28-31-X-
63\ Claudionor Elias’; 1 male, ‘DZUP\ 023654’
‘Passos-MG\ Brazil- 10/1963\ C. & T. Elias leg.’, 2
males (DZUP), idem except ‘023661’, ‘023664’; 1
female, ‘DZUP\ 023113’ ‘Passos-MG\ Brazil 19-24
III-62\ Claudionor Elias’; 1 female, ‘DZUP\ 022249’
‘Patos de Minas-MG\ Brazil-23/11/965\ Claudionor
Elias’; 1 female, ‘DZUP\ 022257’ ‘Patrocínio-MG\
5/10/1965\ Claudionor Elias’; 1 female, ‘DZUP\
022247’ ‘Perdizes-MG\ Brazil, 7-65\ C.Elias Leg’; 1
female, ‘DZUP\ 023462’ ‘Pratápolis-MG\ Brazil 5-II-
63\ Claudionor Elias’; 1 male (AMNH), ‘Brazil,
Minas\ Gerais: Santa Vitoria\ February 1970\ F.H.
Oliveira’; ‘147/407’; 1 female, ‘DZUP\ 022254’ ‘P. de
Caldas-MG\ Brazil XI-61\ Claudionor Elias l.’; 1
male (RPSP), ‘São Roque de Minas\ MG-Brazil-SF
23\ 46°25′W, 20°15′S’ ‘Cerrado-12, 12/01/1992\ Alt.
850 1000 m\ Moure, Camargo,\ Serguei, Pedro leg’
‘920575’; 1 female, ‘DZUP\ 022256’ ‘S.do Salito MG\
Brazil 4 IV 65\ C.Elias leg’, 1 female (DZUP), idem
except ‘022259’; 1 male, ‘DZUP\ 023495’ ‘BRAZIL,
MG, Serra do\ Salitre, RPPN Cachoeira\ do Campo,
24-30.xii.2003,\ AJCAguiar leg.’, 1 male (DZUP),
idem except ‘023499’; 1 male, ‘DZUP\ 023467’ ‘Brazil,
Minas Gerais,\ Serra do Salitre,\ RPPN Cachoeira\
do Campo; 30.xii.2004\ AJCAguiar’, 1 male and 1
female (DZUP), idem except ‘023498’, ‘023460’; 1
female, ‘DZUP\ 023678’ ‘Brazil, Minas Gerais,\ Serra
do Salitre,\ RPPN Cachoeira do\ Campo, vii.2001,\
A.J.C. Aguiar’; 1 female (DZUP), ‘Brazil, Minas
Gerais,\ Serra do Salitre,\ RPPN Cachoeira do\
Campo, vii.2001,\ A.J.C. Aguiar’; 1 female, ‘DZUP\
022271’ ‘Tapira-MG\ Brazil 27-V-65\ C. Elias leg’; 1
male, ‘DZUP\ 022287’ ‘Uberaba MG\ Brazil x-61\
C.Elias leg.’; 1 female, ‘DZUP\ 022284’ ‘Uberlândia-
MG\ Brazil 11.VI.93\ G.A.R.Melo’; 1 male, ‘DZUP\
022282’ ‘Uberlândia-MG\ Brazil, 18.02.92\ A.A.A.
Barbosa’ ‘Est. Ecol. Panga\ no. 20’; 1 male, ‘DZUP\
022285’ ‘Uberlândia-MG\ Brazil,19.03.93\ A.A.A.
Barbosa’ ‘Est. Ecol. Panga\ no. 219’; 1 male, ‘DZUP\
022291’ ‘Brazil, Minas Gerais,\ Uberlândia, Est.
Ecol.\ Panga, 13.v.1993,\ Gabriel A.R.Melo’; 1 female,
‘DZUP\ 022293’ ‘Brazil, Minas Gerais, 9km\ a SW de
© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
395
Capitólio,\ 22.xii.1999, G.A.R. Melo’; 1 male, ‘DZUP\
022290’ ‘Varginha MG\ Brazil IX 1961\ Alvarenga
leg’, 1 male and 4 females (DZUP), idem except
‘023666’, ‘022288’, ‘023458’, ‘023461’, ‘023463’,
‘023470’; Paraná: 1 male (AMNH), ‘Brazil, Paraná:\
Curitiba\ February 4, 1974’ ‘J.G.Rozen\ R.C. Thomp-
son\ collectors’; 1 female (DZUP), ‘Brazil, Paraná\
15 km a E de Tibagi,\ 24°31′S50°17′W,\ 02.xi.2001,\
G.A.R. Melo’; Rio de Janeiro: 1 male, ‘DZUP\ 022313’
‘ITATIAIA GB\ Brazil V 61\ F.M. Oliveira’; Rondônia:
1 female (RPSP), ‘Brazil. Rondônia\ Guajara Mirim\
Sa. Pacaas Novos’ 16/junho/1995\ M.L. Oliveira’
‘PN0210’; 1 female (RPSP), ‘Brazil-RO.\ Guajara
Mirim.\ Pacaás Novos.’ ‘Data 22/10/1995\ M.L.
Oliveira’ ‘PN0002’; 1 female (RPSP), ‘Brazil-RO.\
Guajara Mirim.\ Pacaás Novos.’ ‘Data 16/VI/1995\
M.L. Oliveira’ ‘PN0260’; 1 male (MPEG), ‘Vilhena\
23-2-1961’ ‘Brazil, RO\ J. & B.Bechyné’; 1 male and 1
female (INPA), ‘Brazil, Rondônia\ Guajará Mirim\ Sa
Pacaás Novos’ ‘10°48S’, 65°22′W\ 12-14/X/2001\
Oliveira, Morato &\ Cunha leg.’; 1 female, ‘DZUP\
022245’ ‘Vilhena, RO\ 4.XI.1986\ C.Elias leg.\
POLONOROESTE’; 1 female (DZUP), idem except
‘023270’ ‘4/12/1986’; São Paulo: 1 male (SM), ‘Cerrado-
Est. Ecol. Jataí-SP.\ BR 27 IX 1992, no. 2520\ h:
10-12, Mateus S. leg.’, 2 males (SM), idem except
‘2519’, ‘2513\ h: 14–16’; 1 female (SM), ‘Cerrado-Est.
Ecol. Jataí-SP.\ BR 9 XII 1992, no. 2531\ h: 14–16,
Mateus S. leg.’, 2 females (SM), idem except ‘2584 h:
10–12’, ‘2582 h: 8–10’; 1 females (SM), ‘Cerrado-Est.
Ecol. Jataí-SP.\ BR 16 III 1993, no. 2888\ h: 8–10,
Mateus S. leg.’, 1 female (SM), idem except ‘2898’; 1
female (SM), ‘Cerrado-Est. Ecol. Jataí-SP.\ BR 27 IX
1993, no. 2489\ h: 8–10, Mateus S. leg.’, 1 female
(SM), idem except ‘2514 h: 10–12’; 1 female (SM),
‘Cerrado-Est. Ecol. Jataí-SP.\ BR 03 XI 1992, no.
2477\ h: 8–10, Mateus S. leg.’, 1 female (SM), idem
except ‘2476’; 1 male (SM), ‘Cerrado-Est. Ecol. Jataí-
SP.\ BR 19 IV 1993, no. 2996\ h: 10–12, Mateus S.
leg.’; 1 male (SM), ‘Cerrado-Est. Ecol. Jataí-SP.\ BR
19 XII 1991, no. 0389\ h: 8–10, Mateus S. leg.’; 1 male
(SM), ‘Cerrado-Est. Ecol. Jataí-SP.\ BR 20 I 1993, no.
2709\ h: 8–10, Mateus S. leg.’; 1 male (SM), ‘Cerrado-
Est. Ecol. Jataí-SP.\ BR 22 XII 1992, no. 2611\ h:
10–12, Mateus S. leg.’; 1 female (SM), ‘Cerrado-Est.
Ecol. Jataí-SP.\ BR 06 IV 1993, no. 2935\ h: 10–12,
Mateus S. leg.’; 1 male (SM), ‘Cerrado-Est. Ecol.
Jataí-SP.\ BR 07 I 1993, no. 2677\ h: 14–16, Mateus
S. leg.’; 1 female, ‘DZUP\ 023647’ ‘Pedregulho-SP\
Brazil 8-XI-62\ Claudionor Elias’; 1 female, ‘DZUP\
023000’ ‘Rifaina-Brazil\ SP-28/10/1965\ C.Elias leg.’,
5 females (DZUP), idem except ‘023685’, ‘023679’,
‘023678’, ‘023676’, ‘023673’; 1 female, ‘DZUP\ 022243’
‘Rifaina-SP\ Brazil\ 20-IX-1965\ C. Elias leg.’; 1
female (SEMC), ‘BRAZIL: São Paulo\ George
Deterer\ September 1961\ (W. Bokermann)’; 1 female
396 A. J. C. AGUIAR and G. A. R. MELO
(IBUSP), ‘IB-USP-SP Brazil\ 9:20hs 25/02/03\ planta
col. S. Naxara’.
PARATETRAPEDIA ROMANI (FRIESE, 1923)
(FIGS 59, 73, 102, 134, 177, 178)
Tetrapedia romani Friese 1923: 3; lectotype male,
designated by Aguiar (2007), Brazil: Amazonas,
Manaus (NHRS).
Paratetrapedia romani; Aguiar (2007): 625.
Paratetrapedia (Paratetrapedia) romani; Rasmussen
& Ascher (2008): 90.
Comments and diagnosis: Paratetrapedia romani is
the only species of the lineata group in which the
male S3 lacks a deeply concave ‘U’-shaped area
covered with dense short plumose pubescence.
Despite this, other features clearly associate it with
the lineata group: long hairs on margin of mandible,
lower portion of genal area, coxa, trochanter, and
lower mesepisternum of male; male S2 with one short
isolated row of setae on mid portion of margin; apical
portion of male S7 with lateral lobes in a weakly
acute angle; male hind basitarsus with acute tooth on
anterior margin; gonostyli with large lobe on basal
portion; row of setae on margin of female S2–S3
forming a weakly semicircular line on mid portion
(Fig. 30). The female of P. romani is very similar to
that of P. alsinai, being distinct by the marginal hair
band of T4 occupying about one third of the margin
laterally.
Distribution: Brazil: Amazonas, Pará (Fig. 73).
Redescription: Lectotype male. Body length: 9.6;
maximum head width: 2.9; fore wing length (includ-
ing tegula): 8.8. Colour. Integument mostly orange
yellow. Mandible yellow with apex black; disc of frons,
upper paraocular area and vertex black; frons with
thin yellow stripe on mid line, contacting supracly-
peal area and mid ocellus; genal area completely
yellow; scape, pedicel, and flagellomeres orange
yellow. Mesoscutum black with two thin yellow
stripes on lateral margins (Fig. 59); mid portion of
lateral stripe on mesoscutum weakly enlarged; scutel-
lum yellow; axillar fossa black. Terga orange yellow;
marginal zone weakly brown. Wing membrane yellow
infumate; veins and pterostigma orange yellow;
microtrichia orange yellow; apical third of fore wing
dark because of brown microtrichia. Hind basitarsus
weakly dark brown; tibial spurs white. Pubescence.
Mostly pale yellow, except marginal hair bands of
T3–T4 and scopa of mid and hind basitarsus with
dark brown pubescence. Mandibles, lower mesepister-
num, coxa, and trochanter of fore and mid legs, and
low mesepisternum with long simple thin pale hairs,
© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
about two times longer than width of mandibular
base. Scape with sparse pale yellow setae, c. 0.3 mm
in length. Mesoscutum and scutellum with short
golden yellow, sparse plumose pubescence; plumose
hairs on scutellum about two times longer than on
mesoscutum; metapostnotum with thin setae, weakly
branched, about 0.2 mm in length. T4–T6 with dense
marginal hair band of plumose pale yellow hairs
occupying entire margin. S2 with short row of setae
on mid portion of margin; S3 with row of short simple
setae on margin, with a short interval on mid portion;
S4 and S5 typical of genus; S6 with two rows of stout
plumose setae on lateral margins and short plumose
hairs on margin of apex. Integument sculpture.
Clypeus and supraclypeal area with sparse coarse
punctures (1–2 pd); frons with sparse coarse punc-
tures (1–2 pd). Mesoscutum and scutellum with dense
minute punctures (< 0.5 pd), intermingled amongst
sparse coarser punctures (> 2 pd); metapostnotum
with sparse fine punctures (> 3 pd); mesepisternum
laterally mainly with sparse coarse punctures
(> 2 pd), except lower third with dense coarse punc-
tures (1 pd). Structure. Pronotal collar with lamella
acute throughout its length. Scutellum disc weakly
convex. Hind basitarsus with one thin acute tooth on
anterior margin. Head about 1.3¥ broader than long
(2.9 : 2.25); ratio of lower interocular distance to
upper interocular distance: 0.82 (1.25 : 1.52); clypeus
about 1.9¥ broader than long (2.4 : 1.35); scape:
length 0.75, maximum width, 0.25; length of F1–F3:
0.2, 0.2, 0.25; diameter of F2: 0.2.
Paralectotype female: Body length: 7.9; maximum
head width: 2.9; wing length (including tegula): 8.5.
Colour. Similar to male, except that legs completely
orange yellow. Pubescence. Similar to male, except for
metapostnotum with very short hairs; T4 with mar-
ginal hair band occupying about one third of margin
laterally; marginal hair band on T5–T6 with simple
hairs, on T6 with golden yellow hairs. Integument
sculpture. Similar to male, except antennal scrobe
with dense fine punctures (1 pd); mesepisternum
laterally with dense coarse punctures (0.5–2 pd);
metapostnotum with dense fine punctures (1 pd),
with smooth integument amongst punctures; lateral
margins of metapostnotum with large smooth area.
Structure. Similar to male, except for pronotal collar
with lamella weakly obtuse laterally. Scutellum disc
weakly convex, with profile almost in right angle.
Head about 1.3¥ broader than long (2.93 : 2.25); ratio
of lower interocular distance to upper interocular
distance: 0.9 (1.32 : 1.57); clypeus about two times
broader than long (1.4 : 0.7); scape: length 0.8,
maximum width 0.22; length of F1–F3: 0.25, 0.12,
0.18; diameter of F2: 0.22.
 SYSTEMATICS OF THE GENUS PARATETRAPEDIA
Variation: Most examined specimens have yellow
stripes on lateral margins of mesoscutum with mid
portion very enlarged (Figs 56, 59); however, one
specimen from Manaus has narrow stripes (as in
Figs 57, 58). Half of the specimens have the yellow
stripe on mid line of frons isolated on disc, and in the
other half, the stripe contacts the yellow mark on the
supraclypeal area.
Type material: Tetrapedia romani Friese; lectotype
male (NHRS): ‘Mana\ os’ ‘Amazon\ Roman’ ‘mars’
‘Tetrapedia\ romani\ 씹 1920 Friese Fr. det.’; para-
lectotype male (NHRS): ‘Mana\ os’ ‘Amazon\ Roman’
‘juli’ ‘romani\ 씹 1920 Friese Fr. det.’.
Additional material: See Appendix.
PARATETRAPEDIA TOCANTINENSIS SP. NOV.
(FIGS 58, 73, 103, 135, 179, 180)
Comments and diagnosis: Paratetrapedia tocantinen-
sis is very similar to P. lineata in colour pattern, body
size, and shape. It differs from P. lineata mainly by
the metapostnotum with sparse fine punctures (2 pd),
with smooth and shiny integument amongst punc-
tures; epistomal suture straight above upper margin
of clypeus; lamella of pronotal collar obtuse in the
lateral portions; mesepisternum and propodeum
usually yellow. Males can be differentiated by a con-
spicuously acute tooth on anterior margin of hind
basitarsus, and T4 with marginal hair band occupy-
ing one third of margin laterally. In the females, the
lateral yellow stripes of mesoscutum are fused with
the discal stripes along the anterior margin of the
sclerite (Fig. 58).
Description
Holotype male: Body length: 7.8; maximum head
width: 2.6; fore wing length (including tegula): 7.0.
Colour. Integument bicoloured (reddish brown and
yellow). Mandible yellow with apex black; labrum,
clypeus, and supraclypeal area yellow; paraocular
area with yellow stripe occupying more than the
lower half of paraocular area, with upper portion
obtuse, weakly sinuous; frons dark brown with one
thin yellow stripe along the mid line; genal area
yellow; scape orange yellow; pedicel and flagellomeres
with anterior surface yellow and posterior surface
reddish brown. Mesoscutum black with two thin
yellow stripes on disc and lateral margins; scutellum
and metanotum yellow; axilla brown; anterior surface
of mesepisternum reddish brown. Terga yellow with
marginal zone reddish brown. Wing membrane
mostly hyaline, weakly yellow infumate on radial cell;
microtrichia yellow on basal portion of wing; veins
and pterostigma orange yellow. Legs yellow; hind
© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
397
tibia and basitarsus weakly brown; tibial spurs white.
Pubescence. Mostly white to pale yellow; except
mesoscutum, scutellum, tergal margins, and legs with
pubescence weakly orange yellow; T4 with marginal
hair band brown. Margin of mandible, lower portion
of genal area, coxae, and trochanters of fore and mid
legs, and low mesepisternum with long fine hairs,
length about twice the width of mandibular base.
Scape with long dark brown setae on inner face, about
0.12–0.6 mm in length. Mesoscutum and scutellum
with dense short plumose yellow pubescence; meta-
postnotum with sparse short plumose hairs, about
0.12 mm in length. T4 with marginal hair band of
dark brown hairs on about one third of margin later-
ally; T5–T6 with complete marginal hair band of
dense long orange yellow plumose hairs; S2 with one
short row of simple setae isolated on mid portion of
margin; S3 of male deeply concave in ‘U’ shape,
covered by dense short plumose pubescence; S4 and
S5 typical of genus; S6 with two rows of stout plumose
hairs on lateral margins and short plumose hairs on
margin of apex. Integument sculpture. Clypeus with
coarse punctures, mainly sparse (1–2 pd), inter-
mingled amongst sparse minute punctures (> 2 pd);
frons with coarse punctures (0.5–2 pd). Mesoscutum
and scutellum with dense minute punctures (0.5 pd),
intermingled amongst sparse coarser punctures
(> 2 pd); metapostnotum with sparse fine punctures
(2 pd) intermingled amongst integument smooth and
shiny; mesepisternum laterally mainly with sparse
coarse punctures (2 pd); punctures densely distrib-
uted on omaular area (0.5–1 pd). Structure. Epis-
tomal suture straight above upper margin of clypeus.
Lamella of pronotal collar obtuse laterally in dorsal
view. Scutellum weakly convex, almost flat. Hind
basitarsus with acute tooth on anterior margin. Meta-
soma elongated, about two times longer than broad
(3.7 : 1.8). Head about 1.2¥ broader than long
(2.6 : 2.05); ratio of lower interocular distance to
upper interocular distance: 0.8 (1.12 : 1.40); clypeus
about 1.8¥ broader than long (1.1 : 0.6); scape: length
0.7, maximum width 0.22; length of F1–F3: 0.18, 0.12,
0.24; diameter of F2: 0.22.
Paratype female: Body length: 8.0; maximum head
width: 2.7; fore wing length (including tegula): 6.6.
Colour. Similar to male, except for yellow stripe on
paraocular area with upper portion obtuse; yellow
stripes on mesoscutum contiguous on anterior margin
of mesoscutum (Fig. 85). Pubescence. Similar to male;
except for few long setae on margin of mandible;
metapostnotum with very short hairs (< 0.1 mm); T4
with marginal hair band on whole margin; T5–T6
with marginal band of simple hairs occupying whole
margin; row of setae on sternal margin oval sinuated
on mid portion. Integument sculpture. Similar to
398 A. J. C. AGUIAR and G. A. R. MELO
male, but denser; clypeus and supraclypeal area
with dense coarse punctures (0.5–1 pd); disc of frons
with dense coarse punctures (0.5–1 pd), and lateral
portions with dense fine punctures (< 0.5 pd). Meta-
postnotum with dense fine punctures (1–2 pd), inter-
mingled amongst smooth and shiny integument;
mesepisternum laterally with sparse punctures,
except for dense coarse punctures on omaular area
(0.5–1 pd). Structure. Similar to male. Metasoma
about 1.6¥ longer than broad (4.2 : 2.6); head about
1.2¥ broader than long (2.7 : 2.2); ratio of lower
interocular distance to upper interocular distance: 0.9
(1.35 : 1.5); clypeus about two times broader than long
(1.25 : 0.6); scape: length 0.75, maximum width 0.2;
length of F1–F3: 0.18, 0.12, 0.7; diameter of F2: 0.2.
Variation: Most specimens have the mesepisternum
entirely yellow, but in one specimen from Piauí and
another from Goiás, it is reddish brown.
Etymology: The species name refers to the Tocantins
river in central Brazil.
Distribution: Brazil: Goiás, Maranhão, Piauí,
Tocantins (Fig. 73).
Type material: Holotype male, ‘DZUP\ 022501’
‘Brazil, Goiás, 11 km SE de\ Campos Belos,
13°07′32′S\ 46°44′29′ W, 650 m, 04.iv.2003,\ Melo,
Aguiar, Marchi e\ Gonçalves, em cerradão sobre\
massapé, em Hyptis’; paratypes: BRAZIL: Goiás: 1
female, ‘DZUP\ 021840’ ‘Brazil, Goiás, 11 km SE de\
Campos Belos, 13°07′32′S\ 46°44′29′ W, 650 m,
04.iv.2003,\ Melo, Aguiar, Marchi e\ Gonçalves, em
cerradão sobre\ massapé, em Hyptis’, 2 males and 4
females (DZUP), idem except ‘022504’, ‘022496’,
‘021834’ to ‘021836’; Maranhão: 1 female, ‘DZUP\
026578’ ‘Brazil, MA, Urbano Santos\ Reserva Tonico
Bacelar\ 3°12′28S/43°24′12′W\ 24.i.2004; C.M. Maia’;
Piauí: 1 female (RPSP), ‘30 Km NW Bom Jesus\ PI
Brazil 25,29 I 1993\ SC23, 44°35′W8°57′S’ ‘Camargo,
Tavares\ Pedro leg. 931552’; Tocantins: 1 male,
‘DZUP\ 023439’ ‘Buriti do Tocantins\ TO, Brazil\
15-XII-1999\ C.A.L. de Carvalho leg.’ ‘Paratetrape-
dia\ lineata\ (Spinola, 1851)\ F.F. de Oliveira\ Det.
2000’, 1 female (DZUP), idem except ‘023421’.
PARATETRAPEDIA FLAVIFRONS SP. NOV.
(FIGS 43, 73, 101, 133, 187, 188)
Comments and diagnosis: Paratetrapedia flavifrons is
very similar to P. romani and P. alsinai in the yellow
infumate wing membrane, being distinct, however, by
the presence of a very large yellow spot on disc of
© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
frons (Fig. 43), the orange yellow lateral portions of
mesoscutum, and male basitarsus with only a short
tooth on anterior margin.
Distribution: Brazil: Amazonas, Mato Grosso,
Rondônia, Pará (Fig. 73). [Note: data from a specimen
from Pará labelled ‘BR-14, km 93\ Belém – Brasília’
were not included in the map (Fig. 73) because this
specimen represents a doubtful record from a locality
far away from the main distribution of the species.]
Description
Holotype male: Body length: 9.0; maximum head
width: 2.75; fore wing length (including tegula): 9.2.
Colour. Integument bicoloured (reddish brown and
orange yellow), mostly orange yellow. Mandible
yellow with apex black; labrum, clypeus, and supra-
clypeal area yellow; paraocular area with wide yellow
stripe, with upper portion obtuse (Fig. 43); frons with
one yellow stripe along mid line (about 1.5 ¥ F2 diam-
eter; Fig. 43); genal area yellow; scape yellow; pedicel
and flagellomeres reddish brown. Mesoscutum with
two thin yellow stripes on disc and lateral margins;
disc of mesoscutum black and lateral portions orange
yellow; axilla dark brown. Hind tibia and basitarsus
dark brown; tibial spurs white. Wing membrane
orange yellow infumate; microtrichia yellow; fore
wing membrane with apical portion dark with brown
microtrichia; veins and pterostigma yellow. Terga
completely orange yellow, weakly darker on marginal
zone. Pubescence. Mostly pale yellow, except the basi-
tarsi of mid and hind leg, and marginal hair band of
T4, which have dark brown hairs. Outer margin of
mandible, lower genal area, mesepisternum ventrally,
coxae and trochanters with long fine white hairs
(length more than twice width of mandibular base).
Scutellum with long erect simple setae (about
0.16 mm in length); metapostnotum with short
plumose hairs (about 0.12 mm in length). T4 with
marginal hair band throughout its margin; T5–T6
with dense plumose yellow pubescence throughout its
margin; S2 with one short row of simple setae on mid
portion of margin; S3 deeply concave in ‘U’ shape,
covered by dense short plumose pubescence; S4 and
S5 typical of genus; S6 with two rows of stout plumose
setae on lateral margins and short plumose hairs on
margin of apex. Integument sculpture. Clypeus with
coarse punctures, mostly sparse (2 pd), intermingled
amongst sparse minute punctures (2 pd); supracly-
peal area with disc mostly smooth and lateral
margins with coarse punctures (1–3 pd); disc of frons
with coarse sparse punctures (< 0.5–2 pd); antennal
scrobe with dense fine punctures (< 0.5 pd). Mesoscu-
tum with dense fine punctures (0.5–1 pd), inter-
mingled amongst sparse coarser punctures (> 2 pd);
metapostnotum with dense fine punctures (2 pd),
 SYSTEMATICS OF THE GENUS PARATETRAPEDIA
with integument smooth and shiny amongst punc-
tures; mesepisternum with dense coarse punctures
(1 pd). Structure. Lamella of pronotal collar mostly
acute with lateral portions weakly obtuse. Disc of
scutellum weakly convex, almost flat; mid line
slightly concave. Hind basitarsus with one very small
tooth on proximal third of anterior margin. Head
about 1.2¥ broader than long (2.75 : 2.2); ratio of
lower interocular distance and upper interocular dis-
tance: 0.93 (1.17 : 1.25); clypeus about 1.5¥ broader
than long (1.1 : 0.7); scape: length 0.7, maximum
width 0.2; length of F1–F3: 0.21, 0.21, 0.26; diameter
of F2: 0.23.
Paratype female: Body length: 8.2; maximum head
width: 3.0; fore wing length (including tegula): 8.7.
Colour. Similar to male, except for yellow stripe on
paraocular area with uniform width and upper
portion obtuse; disc of frons completely yellow
(Fig. 43). Mesoscutum with two yellow stripes on disc
and lateral margins; lateral portions of mesoscutum
orange yellow. Pubescence. Similar to male; except for
absence of long hairs on mandible margin, coxae,
trochanters, and mesepisternum ventrally; T4 with
marginal band of dark brown hairs throughout its
margin; T5–T6 with marginal hair bands occupying
whole margins. Integument sculpture. Similar to
male, except for clypeus with dense coarse punctures
(0.5–1 pd) and mesoscutum with dense minute punc-
tures (< 1 pd); metapostnotum with dense fine punc-
tures (1 pd). Structure. Lateral portions of lamella of
pronotal collar weakly obtuse in dorsal view, but
conspicuously acute on most of its length. Disc of
scutellum convex. Head about 1.3¥ broader than long
(3.05 : 2.35); ratio of lower interocular distance to
upper interocular distance: 0.84 (1.37 : 1.62); clypeus
about 1.9¥ broader than long (1.30 : 0.67); scape:
length 0.8; maximum width 0.25; length of F1–F3:
0.25, 0.14, 0.23; diameter of F2: 0.25.
Variation: The specimen from Nova Mamoré
(Rondônia) is smaller than the other specimens
(length: 7.2; maximum head width: 2.5).
Etymology: The species name refers to its yellow
frons.
Distribution: Brazil: Amazonas, Mato Grosso,
Rondônia, Pará (Fig. 73).
Type material: Holotype male, ‘DZUP\ 023650’
‘Vilhena, RO\ 9/X/1986\ C. Elias, leg.\ POLONO-
ROESTE’. Paratypes: BRAZIL: Amazonas: 2 females
(MZUSP), ‘Brazil, Amazonas,\ Novo Airão,
02°38,439′S\ 060°50.407′S, 12.xii.2007,\ R.
Gonçalves leg.’; Mato Grosso, 1 female (MZSP),
© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
399
‘Utiariti\ Rio Papagaio, Mt\.X.1966\ Lenko &
Pereira’; Pará: 1 male (MZSP), ‘PA: BR-14, km 93\
Belém – Brasília\ Ago – out, 1959\ Exp. Dep. Zool.’;
Rondônia: 1 male, ‘DZUP\ 022966’ ‘Vilhena, RO\
29/X/1986\ C. Elias, leg.\ POLONOROESTE’, 3
males and 4 females (DZUP), idem except ‘022966’,
‘27/11/1986’ ‘021837’, ‘27/11/1986’ ‘023658’, ‘022967’,
‘022971’ ‘13/XI/1986’, ‘022970’ ‘4/XI/1986’, ‘022974’
‘4/XI/1986’; 1 male, ‘DZUP\ 026619’ ‘VILHENA
Rondônia\ Brazil XI 1960\ M. Alvarenga leg.’; 1 male
(INPA), ‘BRAZIL, Rondônia, Nova\ Mamoré, Parque
Estadual de\ Guajará-Mirim, Rio Formoso’ ‘101926S-
643388W, 20.\ 27.x.1995. J. Vidal & L.S.\ Aquino,
Arm. de Malaise’ ‘0019729’.
THE FLAVIPENNIS SPECIES GROUP
The species of the flavipennis group have the
lamella of pronotal collar with lateral portions low
and obtuse (Figs 18, 20); epistomal suture straight
above upper margin of clypeus (character 5:1);
female mandibles with a vestigial second preapical
tooth; male hind basitarsus without tooth on ante-
rior margin; row of setae on margin of male S2 with
two short rows of erect setae on mid portion (char-
acters 45:1, 46:1; Figs 28, 29); row of setae on
margin of female S2–S3 forming a ‘V’-shaped line on
mid portion (Fig. 31); female metapostnotum with
dense fine punctures, intermingled amongst coarser
punctures on disc (Fig. 23); male S7 with apical por-
tions almost at right angles laterally; gonostylus
with same width throughout length, without lamel-
late expansion on basal portion ventrally (character
52:0).
Paratetrapedia flavipennis and P. duckei differ
from the remaining species in the group mainly by
male S3 not being deeply concave in ‘U’ shape
(Fig. 29), and lamella of pronotal collar with lateral
portions more obtuse and lower than in the remain-
ing species (Fig. 20). Paratetrapedia flaveola, P.
camargoi, and P. chocoensis are distinct from P.
atlantica, P. rufa, P. albilabris, P. andina, and P.
albopilosa by the bicoloured integument (dark
brown and yellow), by male T5 with band of dense
plumose hairs extending throughout its entire
margin, yellow infumate wing membrane (except in
P. flaveola, whose membrane of forewing is mostly
hyaline to whitish infumate, with radial cells yellow
infumate). The species in the clade containing P.
chocoensis at the basal position are mainly allopat-
ric and distributed over most of the Neotropical
region. Paratetrapedia flavipennis has a very
restricted distribution in central Amazonia, whereas
P. flaveola and P. duckei are widespread in the
Amazon basin.
400 A. J. C. AGUIAR and G. A. R. MELO
PARATETRAPEDIA DUCKEI (FRIESE, 1910)
(FIGS 20, 29, 55, 56, 74, 114, 141, 185, 186)
Tetrapedia duckei Friese, 1910: 62; lectotype male,
designated by Aguiar (2007), Brazil: Pará, Ilha do
Marajó (ZMB, examined).
Chalepogenus xanthaspis Cockerell, 1929: 442; holo-
type male, Guyana: Amatuk, Rio Potaro river
(AMNH, examined).
Tetrapedia duckei nomen nudum; Ducke, 1901:
55; Schrottky, 1902: 537. Ducke, 1910b: 364. Friese,
1923: 3.
Paratetrapedia xanthaspis; Michener, 1954: 116 (misi-
dentification of P. chocoensis).
Paratetrapedia xanthaspis Cockerell [sic]; Steiner,
1985: 1540 [misidentification based on geographical
distribution; floral record: Spachea membranacea
(Malpighiaceae)].
Paratetrapedia duckei; Moure, 1941: 518 (misidenti-
fication based on geographical distribution). Vogel,
1974: 207. Rebêlo et al., 2003: 273 [misidentification
of Tropidopedia punctifrons (Smith, 1854)]. Aguiar,
2007: 622.
Paratetrapedia (Tropidopedia) duckei; Michener &
Moure, 1957: 413. Albuquerque & Mendonça, 1996:
49 (misidentification of Tropidopedia punctifrons).
Paratetrapedia (Paratetrapedia) duckei; Rasmussen
& Ascher, 2008: 45.
Comments and diagnosis: Paratetrapedia duckei is
most clearly distinguished by lamella of pronotal
collar with lateral portions low and obtuse in dorsal
view (Fig. 20), conspicuously acute only on mid third,
and male S3 not deeply concave in ‘U’ shape (Fig. 29).
The male has the same pattern of pubescence on
terga and sterna found in P. flavipennis, with two
small rows of erect setae on mid portion of S2, and S3
mostly glabrous with only a row of simple setae
slightly interrupted in mid portion (Figs 28, 29). This
species is very similar structurally to P. flavipennis,
differing only in the pattern of colour on mesoscutum,
wings, and genal area, and smaller body size. Paratet-
rapedia duckei has the disc of mesoscutum dark
brown and lateral portions orange yellow, and two
thin yellow stripes on disc and lateral margins
(Figs 55, 56); genal area dark brown with thin yellow
stripe. Paratetrapedia duckei exhibits considerable
variation in colour pattern with dark forms in the
western portion of its distribution and pale forms in
the eastern portion.
Redescription: Male (redescription of holotype of
Chalepogenus xanthaspis Cockerell). Body length: 8.7;
maximum head width: 2.6; fore wing length (includ-
ing tegula): 8.7. Colour. Integument bicoloured (brown
and yellow), mostly yellow; paraocular area with wide
© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
yellow stripe obtuse on upper portion; frons, upper
paraocular area and vertex with dark brown integu-
ment; frons with yellow stripe on mid line (width
about 0.5–1.25 ¥ F2 diameter); genal area dark brown
with thin yellow stripe along ocular margin (width
about 0.6 ¥ F2 diameter); scape yellow; pedicel and
flagellomeres orange yellow. Mesoscutum with two
thin yellow stripes on disc and lateral margins,
slightly enlarged on mid portion (Fig. 56); mesoscu-
tum with disc reddish brown and lateral portions
orange yellow; axilla brown; wing membrane yellow
infumate with microtrichia yellow; fore wing dark
brown on apical third because of dense brown microt-
richia; veins and pterostigma orange yellow; hind
basitarsus weakly dark brown; tibial spurs pale
yellow. Terga orange yellow, with marginal zone dark
brown. Pubescence. Mostly yellow, except marginal
hair bands of T2–T4 and hind basitarsus with pubes-
cence dark brown. Scutellum with long erect simple
setae about 0.12–0.19 mm in length; metapostnotum
with short plumose hairs, c. 0.12 mm in length. T4
with marginal hair band almost complete on whole
margin, shortly interrupted on mid portion; T5–T6
with marginal band of plumose pale yellow hairs
throughout its margin; S2 with two short rows of
setae on mid portion; S3 with row of short simple
setae, slightly interrupted on mid portion; S6 with
two rows of stout plumose setae on margins laterally
and short plumose hairs on margin of apex. Integu-
ment sculpture. Clypeus with sparse coarse punc-
tures (0.5–2 pd), intermingled amongst sparse minute
punctures (2 pd); supraclypeal area with disc mostly
smooth and lateral portions with dense coarse punc-
tures (0.5–1 pd); disc of frons mostly with sparse
coarse punctures (< 0.5–2 pd); antennal scrobe with
fine dense punctures (0.5 pd). Mesoscutum with fine
dense punctures (< 0.5 pd) intermingled amongst
sparse coarser punctures (2 pd); metapostnotum with
dense minute punctures on lateral portions (1 pd),
and coarser punctures on disc (0.5–1 pd); transition
zone between metapostnotum and propodeum with
smooth and shiny surface; mesepisternum mostly
with sparse coarse punctures (2 pd). Structure. Prono-
tal collar with lamella obtuse and low on third lateral
portions, and central third of lamella acute in dorsal
view. Scutellum disc convex. Head about 1.15¥
broader than long (2.6 : 2.3); ratio of lower interocular
distance to upper interocular distance: 0.9 (1.2 : 1.4);
clypeus about 1.5¥ broader than long (1.16 : 0.73);
scape: length 0.72, maximum width 0.21; length of
F1–F3: 0.7: 0.23: 0.16: 0.23; diameter of F2: 0.26.
Variation: Besides the variation in body colour men-
tioned above, specimens from the western portion of
the distribution (Brazil: Rondônia, Mato Grosso;
Peru) have pubescence of hind leg mostly dark brown
 SYSTEMATICS OF THE GENUS PARATETRAPEDIA
Figure 74. Geographical distribution of Paratetrapedia flavipennis and Paratetrapedia duckei.
and yellow stripe on genal area interrupted on mid
third, whereas most specimens from the northern and
eastern portions (Brazil: Amapá, Amazonas, Manaus,
Pará; Guyana, French Guiana) have pubescence of
hind leg orange yellow and yellow stripe on genal
area entire along eye margin. In addition, some
females from Amapá and Pará have microsculpture of
minute punctures on T1, and one specimen from
Rondônia has this microsculpture on T2–T3.
Distribution: Bolivia; Brazil: Acre, Amapá, Amazonas,
Mato Grosso, Pará, Rondônia; French Guiana;
Guyana; Peru; Suriname (Fig. 74).
Type material: Tetrapedia duckei Friese; lectotype
female (ZMB): ‘I. Marajó\ R. Anajás\ 10.6.1900\
Ducke’ ‘Tetrapedia\ duckei\ 1900 Friese det.’.
Chalepogenus xanthaspis Cockerell, holotype male
(AMNH), ‘Amatuk\ Brit. Guiana\ May 24.1929\ J.
Ogilvie’ ‘Chalepogenus\ xanthaspis\ Ckll Type’ ‘Ac.
33337’ ‘Lectotype Chalepogenus 씹 xanthaspis Cock-
erell 1929’.
Additional material: See Appendix.
PARATETRAPEDIA FLAVIPENNIS (SMITH, 1879)
(FIGS 54, 74, 115, 140, 181, 182)
Tetrapedia flavipennis Smith, 1879: 129; holotype
female, Brazil: Amazonas, Tefé (BMNH, examined).
© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
401
Tetrapedia flavipennis; Dalla Torre, 1896: 299. Friese,
1899: 300. Cockerell, 1909: 399.
Chalepogenus flavipennis; Cockerell, 1929: 443.
Paratetrapedia flavipennis; Moure, 1941: 518. Moure,
1944a: 74. Moure, 1996: 929. Aguiar, 2007: 623.
Paratetrapedia (Paratetrapedia) flavipennis; Miche-
ner & Moure, 1957: 416. Silveira et al., 2002:
136.
Comments and diagnosis: Paratetrapedia flavipennis
differs from the remaining species in the group
mainly by male S3 not deeply concave in ‘U’ shape
(Fig. 29), and lamella of pronotal collar with lateral
portions more obtuse and lower than in the remaining
species (Fig. 20). Paratetrapedia flavipennis is struc-
turally identical to P. duckei, but differs mainly by
mesoscutum and genal area completely orange yellow
and larger body size (> 9.0 mm in length).
Redescription: Holotype (female). Body length: 9.8;
maximum head width: 3.0; fore wing length (includ-
ing tegula): 8.7. Colour. Integument mostly orange
yellow. Paraocular area with wide yellow stripe with
upper portion obtuse; mandibles, labrum, and clypeus
yellow; frons, upper paraocular area, and vertex
reddish brown; frons with thin yellow stripe on mid
line; scape yellow, pedicel and flagellomeres orange
yellow; genal area completely orange yellow. Mesos-
cutum completely orange yellow, with two thin yellow
stripes weakly faint on disc. Wings with membrane
402 A. J. C. AGUIAR and G. A. R. MELO
yellow infumate; apical portion of fore wing mem-
brane dark because of dark brown microtrichia; veins
and pterostigma orange yellow. Tibial spurs pale
white. Margins of terga weakly dark brown. Pubes-
cence. Mostly orange yellow. Scutellum with long
simple erect setae, about 0.12–0.26 mm in length;
metanotum with short plumose hairs, about 0.10 mm
in length; metapostnotum with dense short plumose
hairs, c. 0.08 mm in length. T4–T6 with marginal
band of short simple hairs throughout margin. Row of
setae on sternal margins sinuated in ‘V’ shape on mid
portion (as in Fig. 32). Integument sculpture. Clypeus
and supraclypeal area with dense coarse punctures
(< 0.5 pd); disc of frons with dense coarse punctures
on mid portion (< 0.5–2 pd). Mesoscutum with dense
fine punctures (< 0.5 pd), intermingled amongst
sparse coarser punctures (1–3 pd); lateral portions of
metapostnotum with dense fine punctures (0.5–1 pd),
with few coarser punctures on disc. Mesepisternum
laterally with sparse coarse punctures (2 pd), omaular
area with denser punctures (0.5–1 pd). Structure.
Lamella of pronotal collar obtuse throughout its
length. Scutellum disc almost flat. Head about 1.2¥
broader than long (3.0 : 2.5); ratio of lower to upper
interocular distance: 0.87 (1.4 : 1.6); clypeus about
1.6¥ broader than long (1.27 : 0.77); scape: length, 0.8,
maximum width, 0.2; length of F1–F3: 0.26, 0.17,
0.17; F2 diameter: 0.2.
Variation: The holotype of P. flavipennis has fore wing
membrane yellow infumate with one thin band of
dark brown microtrichia on posterior margin; width of
band of microtrichia about 1.0 ¥ F2 diameter. Only
one specimen from Tefé (Amazonas) has the same
pattern of microtrichia on fore wing of the holotype.
The remaining specimens examined have a broader
band of microtrichia dark brown on apical portion of
fore wing (c. 3 ¥ F2 diameter).
Distribution: Brazil: Amazonas; Peru (Fig. 74).
Type material: Tetrapedia flavipennis Smith; holotype
female (BMNH), ‘Type\ H.T.’ ‘B.M. Type\ Hym.\
17.B.884\ Tetrapedia\ flavipennis\ (Type) Sm.’ ‘58\
h\\ Ega’.
Additional material: See Appendix.
PARATETRAPEDIA FLAVEOLA SP. NOV.
(FIGS 18, 23, 28, 31, 47, 75, 78, 105, 136, 191, 192)
Paratetrapedia lineata; Melo & Zanella (2003): 2919
(misidentification).
© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
Comments and diagnosis: Paratetrapedia flaveola is
superficially very similar to P. lineata in body size,
structure, and colour pattern. However, they belong to
different species groups and are quite distinct in the
diagnostic characteristics for these groups. Paratetra-
pedia flaveola differs from the species of the lineata
group by the narrowed upper portion of the yellow
stripe on the female paraocular area(Fig. 47); genal
area mostly reddish brown with narrow yellow stripe
along eye margin (about 0.8 ¥ F2 diameter), usually
interrupted on mid third in females; metapostnotum
completely black, with dense minute punctures on
lateral portions and coarser punctures on disc;
lamella of pronotal collar with lateral portions obtuse
in dorsal view; male S2 with two small rows of setae
on mid portion. Paratetrapedia flaveola differs from P.
atlantica, P. rufa, P. albilabris, P. andina, and P.
albopilosa by the bicoloured integument (dark brown
and yellow), and by male T5 with band of dense
plumose hairs extending throughout its entire
margin. Paratetrapedia flaveola differs from P. cama-
rgoi and P. chocoensis mainly by hyaline wing mem-
brane and dark brown disc of terga.
The specimens of P. flaveola collected by Helmut
Heider in Vila Tunari (Cochabamba, Bolivia) repre-
sent the first records of Paratetrapedia on orchid oil
flowers of the genera Ornithophora amazonica
(labelled as Stigmatostalix) and Oncidium.
Description
Holotype male: Body length: 8.5; maximum head
width: 2.4; fore wing length (including tegula): 6.75.
Colour. Integument bicoloured (reddish brown and
yellow), mainly yellow. Mandible yellow with apex
black; labrum yellow; clypeus mostly yellow with
upper lateral portions dark brown; supraclypeal
area mostly yellow, with lower margin dark brown;
yellow stripe on paraocular area acute on upper
portion; frons reddish brown with one thin yellow
stripe (Fig. 47); genal area reddish brown with thin
yellow stripe along ocular margin, brown with thin
yellow stripe along ocular margin. 0.8 ¥ F2 diam-
eter; scape yellow; pedicel and flagellomeres with
anterior surface orange yellow and posterior surface
reddish brown. Mesoscutum with two thin yellow
stripes on disc and lateral margins; scutellum and
metanotum yellow; axilla brown; lateral areas of
metanotum dark brown; metapostnotum completely
reddish brown; propodeum mostly brown with small
yellow marks slightly faint on upper and lower
margins; mesepisternum laterally with upper
portion yellow and lower portion reddish brown.
Tergal disc yellow, with anterior and posterior
margins reddish brown; sterna yellow. Wing mem-
brane mostly white infumate with white microtri-
chia. Fore wing membrane slightly yellow infumate
 SYSTEMATICS OF THE GENUS PARATETRAPEDIA
on proximal third, with some yellow microtrichia;
posterior margin with some dark brown microtri-
chia; veins and pterostigma orange yellow. Mid and
hind basitarsus dark brown; tibial spurs white.
Pubescence. Mostly pale yellow, except marginal
bands of T3–T5 and basitarsus of mid and hind legs
with pubescence dark brown. Inner face of scape
with pale yellow setae, about 0.12–0.26 mm in
length. Mesoscutum and scutellum with dense
velvet plumose pale yellow pubescence; scutellum
with long erect simple setae, about 0.10–0.17 mm in
length; metapostnotum with yellow plumose hairs,
about 0.17 mm in length. T4 with marginal hair
band of brown plumose hairs almost complete, with
two small gaps on lateral portions; T5–T6 with mar-
ginal band of long plumose hairs occupying whole
margin; on T5, with dark brown hairs; on T6, pale
yellow; margin of S2 with two short rows of stout
setae on middle portion (Fig. 28); posterior margin
of S3 of male deeply concave in ‘U’ shape, covered
by dense short plumose pubescence; S4 and S5
typical of genus; S6 with two rows of stout plumose
hairs on lateral margins, and short plumose hairs
on margin of apex. Integument sculpture. Clypeus
with dense coarse punctures (1 pd), intermingled
amongst sparse fine punctures (1–2 pd); supracly-
peal area with dense coarse punctures (0.5–2 pd);
frons with dense coarse punctures (0.5–2 pd); anten-
nal scrobe with dense fine punctures (< 0.5 pd).
Mesoscutum and scutellum with dense fine punc-
tures (< 0.5 pd), intermingled amongst coarser and
sparser punctures (1–3 pd); metapostnotum with
dense fine punctures on lateral portions (0.5 pd),
and coarser punctures on disc (0.5–1 pd); mesepis-
ternum laterally with dense coarse punctures
(< 1 pd). Structure. Lamella of pronotal collar
weakly obtuse on lateral portions, with acute carina
distinctly on mid portion. Disc of scutellum weakly
convex, almost flat. Head about 1.3¥ broader than
long (2.6 : 2.0); ratio of lower interocular distance to
upper interocular distance: 0.71 (1.0 : 1.4); clypeus
about 1.7¥ broader than long (1.03 : 0.58); scape,
length 0.6; maximum width 0.2; length of F1–F3:
0.17, 0.14, 0.21; diameter of F2: 0.2.
Paratype female: Body length: 7.7; maximum head
width: 2.4; fore wing length (including tegula): 6.2.
Colour. Similar to male, except for yellow stripe on
mid line of frons broader (about 0.5–2.0 ¥ F2 diam-
eter). Pubescence. Similar to male, except for mar-
ginal band on T5–T6 with reddish brown simple
hairs; S2–S4 with rows of setae on margin, sinuated
in ‘V’ on mid portion (Fig. 31). Integument sculp-
ture. Similar to male, except for denser punctures
on clypeus and supraclypeal area; clypeus and
supraclypeal area with dense coarse punctures
© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
403
(< 0.5 pd); frons mostly with dense fine punctures
(< 0.5 pd), with few coarser punctures adjacent to
mid line. Mesoscutum and scutellum with dense fine
punctures (0.5 pd), intermingled amongst sparser
coarser punctures; mesepisternum laterally with
dense coarse punctures (0.5–1 pd). Structure.
Similar to male, except for pronotal collar with
lamella more obtuse laterally. Disc of scutellum
weakly convex, almost flat. Head about 1.3¥ broader
than long (1.85 : 2.4); ratio of lower interocular dis-
tance to upper interocular distance: 0.85
(1.25 : 1.46); clypeus about 1.9¥ broader than long
(1.21 : 0.62); scape: length 0.7, maximum width 0.19;
length of F1–F3: 0.22, 0.12, 0.17; diameter of F2:
0.21.
Variation: The females have the yellow stripe on
genal area complete or interrupted on mid third, but
it is always very narrow (c. 0.8 ¥ F2 diameter). The
yellow stripe on paraocular area in a few specimens
has an obtuse apical portion. The pubescence of mid
and hind leg can vary between pale yellow to com-
pletely dark brown.
Etymology: The species name refers to the yellow
pattern of colour of the integument.
Distribution: Bolivia; Brazil: Acre, Amapá, Amazonas,
Bahia, Distrito Federal, Goiás, Maranhão, Mato
Grosso, Mato Grosso do Sul, Minas Gerais, Pará,
Rondônia, Roraima, São Paulo, Tocantins; Ecuador;
Guyana; Peru; Suriname (Fig. 75).
Type material: Holotype male, ‘DZUP\ 022427’
‘Brazil, São Paulo, Cajuru,\ Fazenda Rio Grande,\
21°12′S 47°09′W,\ 25.iii.2000, G.A.R. Melo,\
Stachytarpheta’. Paratypes: BOLIVIA: 1 female
(ZSMC), ‘Bolivien, Chapare\ Villa Tunari, 320 m\
20.November 2002\ leg. H.Heider’ ‘Stigmatostalix\
amazonica’ ‘Monoeca’; 1 female (ZSMC), ‘GG-1950
Bolivien 11/00\ Cochabamba/Vila Tunari\ 320 m\
leg. H. Heider\ Köder: Oncidium’ ‘Monoeca’; 1 female
(ZSMC), ‘GG-1953 Bolivien?/00\ Cochabamba/Vila
Tunari\ 320 m\ leg. H. Heider\ Köder: Stigma-
tostalix amazonica’ ‘Monoeca’; 1 female (ZSMC), ‘Boli-
vien,\ Chapare, Vila Tunari,\ 320 m, 14.3.2001\ leg.
H. Heider’ ‘Paratetra-\ pedia?’; 3 females (INHS),
‘Bolivia La Paz Prov. Mapiri\ Arroyo Tuhiri,
10.IV.2004\ 15°17′26′S 68°15′46′W\ 508 m Leg M.
Hauser’, 1 female (INHS), idem except ‘Paratetrape-
dia\ spp.\ det. Claus Rasmussen2003’; BRAZIL:
Acre: 1 female (RPSP),’Brazil Acre\ Rio Branco\ Res.
Humaitá\ 29/X/1994\ M.L. Oliveira’ ‘no\ 0111’; 1
male (RPSP), ‘Brazil Acre\ Rio Branco\ 30/X/1994\
M.L.Oliveira’; 1 male, ‘DZUP\ 021867’ ‘Cruzeiro do
Sul\ Acre-Brazil II-63\ M. Alvarenga’, 2 males
404 A. J. C. AGUIAR and G. A. R. MELO
Figure 75. Geographical distribution of Paratetrapedia flaveola sp. nov., Paratetrapedia camargoi sp. nov., and
Paratetrapedia chocoensis sp. nov.
(DZUP), idem except ‘021869’, ‘021879’; 1 male,
‘DZUP\ 21865’ ‘R. Branco Acre\ BR 15-20-XI-61\
F.M. Oliveira’, 4 males and 1 female (DZUP), idem
except ‘021866’, ‘021903’, ‘022404’, ‘21901’, ‘021907’; 1
female, ‘DZUP\ 022707’ ‘Rio Branco-AC\ Brazil 8-XI-
61\ F.M. Oliveira leg’; 1 female (INPA), ‘BRAZIL.
Acre\ Rio Branco\ 09°58′S, 67°48′W\ 28/abril/
2001’Em lanterneira\ (Lophanthera\ lactescens)\
Oliveira et al. leg.’; Amapá: 2 females (IEPA), ‘Brazil-
Amapá\ I.E.P.A.’ ‘Brazil-Amapá\ Laranjal do Jarí\
Ponte 1\ 00:32:54S/ 52:11:30W\ 14/IX/2001\ G.A.R.
Melo’, 1 female (IEPA), idem except ‘L. do Jari\ Rio
Cajari\ (ramal do escondido)\ 00:33:27S/ 52: 14:36W\
20/IX/2001\ G.A.R. Melo’; 1 female, ‘DZUP\ 021844’
‘Serra do Navio-AP\ Brazil 7-II-62\ F. M. Oliveira’;
Amazonas: 1 female, ‘DZUP/ 022716’ ‘TEFE Amazo-
nas\ Brazil XII-61\ F.M. Oliveira’, 4 females (DZUP),
idem except ‘022706’, ‘022702’, ‘022703’, ‘022711’; 1
female, ‘DZUP\ 022710’ ‘TEFE Amazonas\ Brazil
1-4-XII-61\ F.M. Oliveira’; 1 female, ‘DZUP\ 022735’
‘TEFE Amazonas\ BR-I-1962\ E. Carvalho’; Bahia: 1
female (RPSP), ‘951604’, Santo Antônio\ Cocos-BA,
Brazil\ 44°40′W,14°8′S′, ‘24, 27-VII-1995\ Mazucato
leg.’; Distrito Federal: 1 male (DZUP), ‘D. Federal\
F.C. Grosso\ 16.10.77’ ‘Paratetrapedia\ lineolata?’; 1
female (DZUP), ‘Brazilia\ Lago Sul\ 27.8.1977\ coll.
A. Raw\ MUDB’ ‘Peixotoa\ Malpighiaceae’ ‘Paratet-
rapedia\ lineolata’; 1 female, ‘DZUP\ 041089’ ‘Brazil
B. Mata\ Loc. J. Botânico\ Data 05-11-96\ Col. J.
© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
Valdecy’ ‘1439’, 1 female (DZUP), idem except ‘041091’
‘19-02-97’ ‘1945’; 1 female, ‘DZUP\ 041090’, ‘Brazil\
Loc. J. Botânico\ Data 19.02.97\ Col. Ana Alves’
‘1944’; 1 male, ‘DZUP\ 041092’ ‘Brazil Ipê\ amarelo\
Loc. B. Mata\ Data 19/9/95\ Col. Dimaren’ ‘193’; 1
male, ‘DZUP\ 041093’ ‘Brazil Cerrado\ Loc. J.
Botânico\ Data 28-05-96\ Col. Ma Araujo’ ‘852’;
Goiás: 2 females (RPSP), ‘Aragarças\ GO. Brazil\
10.24.I.1971\ col. Camargo’; 1 female, ‘DZUP\
022722’ ‘S. Rita Araguaia\ GO Brazil XII-63\ M.
Alvarenga, leg’; 1 female (DZUP), ‘Brazil: GO\ Goiás,
Cor\ Paciência\ A. Raw col\ 7.7.1984’; 1 female
(DZUP), ‘Brazil: GO\ Goiás\ Gold Creek\ A. Raw
col\ 8.7.1984’; 3 females (DBAI), ‘Goiás Velho.\ Corr.
Paciência\ 20.11.1981\ in flight’ ‘Paratetrapedia\
lineata’; 1 male (DBAI), ‘Goiás Velho.\ Corr. Paciên-
cia\ 20.11.1981’ ‘Paratetrapedia\ lineolata’; 1 female
(DBAI), ‘Goiás Velho.\ Corr. Paciência\ 20.11.1981’
‘Desmodium’ ‘Paratetrapedia\ lineata’; 2 males and 2
females (DZUP), ‘Goiás Velho\ C. Paciência\ 31-07-
82’; 1 male (DZUP), ‘Goiás Velho\ C. Paciência\
31.0783’; 1 female (DBAI), ‘BRAZIL\ Goiás Velho\
A.Raw 5.4.1980\ 7 km woow’ ‘Paratetrapedia\ lin-
eolata’; 1 female (DZUP), ‘Brazil: GO\ Ch. Veadeiros\
05-07.12.1996\ Boaventura,\ Freitas & Freitas’
‘Paratetrapedia\ cf. lineolata’; 6 males and 1 female
(DZUP), ‘Brazil: GO\ Goiás, Cor\ Paciência\ A. Raw
col\ 15.10.1984’ ‘Psychotria’; 1 male (DZUP),
‘BRAZIL\ Goiás Velho\ A.Raw 3.2.1980’ ‘pink
 SYSTEMATICS OF THE GENUS PARATETRAPEDIA
Rubiac?’ ‘sp23’; 3 males (DZUP), ‘Brazil: GO\ Goiás,
Cor\ Paciência\ A. Raw col\ 14.10.1984’, 3 males
(DZUP), idem except ‘15.10.1984’, 4 males (DZUP),
idem except ‘13.10.1984’; 5 males (DBAI), ‘Goiás
Velho.\ Corr. Paciência\ 18.11.1981’ ‘Stachytarpheta\
jamaicensis\ Verbenaceae’ ‘Paratetrapedia\ lineata’;
2 males (DZUP), ‘BRAZIL\ Goiás Velho\ A.Raw
7.12.1980\ Corr. da Paciência’ ‘Stachytarpheta\
jamaicensis\ Verbenaceae’ ‘Paratetrapedia\ lineata’;
1 female (DZUP), ‘Brazil: GO\ Itiquira\ Byrsonima
laxiflora\ A.Raw 26.5.1996’; 1 female (DZUP), ‘Brazil:
GO\ Goiás, Cor\ Paciência\ A. Raw col\ 16.5.87’; 1
female (DZUP), ‘Brazil: GO\ Goiás, Cor\ Paciência\
A. Raw col\ 7.7.1984’; 1 female (DZUP), ‘Brazil\ Est.
Goias\ P. Bernardo\ A. Raw 2.10.77’ ‘Byrsonima’
‘Paratetrapedia\ lineata’; 1 female (DZUP), ‘Goiás\ C.
Paciência\ 31-7-82’; 1 male (DZUP), ‘Brazil-Goiás\ S.
José do Tocantins,\ Rio Maranhão,\ 22.VI.1942, F.
Lane’ ‘105251’; 1 male, ‘DZUP\ 021883’ ‘Brazil, Goiás,
11 km SE de\ Campos Belos, 13°07′32′S\ 46°44′29′
W, 650 m, 04.iv.2003,\ Melo, Aguiar, Marchi e\
Gonçalves, em cerradão sobre\ massapé, em Hyptis’,
3 males and 2 females (DZUP), idem except ‘021821’,
‘021825’, ‘021899’, ‘021898’, ‘021884’; 1 female,
‘DZUP\ 021824’ ‘Brazil, Goiás,\ 2 km W de
Teresina de Goiás,\ Fazenda Santa Tereza,\
13°47′43′S47°17′39′W, 800 m,\ 03.iv.2003, Melo,
Aguiar,\ Marchi e Gonçalves’, 2 females (DZUP),
idem except ‘021823’, ‘021860’; 1 male, ‘DZUP\
021893’ ‘Dianópolis-GO\ Brazil – 12-III-62\ F M
Oliveira leg’; Maranhão: 1 female (MPEG), ‘MA
Açailândia\ 29-IX-1978’ ‘Brazil MA\ MF Torres’;
Mato Grosso: 1 female, ‘DZUP\ 021886’ ‘Jaçanã-P.N.
Xingu\ MT Brazil XI-1961\ Alvarenga, Werner’, 1
female (DZUP), idem except ‘021845’; 1 male (RPSP),
‘Serra das Araras-MT\ Brazil 19,27-IX-1987’
‘SD21,57°14′W,15°27′S\ Gimenes leg. 871134’; 1 male
(DZUP), ‘Mato Grosso\ P.N. Araguaia\ Mata\ 21-6-
79’; 1 female (DZUP), ‘Mato Grosso\ Valle dos
Sonhos\ 24.4.1978\ coll. A.Raw’ ‘sp23’; 1 female
(DZUP), ‘Mato Grosso\ P.N. Araguaia\ I. Centopéia\
15-6-79’ ‘Coll. at. Cassia Caesalpinaceae’ ‘sp.23’; 1
female (MPEG), ‘Brazil MT\ Barra do Bugres\ R.E.
Serra das Araras\ 19.2.1986’ ‘Brazil Mato Grosso\
Marcio Zanuto’; 1 male, ‘DZUP\ 22453’ ‘Cáceres, MT\
27.III.1985\ C.Elias leg.\ POLONOROESTE’, 13
males and 19 females (DZUP), idem except ‘022490’,
‘022467’, ‘022473’, ‘022475’ to ‘022479’, ‘022483’ to
‘022487’, ‘022489’, ‘022452’, ‘021819’, ‘021813’ to
‘021815’, ‘022472’, ‘021804’ to ‘021810’, ‘022447’,
‘021799’ to ‘021802’; 1 female, ‘DZUP\ 022461’
‘Cáceres, MT\ 11.XII.1984\ C.Elias leg.\ POLONO-
ROESTE’; 2 males, ‘022465’ ‘Cáceres, MT\
12.II.1985\ C.Elias leg.\ POLONOROESTE’, 1 male
(DZUP), idem except ‘022457’; 1 male, ‘DZUP\
022460’ ‘Cáceres, MT\ 16.I.1985\ C.Elias leg.\
© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
405
POLONOROESTE’, 2 males and 3 females (DZUP),
idem except ‘022482’, ‘022491’, ‘022474’, ‘022480’,
‘022481’; 1 male, ‘DZUP\ 022430’ ‘Cáceres, MT\
21.XI.1984\ C.Elias leg.\ POLONOROESTE’, 2
males (DZUP), idem except ‘022431’, ‘022436’; 1
female, ‘DZUP\ 022469’ ‘Cáceres, MT\ 22.III.1985\
C.Elias leg.\ POLONOROESTE’; 1 female, ‘DZUP\
022450’ ‘Cáceres, MT\ 24.XII.1984\ C.Elias leg.\
POLONOROESTE’; 1 male, ‘DZUP\ 022468’ ‘Cáceres,
MT\ 26.III.1985\ C.Elias leg.\ POLONOROESTE’, 1
male and 2 females (DZUP), idem except ‘022458’,
‘022432’ and ‘022411’; 1 male, ‘DZUP\ 022397’
‘Cáceres, MT\ 27.XII.1984\ C.Elias leg.\ POLONO-
ROESTE’, 4 males (DZUP), idem except ‘022440’,
‘022434’, ‘022448’, ‘022446’; 1 male, ‘DZUP\ 022449’
‘Cáceres, MT\ 28.I.1985\ C.Elias leg.\ POLONO-
ROESTE’, 1 male and 2 females (DZUP), idem
except‘022429’, ‘022463’ and ‘022449’; 1 male, ‘DZUP\
022439’ ‘Cáceres, MT\ 5.XII.1985\ C.Elias leg.\
POLONOROESTE’, 3 males and 1 female (DZUP),
idem except ‘041109’ ‘1984’, ‘022442’, ‘022437’ and
‘022462’; 1 male, ‘DZUP\ 022435’ ‘Cáceres, MT\
9.I.1985\ C.Elias leg.\ POLONOROESTE’, 4 males
and 1 female (DZUP), idem except ‘022438’, ‘022443’
to ‘022445’, ‘022454’; 1 male, ‘DZUP\ 022441’
‘Cáceres, MT\ 9-11.XI.1984\ C.Elias leg.\ POLONO-
ROESTE’, 1 male (DZUP), idem exceopt ‘022433’; 1
male, ‘DZUP\ 22470’ ‘Cáceres, MT\ 7.II.1985\
C.Elias leg.\ POLONOROESTE’, 2 males and 4
females (DZUP), idem except ‘022464’, ‘022456’,
‘022410’, ‘022459’, ‘022451’, ‘022455’; 1 male, ‘DZUP\
021862’ ‘Chap. Guimarães-MT\ (C.A. Buriti)\
08.II.1986\ Pe. Moure & Gorayeb’, 9 males and 2
females (DZUP), idem except ‘021863’, ‘021864’,
‘021885’, ‘021890’, ‘021895’, ‘021896’, ‘021900’,
‘021902’, ‘022396’, ‘022419’, ‘022422’; 1 female,
‘DZUP\ 021831’ ‘Cuiabá MT\ Brazil XI-1962\ M.
Alvarenga’; 1 male, ‘DZUP\ 022406’ ‘Utiariti (325 m)\
Rio Papagaio, MT\ Brazil VII-VIII.961\ K. Lenko
col.’; Mato Grosso do Sul: 1 male, ‘DZUP\ 041106’
‘Brazil, Três Lagoas,\ MS, 18-20.iv.2003\ AJCAguiar
leg’; Minas Gerais: 1 male, ‘DZUP\ 021892’ ‘Cabeceira
do Corr.\ Leitão, Curvelo, MG\ 15-VI-1972, 700 m\
Mielke & Brown leg’; 1 male, ‘DZUP\ 022402’ ‘Passos
MG\ Brazil V 1961\ C. Elias leg.’, 1 female (DZUP),
idem except ‘021842’; 1 female, ‘DZUP\ 021826’
‘Ibiraci-MG\ Brazil-X-61\ C. Elias leg.’; 1 female,
‘DZUP\ 021888’ ‘Brazil Minas Gerais,\ Corinto,
1-15.viii.\ 1979, C. Elias leg.’; 1 female, ‘DZUP\
021847’ ‘Nova Resende MG\ Brazil – VII-1961\ Clau-
dionor Elias’; Pará: 1 female (SEMC), ‘Brazil Pará\
Conceição do Ara-\ guaia July 1959\ (M. Alvarenga)’;
1 female (MPEG), ‘Belem Mocambo\ 05.I.1978’ ‘Brazil
Para\ MF Torres’; 1 female (RPSP), ‘Gorotire.
PA\:Gradaus Brazil\ 20.VIII.-5.IX-1983\ 830847’
‘SB-22,51-Sa\ Camargo leg.’; 1 male (RPSP), ‘Capa-
406 A. J. C. AGUIAR and G. A. R. MELO
nema, PA\ Brazil-15-II-84\ SA-23,47 II-1c\ 840133’
‘Camargo\ Mazucato’; 1 male (RPSP), ‘Capanema,
PA\ Brazil-15-II-84\ SA-23,47 II-1c\ 840132’ ‘Cama-
rgo\ Mazucato’; 1 female (MPEG), ‘Brazil Pará
Belém\ Floresta APEG\ 19a22.XI.1982\ Col. I.S.Go-
rayeb e Equipe’ ‘Armadilha\ suspensa\ 1.6 m’; 1
female (MPEG), ‘Belém Brasília\ km 90 FS Antonio\
30-VII-1972’ ‘Brazil, PA\ M. Helena Col.’; 1 female
(MPEG), ‘Mun. Benevides\ PA-408 km 06\ 5-V-1982’
‘Brazil Pará\ WL Overal’; 1 female (MPEG) ‘Brazil
Pará\ Paragominas\ Faz. Cachoeira\ do Rio Ver-
melho\ 15a18.I.1991’; 1 female (INPA), ‘14 xi-1986\
J.A. Rafael\ Arm. Malaise’ ‘BRAZIL Para\ Benev-
ides\ F.Morelandia’; 1 female, ‘DZUP\ 021841’ ‘Ford-
landia,\ Est. Pará\ R. Damasceno\ -col.\ I.1956’; 1
female, ‘DZUP\ 021843’ ‘Belem\ Pará BRAZIL\
X-1954 O.Rego’; 1 female, ‘DZUP\ 021827’ ‘Aurá\
Pará Brazil\ 16.VI.1956\ E. Lobato’; Rondônia: 1
female, ‘DZUP\ 021838’ ‘Brazil, Rondônia,\
Cacaulândia, 9-19.xi.1994\ O.H.H. Mielke’; 1 female
(RPSP), ‘Brazil\ Rondônia\ Costa Marques’ ‘3509-21/
11/96\ S12°17.653′\ W64°01.804′\ Brown, Boina,
Vieira’; 1 female, ‘DZUP\ 021830’ ‘Fte. P. da Beira\
RO-Brazil 7-XI-61\ F.M. Oliveira’; 1 female (RPSP),
‘Brazil, Rondônia\ Guajara Mirim\ Sa. Pacaas Novos’
‘22/agosto/1995\ F.M. Dantas, col’ ‘PN0403’, 2 females
(RPSP), idem except ‘PN0440’, ‘Data 24/IV/1995\
M.L. Oliveira’ ‘PN0038’; 4 females (RPSP), ‘Brazil –
RO\ Guajará Mirim.\ Pacaás Novos.’ ‘Data 23/VII/
1995\ M.L. Oliveira’ ‘PN0316’, 3 females (RPSP),
idem except ‘PN0350’, ‘PN0315’, ‘PN0317’; 1 female
(RPSP), ‘Brazil – Rondônia\ Guajara Mirim\ Sa.
Pacaás Novos’ ‘08/setembro/1995\ F.M. Dantas, col.’
‘PN0543’; 1 male (RPSP), ‘Brazil, Rondônia\ Guajará
Mirim\ Sa. Pacaás Novos’ ‘20/janeiro/1996\ M.L.
Oliveira col’ ‘PN0844’; 3 males and 1 female (INPA),
‘Brazil, Rondônia\ Próx. Guajará Mirim\ 10°48′S,
65°22′W’ ‘12-14/X/2001\ Oliveira, Morato & Cunha
leg.’; 1 female (INPA), ‘Brazil, Rondônia\ Guajara
Mirim\ 10°48′S, 65°22′W’ ‘12-14/X/2001\ Oliveira,
Morato & Cunha leg.’; 1 female (RPSP), ‘Brazil\
Rondônia\ Mirante da Serra’ ‘2969-07/11/96\
S11°01.516′\ W62°48.750′\ Brown, Boina, Vieira’; 1
female (RPSP), ‘RO-13342\ Brazil.Rondônia\
Mirante da Serra\ Linha 81’ ‘19-Agosto-1997\
11°00′04.6′S-62°49′05.3′W\ Brown, Boina, Vieira\
no.’; 1 female (RPSP), ‘RO-13340\ Brazil.Rondônia\
Mirante da Serra\ Linha 81’ ‘19-Agosto-1997\
11°00′04.6′S-62°49′05.3′W\ Brown, Boina, Vieira\
no.’; 1 female (RPSP), ‘Brazil\ Rondônia\ Nova
Mamoré’ ‘367-12/09/96\ S10°19.440′\ W64°46.775′\
Brown, Boina, Vieira’; 1 female (RPSP), ‘Brazil\
Rondônia\ Nova Mamoré’ ‘440-12/09/96\
S10°19.440′\ W64°46.775′\ Brown, Boina, Vieira’; 1
female (RPSP), ‘Brazil\ Rondônia\ Nova Mamoré’
‘366-12/09/96\ S10°19.440′\ W64°46.775′\ Brown,
© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
Boina, Vieira’; 1 female, ‘DZUP\ 021856’ ‘Vilhena,
RO\ 12/XI/1986\ C.Elias, leg\ Polonoroeste’, 1 female
(DZUP), idem except ‘021851’; 1 female, ‘DZUP\
021857’ ‘Vilhena, RO\ 29/X/1986\ C.Elias, leg\ Polo-
noroeste’, 1 males and 2 females (DZUP), idem except
‘022407’, ‘021850’, ‘021858’; 1 female, ‘DZUP\ 021849’
‘Vilhena, RO\ 22/X/1986\ C.Elias, leg\ Polonoroeste’;
2 males (DZUP), idem except ‘021877’ ‘6/12/1986’,
‘021880’ ‘2/12/1986’; 1 female, ‘DZUP\ 021872’
‘Vilhena, RO\ 4/XI/1986\ C.Elias, leg\ Polonoroeste’,
3 females (DZUP), idem except ‘021853’, ‘021854’,
‘021873’; Roraima: 1 female (INPA), ‘Brazil, Roraima\
Parafuri, 031759N\ 640030W, 20.iv.1994’ ‘U.C.
Barbosa, extrato de\ cravo da Índia’; 1 female (INPA),
‘Brazil\ Roraima\ 23-à 28-II-2003’ ‘M.L.Oliveira\
S.J.R. Silva leg.\ Fazenda Trinta\ Br-210’; 1 female
(INPA), ‘Brazil\ Roraima\ 23-á 28-II-2003’ ‘M.L.
Oliveira &\ S.J.R. Silva leg.\ A.I. São\ Marcos\
Guaribas\ A vulso’; 1 female (INPA), ‘Brazil\
Roraima\ 23-à 28-II-2003’ ‘ML Oliveira &\ S.J.R.
Silva leg.\ Fazenda Trinta\ Br-210’; São Paulo: 1
male, ‘DZUP\ 022427’ ‘Brazil, São Paulo, Cajuru,\
Fazenda Rio Grande,\ 21°12′S 47°09′W,\ 25.iii.2000,
G.A.R. Melo,\ Stachytarpheta’; 1 male (RPSP), ‘Faz.
Sta Carlota\ Cajuru-SP-Brazil\ 28-XII-1986\
Camargo leg\ 863364’; 2 males (RPSP), idem except
‘863359’ ‘0863362’; 1 female, ‘DZUP\ 021822’ ‘Brazil,
São Paulo, Cajuru,\ Fazenda Rio Grande,\ 21°12′S-
47°07′W, 13.iv.2001,\ G.A.R.Melo & M.C.
Gaglianone’; 1 male, ‘DZUP\ 022426’ ‘Brazil, São
Paulo\ Cajuru, Faz. Rio\ Grande, 21°12′S\ 47°09′W,
18.iii.2000\ Gabriel A. R. Melo’; 1 female, ‘DZUP\
021828’ ‘Brazil, São Paulo, Cajuru,\ Faz. Rio Grande,
21°12′S-\ 47°09′W, 02-18.xii.\ 1999, Melo & Nasci-
mento,\ Malaise Preta’; 1 male, ‘DZUP\ 022401’
‘Brazil, São Paulo,\ Cajuru, Cachoeira,\ do Mangue,
21°11′S,\ 47°10′W, 2.xii.1999,\ Gabriel A.R. Melo’; 1
male, ‘DZUP\ 022514’ ‘Brazil, São Paulo,\ Luís
Antonio, Est.\ Ecol. de Jataí, 22.iv.\ 1999, G.A.R.
Melo’, 2 males and 2 females (DZUP), idem except
‘022513’, ‘022398’, ‘021859’, ‘021829’; 1 female (RPSP),
‘920151’ ‘Pedregulho-SP.\ Brazil-26-II-92\ Camilo,
Serrano’; 1 male, DZUP\ 022428’ ‘Brazil, São Paulo,\
Ribeirão Preto,\ Campus da USP,\ 12.III.2000, Melo’,
1 male and 1 female (DZUP), idem except ‘022408’ ‘em
Lamiaceae’, ‘021820’; 1 male, ‘DZUP\ 022409’ ‘Brazil,
São Paulo,\ Ribeirão Preto,\ Campus da USP,\
20.XI.1998, Melo\ nascido em 16.xi de larva coletada
em ninho em 02.xi’; Tocantins: 1 female, ‘DZUP\
023418’ ‘Buriti do Tocantins\ TO, Brazil\ 15-XII-
1999\ C.A.L. de Carvalho leg.’ ‘Paratetrapedia\ lin-
eata\ F.F. de Oliveira\ Det. 2000’, 2 females (DZUP),
idem except ‘023419’, ‘023420’; 1 male, ‘DZUP\
021603’ ‘Porto Nacional\ TO-Brazil\ F. Ciliar-Área
B\ Rede Entomológica\ 18.xi.1999-16:50 hs\
Japiassú, M. col.’, 1 male (DZUP), idem except
 SYSTEMATICS OF THE GENUS PARATETRAPEDIA
‘30.i.1999-16:42 hs\ Japiassú, M. col.’ ‘Paratetrape-
dia\ lineata\ (Spinola, 1851)’, 1 female (DZUP), idem
except ‘ Área A\ Rede Entomológica\ 12.i.1999-16:32
hs’; ECUADOR: 3 males and 1 female (AMNH),
‘ECUADOR, Coca\ on Rio Napo, Napo\ Pastaza
Prov.\ V. 1965’ ‘L.E. Pena\ Collector’; 1 male and
2 females (AMNH), ‘ECUADOR, Cumbaratza,
Santiago-\ Zamora Prov., S. E.\ Loja, Loja Prov.\
III-31-1965’; 1 female (AMNH), ‘Limon 900 m\ E
Ecuador\ II-1948\ Z Muller’; 1 male (SEMC),
‘ECUADOR: Sucumbios\ Sacha Lodge, 0.5°S\
76.5°W, 270 m, 4-14-III-\ 1994, Hibbs, ex: malaise’;
GUYANA: 1 female (AMNH), ‘BRITISH GUIANA:\
Kartabo, Bartica\ Dist. 1920’ ‘Trop. Research Sta-
tion\ New York Zool. Society\ no. 20275’ ‘Kartabo\
Bartica, District\ British Guiana\ 22-x-1920’ ‘Gift
of New York\ Zoo. Soc. Department.\ Tropical
Research\ William Beebe. Dir.’ ‘Chalepogenus\ xan-
thaspis F\ Cockerell\ Det. H. F. Schwarz’; 1 male
(AMNH), ‘Kartabo\ Bartica District\ British
Guiana\ 11-VI-1926’ ‘Gift of New York\ Zoo. Soc.
Department.\ Tropical Research\ William Beebe.
Dir.’; 1 female (AMNH), ‘CAMARIA\ British Guiana\
31.vii.1924’ ‘Collected by\ Jay F. W. Pearson’ ‘Chale-
pogenus\ xanthaspis F\ Cockerell\ Det. H.F.
Schwarz’; FRENCH GUIANA: 1 female (MNHP),
‘MARS’ ‘Guyane\ Nouveau Chantier\ Bas-Maroni’
‘Museum Paris\ Coll. J. Vachal 1911’ ‘Tetrap. F\
lacteipennis\ Vach’ ‘holotype’ ‘Tetrapedia\ lacteipen-
nis Vachal’; PERU: 1 male (INHS), ‘INHS\ Insect
Collection\ 53.050’ ‘Peru, JU\ 8 km N Satipo\ MEI
win 2.iii.79’ ‘INTSOY’ ‘Paratetrapedia\ Det. C. Ras-
mussen, 2003’; 1 female (MNHP), ‘Pérou\ Paquitea’
‘Museum Paris\ Coll. J. Vachal 1911’ ‘Tetrapedia\
lacteipennis\ Vach.’; 1 male and 1 female (CLAUS),
‘PERU, SM, San\ Antonio de Cumbaza\ 0625/
7624 400 masl\ 12.v.2002 C.Rasmussen\ Ex. wet
riverbank’ ‘Paratetrapedia\ sp.1\ Det. Claus Rasmus-
sen, 2002’; 1 female (AMNH), ‘Tingo Maria\ Huan.
Peru\ October 11.1946\ Alt. 2200 ft.’ ‘J.C. Pallister\
Coll. Donor\ Frank Johson’ ‘AMNH’, 1 male (AMNH),
idem except ‘December 28 1946’; 1 male (SEMC),
‘PERU: Junin Department\ Santa Ana, 7 km E\ La
Merced-Satipo Road, Km 29\ 840 m, 10°56′6′S,
75°10′54′W\ 21 OCT 1999; R Brooks\
PERU1B99094, ex: Bidens’ ‘SM0144813\ KUNHM-
ENT’, 3 males (SEMC), idem except ‘SM0146673’,
‘SM0144771’, ‘SM0146672’; 1 female (SEMC), ‘PERU:
Huanuco Department.\ Rio Pumahuasi, 930 m\
9°11′33′S, 75°57′24′W\ 11 OCT 1999, R. Brooks\ D.
Bizoska PERU1B99004’ ‘SM0147506\ KUNHM-
ENT’; SURINAME: 1 female (SEMC), ‘SURINAME:
Para\ Carolina Creek, 11 km SE\ Zanderij Airport,
30 m\ 5°23′36′N, 55°9′29′W\ 20 JUN 1999; Z.H.
Falin\ SUR1F99,ex: sweeping\ roadside vegetation’
‘SMO142227\ KUNHM-ENT’.
© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
407
PARATETRAPEDIA CAMARGOI SP. NOV.
(FIGS 75, 106, 138, 193, 194)
Comments and diagnosis: Paratetrapedia camargoi
differs from P. atlantica, P. rufa, P. albilabris, P.
andina, and P. albopilosa by the bicoloured integu-
ment (dark brown and yellow), by male T5 with band
of dense plumose hairs extending throughout its
entire margin, and yellow infumate wing membrane.
Paratetrapedia camargoi is very similar to P. chocoen-
sis in the yellow infumate wing membrane and mostly
yellow terga, differing, however, from it by the mesos-
cutum with orange yellow lateral portions (as in P.
duckei).
Description
Holotype male: Body length: 7.3; maximum head
width: 2.5; fore wing length (including tegula): 7.5.
Colour. Integument mostly yellow. Mandible yellow
with apex black; frons, upper paraocular area and
vertex mostly black; yellow stripe on paraocular area
wide, with upper portion obtuse; frons with thin
yellow stripe on mid line (about 0.5 ¥ F2 diameter);
genal area brown with thin yellow stripe (about
1.2 ¥ F2 diameter), occupying all ocular margin; scape
yellow; pedicel and flagellomeres with anterior
surface pale yellow and ventral surface reddish
brown. Disc of mesoscutum black and lateral portions
orange yellow, with two thin yellow stripes on disc
and lateral margins; axilla dark brown. Terga mostly
yellow, marginal zone weakly orange yellow; mid
portion of marginal zone dark brown. Basitarsus of
mid and fore legs weakly dark brown; tibial spurs
pale yellow. Wing membrane yellow infumate with
yellow microtrichia; fore wing membrane with apical
third dark because of brown microtrichia; veins and
pterostigma orange yellow. Pubescence. Mostly
yellow, except mid and fore basitarsus with dark
brown pubescence. Scape with brownish long setae,
about 0.10–0.16 mm in length. Mesoscutum and
scutellum with dense short plumose yellow pubes-
cence; scutellum with short erect simple setae, about
0.10–0.17 mm in length; metapostnotum with dense
fine plumose hairs, about 0.10 mm in length. T4 with
band of dark brown hairs throughout margin; T5–T6
with marginal band of yellow hairs throughout its
margin; S2 with row of setae adjacent to margin, with
two small rows of simple erect setae isolate on mid
third of margin; S3 of male deeply concave area in ‘U’
shape, covered by dense short plumose pubescence;
S6 with two rows of stout plumose hairs on margins
laterally, and short plumose hairs on margin of apex.
Integument sculpture. Clypeus with coarse punctures
(1 pd), intermingled amongst sparse minute punc-
tures (2 pd); supraclypeal area with dense coarse
punctures (0.5 pd); disc of frons with coarse punc-
408 A. J. C. AGUIAR and G. A. R. MELO
tures, mostly sparse (0.5–2 pd); antennal scrobe with
dense minute punctures (0.5 pd). Mesoscutum with
dense fine punctures (< 0.5 pd), intermingled amongst
sparse coarser punctures (2 pd); metapostnotum with
dense fine punctures (1 pd); mesepisternum mostly
with sparse fine punctures (> 3 pd). Structure.
Lamella of pronotal collar with lateral portions
obtuse, in dorsal view. Disc of scutellum convex. Head
about 1.2¥ broader than long (2.3 : 1.8); ratio of lower
to upper interocular distance: 0.83 (1.0 : 1.2); clypeus
about 1.5¥ broader than long (1.0 : 0.66); scape,
length 0.6; maximum width, 0.2; length of F1–F3:
0.17, 0.14, 0.19; diameter of F2: 0.2.
Etymology: The species name is dedicated to Dr João
Maria Franco de Camargo.
Distribution: Bolivia; Brazil: Acre, Amapá, Amazonas
(Fig. 75).
Type material: Holotype male, ‘DZUP\ 022940’
‘Brazil, Amazonas,\ Manaus, Hotel Tropical\
19.vi.2003, G.A.R. Melo’. Paratypes: BOLIVIA: 1 male
(AMNH), ‘BOLIVIA, Beni: Rio Itenez\ opposite Costa
Marques\ (Brazil), September 1–3,1964\ Bouseman
& Lussenhop’; BRAZIL: Acre: 1 male, ‘DZUP\ 021913’
‘R. Branco Acre\ BR 15-20-XI-61\ F.M. Oliveira’, 10
males (DZUP), idem except ‘021929’, ‘022941’ to
‘022943’, ‘022936’, ‘022946’, ‘022951’, ‘022948’ to
‘022950’, ‘023662’; Amapá: 1 male (IEPA), ‘BRAZIL-
Amapá\ I.E.P.A.’ ‘BRAZIL – Amapá\ Mun. Serra do
Navio\ Serra do Canga\ Ambiente de mata\ 08-II-
2001\ J. Madson/C. Henrique’; 1 male (IEPA),
‘BRAZIL/Amapá\ I.E.P.A.’ ‘BRAZIL-Amapá\ Lar.do
Jari/Centrinho\ Ambiente de mata\ 00:36:06S/
52:20:46W\ 21/V/2001\ J.Chaves’ ‘Percurso\ 3’; Ama-
zonas: 1 male, ‘DZUP\ 022933’ ‘MANAUS-AM\
Brazil-VII 62\ F.M. OLIVEIRA’; 1 male, DZUP\
022935’ ‘Brazil, Amazonas, Rio\ Negro, Santa
Isabel,\ 0°25′S, 65°01′W,\ 11.vii.1999, G.A.R. Melo’.
PARATETRAPEDIA CHOCOENSIS SP. NOV.
(FIGS 57, 75, 109, 139, 201, 202)
Comments and diagnosis: Paratetrapedia chocoensis
differs from P. atlantica, P. rufa, P. albilabris, P.
andina, and P. albopilosa by the bicoloured integu-
ment (dark brown and yellow), by male T5 with band
of dense plumose hairs extending throughout its
entire margin, and yellow infumate wing membrane.
Paratetrapedia chocoensis is very similar to P. fla-
veola, differing by having the lamella of pronotal
collar with lateral portions diverging of margins of
mesoscutum, yellow infumate wing membrane,
completely orange yellow terga, yellow stripe on
© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
paraocular area weakly sinuous with upper portion
obtuse, and smaller body size.
Description
Holotype male: Body length: 6.5; maximum head
width: 2.3; fore wing length (including tegula): 6.3.
Colour. Integument mostly yellow. Mandible yellow
with apex black; labrum, clypeus, and supraclypeal
area yellow; yellow stripe on paraocular area on
more than half of lower portion, and weakly sinuous
and obtuse on upper portion; frons dark brown with
thin short yellow stripe on mid line; genal area
yellow; scape orange yellow; pedicel and flagellom-
eres reddish brown. Mesoscutum black with two
thin yellow stripes on disc and lateral margins;
scutellum and metanotum yellow; lateral area of
metanotum yellow; axilla brown laterally; metapost-
notum orange yellow, slightly dark; mesepisternum
with anterior surface reddish brown and lateral
surface yellow. Terga completely orange yellow. Wing
membrane yellow infumate, with microtrichia
yellow; fore wing membrane with apical third dark
by brown microtrichia; veins and pterostigma
orange yellow. Legs yellow; hind basitarsus brown;
tibial spurs white. Pubescence. Mostly orange
yellow, except hind basitarsus with dark brown
pubescence. Scape with long pale yellow setae on
inner face, about 0.2 mm in length; mesoscutum and
scutellum with dense yellow short plumose pubes-
cence; scutellum with long erect simple setae, about
0.16 mm in length; metapostnotum with short
plumose hairs, about 0.1 mm in length. T4–T6 with
marginal band of plumose yellow hairs throughout
its margin; S2 with two short rows on mid portion
of margin; S3 of male deeply concave in ‘U’ shape,
covered by dense short plumose pubescence; S6 with
two rows of stout plumose hairs on margins later-
ally, and short plumose hairs on margin of apex.
Integument sculpture. Clypeus and supraclypeal
area with coarse punctures (1–2 pd), intermingled
amongst sparse minute punctures (> 2 pd); disc of
frons with coarse punctures (2 pd), and minute
punctures on lateral portions (1 pd). Mesoscutum
and scutellum with fine dense punctures (< 0.5 pd);
metapostnotum with fine punctures, mostly sparse
(1–2 pd); mesepisternum with dense coarse punc-
tures laterally (0.5–1 pd). Structure. Lamella of
pronotal collar with lateral portions weakly obtuse.
Scutellum disc weakly convex, almost flat. Head
about 1.2¥ broader than long (2.4 : 1.8); ratio of
lower interocular distance to upper interocular
distance: 0.81 (1.1 : 1.35); clypeus about 1.75¥
broader than long (1.05 : 0.6); scape: length 0.6,
width 0.2; length of F1–F3: 0.17, 0.12, 0.17; diam-
eter of F2: 0.2.
 SYSTEMATICS OF THE GENUS PARATETRAPEDIA
Paratype female: Body length: 6.6; maximum head
width: 2.5; fore wing length (including tegula): 7.0.
Colour. Similar to male; wing membrane yellow infu-
mate, darker than in female. Pubescence. Similar to
male; scutellum with erect simple setae, about
0.2 mm in length; marginal bands on tergal margin,
similar to male, except for T4–T6 with marginal hair
band of simple orange yellow hairs occupying whole
margin. Integument sculpture. Similar to male;
clypeus with dense coarse punctures (< 0.5 pd); supra-
clypeal area with dense coarse punctures inter-
mingled amongst minute sparse punctures (> 2 pd);
frons with sparse coarse punctures on disc and dense
fine punctures on lateral portions (< 0.5 pd); meta-
postnotum with dense minute punctures (1 pd) on
lateral portions, with coarser punctures on disc.
Structure. Lamella of pronotal collar with lateral
portions obtuse. Disc of scutellum almost flat, and
profile slightly truncated. Head about 1.2¥ broader
than long (2.5 : 2.0); ratio of lower to upper interocu-
lar distance: 0.81 (1.2 : 1.4); clypeus about two times
broader than long (1.1: 0.55); scape: length, 0.7,
maximum width, 0.2; length of F1–F3: 0.22, 0.2, 0.17;
diameter of F2: 0.2.
Variation: Most male specimens have marginal hair
band on T4 incomplete, occupying less than one-third
of margin laterally, but a few specimens have one
isolated band on the mid portion of margin. Males
and females can have the colour of mesepisternum,
propodeum, and metapostnotum completely yellow or
reddish brown. Three specimens from Ecuador have
wing membrane pale brown infumate, leg pubescence
completely yellow, metapostnotum and ventral
portion of mesepisternum reddish brown. The speci-
men from Colombia, collected in the Chocó region, has
the same colour pattern as the holotype, except for
the hind basitarsus with mostly yellow pubescence
and a few sparse long dark brown setae.
Etymology: The species name refers to the Choco
region.
Distribution: Colombia; Costa Rica; Ecuador; Panama
(Fig. 75).
Type material: Holotype male (BLCU), ‘PANAMA:
Panama Prov\ Gamboa- Pipeline Road\ 8 km N
Turnera pana\ mensis 12 may 1985\ 10-1130 am D
Roubik’ ‘NativeBeeSurvey\ USDA, Logan, Utah\
BBSL517242’. Parátipos: COLOMBIA: 1 female
(SEMC), ‘COLOMBIA: Valle.\ Rio Anchicaya,
400 m.\ IX-28-76. Bell,\ Breed & Michener’; COSTA
RICA: 1 female (SEMC), ‘Costa Rica San José Prov.\
Rio Damitas, 14,5 km.\ N. Puerto Quepos\ 200 m.’
‘16 August 1962\ (A.Wille & C.D. Michener)’; 1 male
© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
409
and 5 females (SEMC), ‘Costa Rica (S.) Puntarenas\
Prov., Gromaco, 34 km. SE of\ Potrero Grande, on Rio
Coto\ Brus. 21 July 1963, 1000ft.\ (C.D. Michener &
W. Kerfoot)’; 1 female (SEMC), ‘Costa Rica San José\
Playon, 8 km. N. of\ Parrita, 30 m., 14/19/August
1962 (C.D.\ Michener and A. Wille)’; ECUADOR: 1
male (SEMC), ‘ECUADOR\ CA. PUERTO QUITO\
9-VI-1979\ C. Porter, A. Cerbone’, 1 male (SEMC),
idem except ‘Paratetrapedia\ det RW Brooks’; 1 male
(AMNH), ‘ECUADOR: Pichincha:\ Tinalandia, 17 km
SE\ Sto. Domingo de los\ Colourados, 3.000’\
October 18. 1988\ leg. J.S. Miller’; PANAMA: 1
female (BLCU), ‘PANAMA Panama Prov\ Barro
Colourado Is.\ Snyder-Molino tr.\ UV light trap, 5\
may 1992 D. Roubik’; 1 female (BLCU), ‘PANAMA
Panama Prov\ Barro Colourado Is.\ Snyder-Molino
tr.\ UV light trap, 5\ May 1992 D. Roubik’ ‘Paratet-
rapedia\ aff.\ F lineata\ TGriswold det. 93’ ‘Native-
BeeSurvey\ USDA, Logan, Utah\ BBSL517244’; 1
female (BLCU), ‘PANAMA: Panama Prov\ Parque
Nac. Soberania\ Gamboa-10 kmN on\ Turnera pana-
mensis\ 5 April 1988 D Roubik’ ‘NativeBeeSurvey\
USDA, Logan, Utah\ BBSL517243’; 6 females
(SEMC), ‘Panama Colon Prov.\ Pipeline Road.,
5 km.\ NW. Gamboa (C.Z.)\ on Aeschnyomena\
americana. 12 January \ 1981. C.D. Michener’; 1
female (SEMC), ‘Barro Colourado\ Isl. Panama’ ‘R.W.
Dawson\ March 1/1937’ ‘Chalepogenus\ sp.?\ Det.\
C.D. Michener ‘51’; 4 females (SEMC), ‘Panamá:
Panama Prov.\ 2 km E. Fort Sherman\ 30 January
1980\ Brian H. Smith’; 1 male (SEMC), ‘Barro Colo.
Is. C.Z.\ 24V1952 No.571\ Carl W. Rettenmeyer’, 1
female (SEMC), idem except ‘06IV1952 No.336’; 1
female (SEMC), ‘Barro Colo. Is. C.Z.\ 06IV1952
No.336\ Carl W. Rettenmeyer’; 1 female (SEMC),
‘PANAMA-Canal Zone\ Barro Colourado Island\
7.VII.1956\ Carl.W. & Marian E.\ Rettenmeyer,
No.2088’; 1 male (SEMC), ‘Panama: Darién, Cana\
Biological Station 500–550 m\ 7°45′18′N,77°41′6′W\
03 June 1996; J.Ashe,\ R. Brooks PAN1AB96015\ ex:
Ludwigia’.
PARATETRAPEDIA ATLANTICA SP. NOV.
(FIGS 76, 110, 145, 195, 196)
Comments and diagnosis: Paratetrapedia atlantica is
very similar to the species with dark brown integu-
ment of the flavipennis group. It differs from these
species by pubescence on posterolateral margin of
scutellum composed of dark brown plumose hairs and
mesoscutum completely black with faint yellow
stripes on disc. The female is also distinct by T1 with
minute punctures (mainly on vertical surface), and
transition zone between metapostnotum and propo-
deum with minute punctures. The male of P. atlantica
410 A. J. C. AGUIAR and G. A. R. MELO
is distinct from P. albopilosa by T5 with completely
glabrous margin.
Description
Holotype male: Body length: 8.1; maximum head
width: 2.6; fore wing length (including tegula): 8.1.
Colour. Integument mostly dark brown; mandible
yellow with apex black; labrum black, weakly yellow;
clypeus with two weak faint yellow spots on lateral
margins; lower paraocular area with thin faint yellow
stripe, occupying more than lower half of paraocular
area; scape, pedicel, and flagellomeres reddish brown.
Central portion of metanotum weakly orange yellow.
Wing membrane brown infumate with dark microtri-
chia dispersed; veins and pterostigma orange yellow.
Tibial spurs white. Pubescence. Mostly dark brown,
except paraocular areas, mesepisternum, and propo-
deum with white pubescence. Mesoscutum and scutel-
lum with dense plumose pale brown pubescence;
posterolateral margin of scutellum with long plumose
dark brown hairs; metapostnotum with plumose
hairs, about 0.12 mm in length; T4 with few brown
plumose hairs on less than one third of margin lat-
erally; T5–T6 with margin glabrous; S2 with two
short rows of simple erect setae on mid portion of
margin; S3 of male with deeply concave area in ‘U’
shape covered by dense short plumose pubescence; S6
with two rows of plumose hairs on margin laterally
and short plumose hairs on margin of apex. Integu-
ment sculpture. Clypeus and supraclypeal area with
shallow coarse punctures (0.5–2 pd), intermingled
amongst sparse minute punctures (> 2 pd); frons with
coarse punctures, mostly sparse (2 pd), intermingled
amongst minute punctures on upper portion; anten-
nal scrobe with dense minute punctures (< 0.5 pd).
Mesoscutum and scutellum with minute punctures
(< 0.5 pd), intermingled amongst sparse coarse punc-
tures (2 pd); metapostnotum with dense minute punc-
tures on lateral portions (< 0.5 pd), and coarser
punctures on disc (0.5 pd); mesepisternum with
coarse sparse punctures (> 3 pd), with few dense
punctures on omaular area (0.5–1 pd). Structure.
Lamella of pronotal collar obtuse laterally and weakly
divergent. Scutellum disc strongly convex. Head
about 1.2¥ broader than long (2.65 : 2.1); ratio of
lower to upper interocular distance: 0.78 (1.1 : 1.4);
clypeus about 1.7¥ broader than long (1.1 : 0.65);
scape: length 0.65, maximum width 0.17; length of
F1–F3: 0.2, 0.15, 0.18; diameter of F2: 0.18.
Paratype female: Body length: 8.0; maximum head
width: 2.65; fore wing length (including tegula): 8.0.
Colour. Similar to male, except for paraocular area
and frons without yellow marks. Pubescence.
Similar to male, except for T5–T6 with marginal
band throughout margin. Integument sculpture.
© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
Similar to male except for clypeus and supraclypeal
area with dense coarse punctures (0.5–1 pd); disc of
frons with dense coarse punctures (< 0.5–2 pd);
coarse punctures on disc of frons almost with same
diameter; lateral portions of frons and antennal
scrobe with dense fine punctures (< 0.5 pd). Struc-
ture. Lamella of pronotal collar with lateral portions
conspicuously obtuse. Scutellum disc weakly convex.
Head about 1.2¥ broader than long (2.65 : 2.15);
ratio of lower interocular distance to upper interocu-
lar distance: 0.86 (1.25 : 1.45); clypeus about 1.8¥
broader than long (1.1 : 0.6); scape: length 0.75,
maximum width 0.2; length of F1–F3: 0.22, 0.12,
0.17; diameter of F2: 0.25.
Variation: Most specimens have the only a very faint
yellow stripe on paraocular area, clypeus completely
brown with narrow yellow stripes on margins later-
ally, and genal area completely brown. Colour of terga
of P. atlantica can vary between pale brown to dark
brown, almost black. One male specimen has a com-
pletely smooth frons.
Etymology: The species name refers to the Atlantic
Forest.
Distribution: Brazil: Espírito Santo, Rio de Janeiro,
São Paulo (Fig. 76).
Type material: Holotype male, ‘DZUP\ 022893’
‘REPRESA RIO GRANDE\ Guanabara Brazil\ abril
1961\ F.M. Oliveira’; parátipos: BRAZIL: Espírito
Santo: 1 female, ‘DZUP\ 022016’ ‘Brazil, Espirito
Santo,\ Linhares, V.1982,\ C.Elias’; 1 male, ‘DZUP\
022898’ ‘Conc. da Barra-E. Sto\ Brazil 2-8/V/68\ C. &
C.T. Elias leg.’, 4 males (DZUP), idem except ‘26/4/
1968’ ‘022828’, ‘26/04/1968’ ‘022830’, ‘022912’ ‘26/8/
1969’, ‘022995’ ‘26/8/1969’; 1 male, ‘DZUP\ 022925’
‘Conc. da Barra-E.Sto\ Brazil 2-7/X/68\ C. & C.T.
Elias leg.’; 1 female, ‘DZUP\ 022771’ ‘Conc. da Barra-
ES\ Brazil-11/8/1969\ C.T. & C.Elias’, 2 females
(DZUP), idem except ‘022915’ ‘12/5/1969’ ‘022908’; 1
male, ‘DZUP\ 022848’ ‘Conc. da Barra-ES\ Brazil-17/
9/1969\ C.T. & C.Elias’; 1 female, ‘DZUP\ 022770’
‘Conc. da Barra-ES\ Brazil-19/5/1969\ C.T. & C.
Elias’, 1 female (DZUP), idem except ‘022834’; 1
female, ‘DZUP\ 022837’ ‘Conc. da Barra-ES\ Brazil-
4/9/1969\ C.T. & C. Elias’; 1 male, ‘DZUP\ 022992’
‘Conc. da Barra-ES\ BRAZIL-5/5/1968\ C.T. & C.
Elias’; 1 male, ‘DZUP\ 022847’ ‘Conceição da Barra\
ES-BR 16-23/IV/69\ C. & C.T. Elias leg.’; 1 female,
‘DZUP\ 022919’ ‘Conceição da Barra\ ES-BR 1-8/IV/
69\ C. & C.T. Elias leg.’, 2 females (DZUP), idem
except ‘022931’, ‘022917’; 1 male, ‘DZUP\ 022917’
‘Conceição da Barra\ ES-BR 4/7/1969\ C.T. & C.
Elias leg.’; 1 male, ‘DZUP\ 022774’ ‘Conceição da
 SYSTEMATICS OF THE GENUS PARATETRAPEDIA 411

Figure 76. Geographical distribution of Paratetrapedia atlantica sp. nov., Paratetrapedia rufa sp. nov., Paratet-
rapedia albilabris, Paratetrapedia andina sp. nov., and Paratetrapedia albopilosa sp. nov.

Figures 77–84. Sternum 6 (S6), male; 77: Paratetrapedia hyalinata sp. nov. (Brazil, Pará); 78: Paratetrapedia
flaveola sp. nov. (Brazil, Minas Gerais, Três Marias); 79: Paratetrapedia scaposa sp. nov. (Brazil, Acre, Cruzeiro do
Sul); 80: Paratetrapedia lineata (Brazil, Pará, Óbidos); 81: Paratetrapedia vogeli sp. nov. (Peru, Cuzco, Quincemil); 82:
Paratetrapedia rufa sp. nov. (Bolivia, Santa Cruz, Santa Cruz de La Sierra); 83: Paratetrapedia bicolor (Brazil,
Espírito Santo, Conceição da Barra); 84: Paratetrapedia mexicana sp. nov. (Mexico, Jalisco, San Patricio); scale
bars = 0.1 mm (Fig. 77); 0.05 mm (remaining figures).

© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
412 A. J. C. AGUIAR and G. A. R. MELO

Barra-ES\ Brazil-5/5/1968\ C. & C.T. Elias leg.’, 2
males (DZUP), idem except ‘022829’, ‘022914’; Rio de
Janeiro: 1 male, ‘DZUP\ 022855’ ‘Rep. Rio Grande-
GB\ Brazil 10-IV-1962\ F.M.Oliveira leg’, 1 male
(DZUP), idem except ‘022863’; 1 male, ‘DZUP\
022928’ ‘REPRESA RIO GRANDE\ Guanabara
Brazil\ abril 1961\ F.M. Oliveira’, 5 males (DZUP),
idem except ‘022852’, ‘022768’, ‘022782’, ‘022926’,
‘022994’; 1 female, ‘DZUP\ 022978’ ‘Km.47 Estrada
Rio-S.Paulo\ Mun. Itaguay Est.Rio\ 10/947 Wygod
col.’; 1 male, ‘DZUP\ 022923’ ‘Itatiaia GB\ Brazil
IV-61\ F.M. Oliveira’, 2 males (DZUP), idem except

© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
 SYSTEMATICS OF THE GENUS PARATETRAPEDIA 413

Figures 85–100. Sternum 7 (S7), male; 85: Paratetrapedia lugubris (Brazil, Rondônia, Vilhena); 86: Paratetrapedia
connexa (Guatemala); 87: Paratetrapedia hyalinata sp. nov. (Brazil, Pará); 88: Paratetrapedia volatilis (Brazil,
Paraná, Piraquara); 89: Paratetrapedia leucostoma (Brazil, Maranhão, São Luís); 90: Paratetrapedia moesta (Mexico,
Jalisco, San Patricio); 91: Paratetrapedia scaposa sp. nov. (Brazil, Acre, Cruzeiro do Sul); 92: Paratetrapedia
bifrons sp. nov. (Mexico, Veracruz, Fortin de las Flores); 93: Paratetrapedia basilaris sp. nov. (Brasil, Amazonas,
Manaus); 94: Paratetrapedia vogeli sp. nov. (Peru, Cuzco, Quincemil); 95: Paratetrapedia bicolor (Brazil, Espírito
Santo, Conceição da Barra); 96: Paratetrapedia testacea (Brazil, Pará, Óbidos); 97: Paratetrapedia calcarata (Costa Rica,
Puntarenas); 98: Paratetrapedia punctata sp. nov. (Brazil, Minas Gerais, Pratinha); 99: Paratetrapedia fervida
(Brazil, Espírito Santo, Santa Teresa); 100: Paratetrapedia mexicana sp. nov. (Mexico, Jalisco, San Patricio); scale
bars = 0.1 mm.
䉳

‘022897’, ‘022015’; 1 female, ‘DZUP\ 022773’
‘Sumaré\ D. Federal\ BR.II.1955\ M.Alvarenga’; 7
males and 4 females (SEMC), ‘BRAZIL Rio de Jan-
eiro\ Baia da Guanabara\ Floresta dos Macacos\
April’61 (M. Alvarenga)’; 1 male and 1 female
(SEMC), ‘BRAZIL Rio de Janeiro\ Baia da Guana-
bara\ Floresta dos Macacos\ 16 February 1961\ (M.
Alvarenga)’; 1 male (SEMC), ‘BRAZIL Rio de Janeiro\
Represa Rio Grande\ Dec’60 (M. Alvarenga)’; 1 male,
‘DZUP\ 022783’ ‘Est. Vista Chinesa\ Distrito Federal
Brazil\ 17-II-953\ C.A.C. Seabra’; São Paulo: 1
female (MZSP), ‘Ubatuba, SP.\ Km.240, Brazil\
31.III.1962\ K. Lenko col.’.
PARATETRAPEDIA RUFA SP. NOV.
(FIGS 50, 62, 76, 82, 111, 142, 189, 190)
Comments and diagnosis: Paratetrapedia rufa is very
similar structurally to P. atlantica, differing mainly
by its orange yellow integument and mostly smooth
female T1. Females of P. rufa can be confused with
those of P. testacea because of their orange yellow
integument, but the former has an obtuse lamella on
the pronotal collar and metapostnotum mostly with
densely fine punctures (1 pd).
brous, with few setae on margin laterally; S2 with two
small rows of simple setae on posterior margin; S3 of
male with deeply concave area in ‘U’ shape covered by
dense short plumose pubescence; S4 and S5 typical of
genus; S6 with two rows of stout plumose hairs on
margin laterally, and short plumose hairs on margin
of apex. Integument sculpture. Clypeus with dense
coarse punctures (1 pd) intermingled amongst dense
minute punctures (1 pd); supraclypeal area with
dense coarse punctures (< 0.5 pd) intermingled
amongst minute punctures (1–2 pd); disc of frons with
dense coarse punctures (< 0.5 pd), and dense fine
punctures on lateral portions and antennal scrobe
(< 0.5 pd). Mesoscutum and scutellum with dense fine
punctures (< 0.5 pd); metapostnotum with dense fine
punctures (1 pd), and minute punctures on lateral
margins (1 pd); mesepisternum with dense coarse
punctures, mostly sparse (2 pd). Structure. Lamella of
pronotal collar with lateral portions obtuse. Scutel-
lum disc convex, weakly concave on mid line. Head
about 1.2¥ broader than long (2.5 : 2.1); ratio of lower
to upper interocular distance: 0.82 (1.15 : 1.4); clypeus
about 1.7¥ broader than long (0.57 : 0.32); scape:
length 0.65, maximum width 0.22; length of F1–F3:
0.17, 0.15, 0.2; diameter of F2: 0.2.

Description Paratype female: Body length: 7.4; maximum head

Holotype male: Body length: 8.7; maximum head
width: 2.5; fore wing length (including tegula): 8.5.
Colour. Integument mostly orange yellow; face with
yellow marks weakly conspicuous (Fig. 50). Mesoscu-
tum with two thin yellow stripes on disc and lateral
margins; disc of mesoscutum dark brown and lateral
portions orange yellow (Fig. 62); axilla dark brown
laterally. Wing membrane orange yellow infumate,
with brown microtrichia; veins and pterostigma
orange yellow. Tibial spurs pale yellow. Pubescence.
Mostly orange yellow. Legs with orange yellow pubes-
cence. Scape with brown short setae on inner face,
about 0.2 mm in length. Mesoscutum and scutellum
with pale yellow plumose hairs; scutellum with long
erect simple setae, about 0.2 mm in length; metapost-
notum with dense short plumose hairs, about
0.14 mm in length. Margins of T4–T6 completely gla-
width: 2.6; fore wing length (including tegula): 7.4.
Colour. Similar to male, except for disc of mesoscutum
pale orange. Pubescence. Similar to male, except for
inner surface of scape with short simple yellow setae,
about 0.04 mm in length; mesoscutum and scutellum
with short pale yellow plumose pubescence; scutellum
with long erect simple setae, about 0.2 mm in length;
metapostnotum with dense short plumose pubes-
cence, about 0.08 mm in length. T5–T6 with marginal
hair band distributed throughout its margin. Integu-
ment sculpture. Similar to male, except for clypeus
with dense coarse punctures (0.5–1 pd); supraclypeal
area with dense coarse punctures (< 0.5 pd); disc of
frons with dense coarse punctures (< 0.5 pd); lateral
portions of frons and antennal scrobe with dense fine
punctures (0.5 pd); mesoscutum and scutellum with
dense fine punctures (0.5 pd) intermingled amongst

© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
414 A. J. C. AGUIAR and G. A. R. MELO

© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
 SYSTEMATICS OF THE GENUS PARATETRAPEDIA 415

Figures 101–115. Sternum 7 (S7), male; 101: Paratetrapedia flavifrons sp. nov.; 102: Paratetrapedia romani (Brazil,
Amazonas, Manaus); 103: Paratetrapedia tocantinensis sp. nov. (Brazil, Goiás, Campos Belos); 104: Paratetrapedia
lineata (Brazil, Pará, Óbidos); 105: Paratetrapedia flaveola sp. nov. (Brazil, Minas Gerais, Três Marias); 106:
Paratetrapedia camargoi sp. nov. (Brazil, Acre, Rio Branco); 107: Paratetrapedia albilabris (Costa Rica, Montezuma);
108: Paratetrapedia andina sp. nov. (Bolivia, Yungas, Puerto Villa); 109: Paratetrapedia chocoensis sp. nov.
(Panama, Barro Colorado); 110: Paratetrapedia atlantica sp. nov. (Brazil, Espírito Santo, Conceição da Barra); 111:
Paratetrapedia rufa sp. nov. (Bolivia, Santa Cruz, Santa Cruz de la Sierra); 112: Paratetrapedia albopilosa sp. nov.
(Brazil, Maranhão, São Luís); 113: Paratetrapedia flavescens sp. nov. (Mexico, San Luis Potosi, Xilitla); 114.
Paratetrapedia duckei (French Guiana); 115: Paratetrapedia flavipennis (Brasil, Amazonas, Manaus); scale bar = 0.1 mm.
䉳

coarser and sparser punctures (1–4 pd); meta-
postnotum mostly with dense minute punctures
(0.5–1 pd), with some coarser punctures on disc;
mesepisternum mostly with sparse coarse punc-
tures (1–3 pd). Structure. Lamella of pronotal collar
with lateral portions obtuse. Scutellum disc weakly
convex, almost flat. Head about 1.2¥ broader than
long (2.9 : 2.1); ratio of lower to upper interocular
distance 0.85 (1.4 : 1.2); clypeus about two times
broader than long (1.2 : 0.6); scape: length 0.7;
maximum width 0.2; length of F1–F3: 0.17, 0.12, 0.17;
diameter of F2: 0.2.
Etymology: The species name refers to the red orange
colour of the integument.
disc of frons; mandible, labrum, clypeus, and genal
area completely dark brown in the male of P. albila-
bris. Male and female have T4 with only a short band
of simple hairs, occupying less than one third of
margin laterally; female T5–T6 with marginal band
throughout its margin, and completely glabrous in
males.
Distribution: Mexico; Costa Rica (Fig. 76).
Type material: Tetrapedia albilabris Friese, holotype
male (ZMB), ‘Costa Rica\ San Jose’ ‘Tetrapedia\ albi-
labris\ M 1915 Friese det. 1915 Fr.’ ‘Type’ ‘Zool.
Mus.\ Berlin’. The row of hairs on mid portion of S2
margin is slightly deformed in the holotype.

Distribution: Bolivia; Brazil: Pará (Fig. 76). Additional material: See Appendix.

Type material: Holotype male, ‘DZUP\ 026537’

‘LIMBO BOLIVIA\ COCHABAMBA IV 53\ Mar-
tinez, leg’; paratypes: BOLIVIA: 1 male (AMNH),
‘BOLIVIA: Santa Cruz\ Santa Cruz de la Sierra\
Jardim Botánico\ 31-VII-1-VIII-1976\ C. Porter, C.
Calmbacher’; BRAZIL: 1 female, ‘DZUP\ 022524’
‘Coleção\ Campos Seabra’ ‘Rio Acará\ Pará BRAZIL\
24-VIII-1954\ N. Cerqueira’, 2 females (DZUP), idem
except ‘022528’, ‘022525’.
PARATETRAPEDIA ALBILABRIS (FRIESE, 1916)
(FIGS 65, 76, 107, 146, 205, 206)
Tetrapedia albilabris Friese 1916: 335; holotype male,
Costa Rica: San José, San Jose (ZMB, examined).
Tetrapedia albilabris; Lutz & Cockerell (1920): 568.
Paratetrapedia albilabris; Aguiar (2007): 621.
Paratetrapedia (Paratetrapedia) albilabris (Friese,
1916); Rasmussen & Ascher (2008): 22.
Comments and diagnosis: Paratetrapedia albilabris is
very similar to P. albopilosa, both having the poste-
rolateral portions of the scutellum with numerous
white plumose hairs (Fig. 65). The female differs from
P. albopilosa by T1 with dense minute punctures on
vertical surface, and the male by lack of yellow marks
on face, except for a thin yellow stripe on mid line of
PARATETRAPEDIA ALBOPILOSA SP. NOV.
(FIGS 76, 112, 143, 203, 204)
Comments and diagnosis: Paratetrapedia albopilosa
is very similar to P. atlantica and P. albilabris. It
differs from both P. atlantica and P. albilabris
by frons weakly convex and area of transition
between metapostnotum and scutellum smooth, and
from P. atlantica by white plumose pubescence on
posterolateral portions of scutellum (as in P. albila-
bris, Fig. 65). The male is distinct by large yellow
marks on frons, labrum completely yellow, paraocu-
lar area with large yellow stripe and genal area
with thin yellow stripe on lower third. The male of
P. albopilosa also differs from P. albilabris by hind
basitarsus without an acute tooth on anterior
margin.
Description
Holotype male: Body length: 6.8; maximum head
width: 2.5; fore wing length (including tegula): 7.3.
Colour. Integument mostly dark brown; terga and
sterna pale brown, almost yellow; mandible yellow
with apex black; labrum pale yellow; clypeus with two
pale yellow spots on lateral margins; supraclypeal
area weakly pale yellow; paraocular area with yellow
stripe; disc of frons with thin yellow stripe on mid

© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
416 A. J. C. AGUIAR and G. A. R. MELO

© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
 SYSTEMATICS OF THE GENUS PARATETRAPEDIA 417

Figures 116–128. Sternum 8 (S8), male; 116: Paratetrapedia lugubris (Brazil, Rondônia, Vilhena); 117: Paratetrapedia
connexa (Guatemala); 118: Paratetrapedia hyalinata sp. nov. (Brazil, Pará); 119: Paratetrapedia volatilis (Brazil,
Paraná, Piraquara); 120: Paratetrapedia moesta (Mexico, Jalisco, San Patricio); 121: Paratetrapedia leucostoma (Brazil,
Maranhão, São Luís); 122: Paratetrapedia scaposa sp. nov. (Brazil, Acre, Cruzeiro do Sul); 123: Paratetrapedia
bifrons sp. nov. (Mexico, Veracruz, Fortin de las Flores); 124: Paratetrapedia bicolor (Brazil, Espírito Santo, Conceição
da Barra); 125: Paratetrapedia calcarata (Costa Rica, Puntarenas); 126: Paratetrapedia testacea (Brazil, Pará, Óbidos);
127: Paratetrapedia vogeli sp. nov. (Peru, Cuzco, Quincemil); 128: Paratetrapedia basilaris sp. nov. (Brazil,
Amazonas, Manaus); scale bars = 0.1 mm.
䉳

line, weak; genal area with small yellow stripe on
lower third; scape, pedicel, and flagellomeres reddish
brown. Mesoscutum with two thin yellow stripes very
faint on disc. Fore wing membrane brown infumate
because of dark brown microtrichia; veins and
pterostigma yellow. Tibial spurs white. Pubescence.
Genal area, mesepisternum, propodeum, and sterna
with pubescence mostly white; remaining body with
pubescence brown. Mesoscutum and scutellum with
dense plumose pale brown pubescence; posterolateral
portions of scutellum with dense pale brown pubes-
cence; metapostnotum with short plumose hairs,
about 0.08 mm in length. T4–T6 with margins mostly
glabrous; S2 with two short rows of simple erect setae
on mid portion; S3 deeply concave in ‘U’ shape,
covered by dense short plumose pubescence; S4 and
S5 typical of the genus; S6 with two rows of stout
plumose hairs on margins laterally, and short
plumose hairs on margin of apex. Integument sculp-
ture. Clypeus with shallow coarse punctures
(0.5–2 pd), intermingled amongst sparse minute
punctures (2 pd); supraclypeal area with dense
coarse punctures on lateral portions (< 1 pd), and
sparse coarse punctures on disc (> 2 pd); frons with
heterogeneous punctures: coarse punctures mostly
sparse (2 pd), intermingled amongst sparse minute
on upper portion of disc; integument amongst
punctures smooth and shiny; antennal scrobe with
dense minute punctures (< 0.5 pd). Mesoscutum and
scutellum with dense minute punctures (< 1 pd),
intermingled amongst sparse coarser punctures
(2 pd); metapostnotum with dense fine punctures on
disc (0.5 pd) and dense minute punctures on lateral
margins (< 0.5 pd); transition area between meta-
postnotum and propodeum with smooth and shiny
area; mesepisternum with coarse punctures, mostly
sparse (> 3 pd), with some denser punctures on
omaular area (0.5–1 pd). Structure. Lamella of
pronotal collar with lateral portions obtuse. Scutel-
lum conspicuously convex. Head about 1.2¥ broader
than long (2.5 : 2.1); ratio of lower to upper interocu-
lar distance: 0.8 (1.05 : 1.3); clypeus about 1.5¥
broader than long (1.0 : 0.65); scape: length 0.82,
maximum width 0.8; length of F1–F3: 1.8, 0.14, 0.2;
diameter of F2: 0.16.
Paratype female: Body length: 8.2; maximum head
width: 2.7; fore wing length (including tegula): 7.6.
Colour. Similar to male, except yellow marks on
frons faint; pedicel and flagellomeres weakly orange
yellow; metasoma pale yellow ventrally. Pubescence.
Similar to male, except for T5–T6 with marginal
band of simple setae occupying whole margins.
Integument sculpture. Similar to male, but denser;
clypeus with dense coarse punctures (0.5 pd), inter-
mingled amongst sparse minute punctures (> 2 pd);
supraclypeal area with dense coarse punctures
(0.5 pd); disc of frons with dense coarse punctures
intermingled amongst fine punctures (< 0.5 pd);
antennal scrobe with dense minute punctures
(< 0.5 pd), with small smooth area above the anten-
nal socket. Metapostnotum with dense minute punc-
tures on lateral portions (1 pd), and coarser
punctures on disc (1–2 pd); transition zone between
metapostnotum and propodeum with small smooth
and shiny area; mesepisternum with sparse coarse
punctures (2 pd), and some dense punctures on
omaular area (0.5 pd). Structure. Similar to male;
lamella of pronotal collar with lateral portions
obtuse; scutellum disc convex, almost flat. Head
about 1.2¥ broader than long (2.7 : 2.2); ratio of
lower to upper interocular distance: 0.85
(0.12 : 0.14); clypeus about 2.1¥ broader than long
(1.2 : 0.5); scape: length 0.75, maximum width
0.2; length of F1–F3: 0.18, 0.12, 0.2; diameter of
F2: 0.2.
Variation: Most males have T5 with margins
completely glabrous, but a few specimens have
short marginal band occupying less than one third
of the margin laterally. The colour of sterna and
terga can vary from pale brown, almost yellow, to
dark brown.
Etymology: The species name refers to white pubes-
cence on posterolateral portion of scutellum.
Distribution: Brazil: Goiás, Maranhão, Pará (Fig. 76).
Type material: Holotype male, ‘DZUP\ 022833’
‘Coleção\ Campos Seabra’ ‘Ilha Maiandeua\

© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
418 A. J. C. AGUIAR and G. A. R. MELO

Figures 129–137. Sternum 8 (S8), male; 129: Paratetrapedia flavescens sp. nov. (Mexico, San Luis Potosi, Xilitla);
130: Paratetrapedia mexicana sp. nov. (Mexico, Jalisco, San Patricio); 131: Paratetrapedia fervida (Brazil, Espírito
Santo, Santa Teresa); 132: Paratetrapedia punctata sp. nov. (Brazil, Minas Gerais, Pratinha); 133: Paratetrapedia
flavifrons sp. nov. (Brazil, Rondônia, Vilhena); 134: Paratetrapedia romani (Brazil, Amazonas, Manaus); 135: Paratet-
rapedia tocantinensis sp. nov. (Brazil, Goiás, Campos Belos); 136: Paratetrapedia flaveola sp. nov. (Brazil, Minas
Gerais, Três Marias); 137: Paratetrapedia lineata (Brazil, Pará, Óbidos); scale bars = 0.1 mm.

Maracanã Pará\ BRAZIL ii-1955\ R. Damasceno’; Freitas&Freitas’ ‘Paratetrapedia\ sp4’; 1 female

paratypes: Goiás: 1 male (DZUP), ‘BRAZIL:GO\
Goiás, Cor\ Paciência\ A.Raw col\ 15-10-1984’ ‘Psy-
chotria’; 1 male (DZUP), ‘Goiás Velho\ C. Paciência\
31-7-82’; 1 female (DZUP), ‘Goiás Velho\ C. Paciên-
cia\ 31-7-82’ ‘sp45’; 1 female (DZUP), ‘Brazil;GO\
Ch. Veadeiros\ 05-07.12.1996\ Boaventura\
(DZUP), ‘Goiás Velho\ C. Paciência\ 31-7-82’; Mara-
nhão: 1 female (RPSP), ‘SÃO LUIS-MA.\ Brazil.
5.I.1983\ 830297’ ‘Mazucato, Aily, Camargo leg.’; 4
males (RPSP), idem except ‘830307’, ‘830301’; 1 male
(RPSP), ‘SÃO LUIS-MA.Brazil\ 23.VIII.1982\ J.M.F-
.Camargo leg.\ 621296’; 1 male (RPSP), ‘SÃO LUIS-

© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
 SYSTEMATICS OF THE GENUS PARATETRAPEDIA 419

Figures 138–146. Sternum 8 (S8), male; 138: Paratetrapedia camargoi sp. nov. (Brazil, Acre, Rio Branco); 139:
Paratetrapedia chocoensis sp. nov. (Panama, Barro Colorado); 140: Paratetrapedia flavipennis (Brazil, Amazonas,
Manaus); 141: Paratetrapedia duckei (French Guiana); 142: Paratetrapedia rufa sp. nov. (Bolivia, Santa Cruz, Santa
Cruz de La Sierra); 143: Paratetrapedia albopilosa sp. nov. (Brazil, Maranhão, São Luís); 144: Paratetrapedia
andina sp. nov. (Bolivia, Yungas, Puerto Villa); 145: Paratetrapedia atlantica sp. nov. (Brazil, Espírito Santo,
Conceição da Barra); 146: Paratetrapedia albilabris (Costa Rica, Montezuma); scale bars = 0.1 mm.

MA\ Brazil. 6.I.1983\ Mazucato, Aily\ 830487’; 2
males (RPSP), ‘SÃO LUIS-MA.\ Brazil. 5.I.1983\
830300’ ‘Mazucato, Aily, Camargo leg.’ ‘Paratetrape-
dia\ sp.\ Det. Moure 1984’; 2 males (RPSP), ‘SAO
LUIS-MA.\ Brazil. 5.I.1983\ 830303’ ‘Mazucato,
Aily\ Camargo leg.’; 1 female (LEA), ‘São Luís-MA,
Brazil\ 9-VI-2000\ Cruz e Sodré leg’ ‘Canhão Roxo\
8-9 h’ ‘Paratetrapedia sp3\ 1003’; 1 female (LEA),
‘São Luís-MA, Brazil\ 25-VII-2000\ Cruz e Sodré leg’
‘Pl. 13\ Hr.14-15’ ‘Paratetrapedia sp3\ 1124’, 4
females (LEA), idem except ‘10-VII-2000’ ‘Saco\ Hr
7-8’ ‘854’, ‘24/VI/2000’ ‘Pl. 34\ Hr. 7-8’ ‘860’, ‘13/V/

© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
420 A. J. C. AGUIAR and G. A. R. MELO

Figures 147–158. Male genital capsule, ventral and dorsal views; 147–148: Paratetrapedia lugubris (Brazil, Rondônia,
Vilhena); 149–150: Paratetrapedia connexa (Guatemala); 151–152: Paratetrapedia hyalinata sp. nov. (Brazil, Pará);
153–154: Paratetrapedia volatilis (Brazil, Paraná, Piraquara); 155–156: Paratetrapedia flavescens sp. nov. (Mexico,
San Luis Potosi, Xilitla); 157–158: Paratetrapedia moesta (Mexico, Jalisco, San Patricio); scale bars = 0.1 mm. Abbrevia-
tions: beg, expansion on basal portion of gonostylus; ea, expansion of apodeme of gonapophyses; lbg, lamellate inner lobe
of gonapophyses; lg, inner lamella of gonapophyses.

© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
 SYSTEMATICS OF THE GENUS PARATETRAPEDIA 421

Figures 159–170. Male genital capsule, ventral and dorsal views; 159–160: Paratetrapedia bifrons sp. nov. (Mexico,
Veracruz, Fortin de las Flores); 161–162: Paratetrapedia bicolor (Brazil, Espírito Santo, Conceição da Barra); 163–164:
Paratetrapedia basilaris sp. nov. (Brazil, Amazonas, Manaus); 165–166: Paratetrapedia calcarata (Costa Rica,
Puntarenas); 167–168: Paratetrapedia testacea (Brazil, Pará, Óbidos); 169–170: Paratetrapedia leucostoma (Brazil,
Maranhão, São Luís); scale bar = 0.1 mm.

© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
422 A. J. C. AGUIAR and G. A. R. MELO

Figures 171–184. Male genital capsule, ventral and dorsal views; 171–172: Paratetrapedia fervida (Brazil, Espírito
Santo, Santa Teresa); 173–174: Paratetrapedia punctata sp. nov. (Brazil, Minas Gerais, Pratinha); 175–176: Paratet-
rapedia mexicana sp. nov. (Mexico, Jalisco, San Patricio); 177–178: Paratetrapedia romani (Brazil, Amazonas,
Manaus); 179–180: Paratetrapedia tocantinensis sp. nov. (Brazil, Goiás, Campos Belos); 181–182: Paratetrapedia
flavipennis (Brazil, Amazonas, Manaus); 183–184: Paratetrapedia lineata (Brazil, Pará, Óbidos); scale bars = 0.1 mm.
Abbreviation: lbg, lamellate inner lobe of gonapophyses.

2000’ ‘Pl. 34\ Hr. 14-15’ ‘826’, ‘24/V/2000’ ‘Saco/Hr. 7-8’
‘1069’; 1 female (LEA), ‘Turu, S. Luís Ma, BR\ 07/VI/
03\ ARAGÃO-LEG’ ‘Paratetrapedia\ nasuta (F)\
07:00-08:00\ 1261’, 1 male (LEA), idem except
‘12/IV/03’ ‘(M)\ 06:00-07:00\ 1218’; 1 female (LEA),
‘Alcântara-Ma-Br\ 30/V/1993\ Araújo & Gonçalves’
‘Pl. n.020\ Hr. 9-10’ ‘Paratetrapedia\ sp. (1)’; Pará, 1
female (DZUP), ‘Brazil: Pará\ Belém: Cotijuba\
07.IV.2002\ A.A.S.Matos’ ‘Diag 2’; 1 female (DZUP),
‘Brazil: PA, Bragança\ Vivenda Vitória\ 24.VII.2002\
Leg: V.F.O.Raiol’ ‘Diag 265’; 1 female (MNRJ),
‘Utinga, Belém\ Pará, Brazil’ ‘Arlindo\ X-71’; 1 male,
‘DZUP\ 022993’ ‘BELEM\ Pará BRAZIL\ XII-1954\
R. Damasceno’.

© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
 SYSTEMATICS OF THE GENUS PARATETRAPEDIA
PARATETRAPEDIA ANDINA SP. NOV.
(FIGS 76, 108, 144, 207, 208)
Comments and diagnosis: Paratetrapedia andina is
very similar to P. atlantica, both having brown
plumose hairs on the posterolateral margins of scutel-
lum and very faint yellow stripes on mesoscutum. It
differs from this species by male T5 with marginal
band occupying about one-quarter of margin laterally.
Paratetrapedia andina is known only from three male
specimens.
Distribution: Bolivia; Ecuador (Fig. 76).
Description
Holotype male: Body length: 8.0; maximum head
width: 2.5; fore wing length (including tegula): 8.1.
Colour. Integument mostly dark brown; mandible
pale yellow with apex black; labrum pale yellow;
clypeus with two pale yellow spots on margins later-
ally, contacted by a thin yellow stripe on lower margin
of clypeus; supraclypeal area with yellow spot on disc;
lower paraocular area with wide yellow stripe; disc of
frons with thin and short yellow stripe; scape, pedicel,
and flagellomeres reddish brown. Mesoscutum with
two thin faint yellow stripes on disc and lateral
margins. Fore wing membrane brown infumate with
dark brown microtrichia; veins and pterostigma
yellow; marginal cell with membrane slightly yellow
infumate. Tibial spurs whitish. Pubescence. Paraocu-
lar area, genal area, mesepisternum, propodeum, and
sterna with pubescence mostly pale white; remaining
body pubescence brown. Scape with long white hairs
on inner surface, about 0.2 mm in length. Mesoscu-
tum and scutellum with dense plumose pale brown
pubescence; posterolateral portions of scutellum with
plumose brown hairs; metapostnotum with short
plumose hairs, about 0.08 mm in length. T4–T5 with
marginal hairs band occupying less than one-quarter
of margin laterally; margins of T6 completely gla-
brous. S2 with two small rows of simple erect setae on
mid portion of margin; S3 deeply concave in ‘U’ shape,
covered by dense short plumose pubescence; S4 and
S5 typical of genus; S6 with two rows of stout plumose
hairs on margins laterally, and short plumose hairs
on margin of apex. Integument sculpture. Clypeus
with shallow coarse punctures (1 pd), intermingled
amongst sparse minute punctures (> 2 pd); supracly-
peal area with coarse punctures, denser on lateral
portions (< 1 pd) and sparser on disc (2 pd); disc of
frons with coarse punctures (1–2 pd); antennal scrobe
with dense fine punctures (< 1 pd). Mesoscutum and
scutellum with dense minute punctures (< 1 pd),
intermingled amongst sparse coarser punctures
(2 pd); metapostnotum with fine dense punctures
© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
423
(0.5 pd), and dense minute on lateral margins
(< 0.5 pd); transition zone of metapostnotum and pro-
podeum with smooth area; mesepisternum with
coarse punctures, mostly sparse (> 2 pd). Structure.
Lamella of pronotal collar with lateral portions
obtuse. Scutellum conspicuously convex. Head about
1.2¥ broader than long (2.5 : 2.1); ratio of lower
interocular distance to upper interocular distance:
0.82 (1.17 : 1.42); clypeus about 1.7¥ broader than
long (1.12 : 0.65); scape: length 0.65, maximum width
0.18; length of F1–F3: 0.2, 0.15, 0.22; diameter of F2:
0.17.
Variation: The specimen from Ecuador has the yellow
marks on face very reduced, mandible, labrum, and
genal area mostly dark brown, clypeus with only two
small yellow spots on lateral margins.
Etymology: The species name refers to the Andean
cordillera.
Type material: Holotype male, ‘DZUP\ 022839’
‘Nigrillani-2500 m\ Enero 1950\ G. Williner leg.\ N.
Yungas-Bolívia’; paratypes: BOLIVIA: 1 male,
‘DZUP\ 022916’ ‘Puerto Villa 1200 m\ Yungas
Bolívia\ 13-20/XII/55 Peña’; 1 male (ZSMC),
‘BOLIVIA\ Yungas, Coroico\ 1900 m\ 19.5.975\ leg.
W. Forster\ Zoolog.\ Staatssig’; ECUADOR: 1 male
(SEMC), ‘ECUADOR: Sucumbios\ Sacha Lodge,
0.5°S\ 76.5°W, 270 m, 12-22-III\ 1994, Hibbs, ex:
malaise’.
UNPLACED SPECIES (WITHOUT DEFINED
SPECIES GROUP)
PARATETRAPEDIA MEXICANA SP. NOV.
(FIGS 44, 71, 84, 100, 130, 175, 176)
Comments and diagnosis: Paratetrapedia mexicana is
distinguished by female prementum with numerous
long curved setae, male S2 with continuous row of
setae and posterior margin of S3 widely concave and
covered by short plumose hairs, male and female
T4–T6 with marginal hair band occupying whole
margins, male hind basitarsus with one conspicuous
tooth on proximal third of anterior margin and meta-
postnotum with coarse punctures intermingled
amongst fine punctures.
Description
Holotype male: Body length: 7.8; maximum head
width: 2.75; fore wing length (including tegula): 8.9.
Colour. Integument mostly dark brown; mandible
pale yellow with apex black; labrum, clypeus, and
supraclypeal area pale yellow; paraocular area with
wide yellow stripe; scape pale yellow; pedicel and
424 A. J. C. AGUIAR and G. A. R. MELO

© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
 SYSTEMATICS OF THE GENUS PARATETRAPEDIA 425

Figures 185–196. Male genital capsule, ventral and dorsal views; 185–186: Paratetrapedia duckei (French Guiana);
187–188: Paratetrapedia flavifrons sp. nov. (Brazil, Rondônia, Vilhena); 189–190: Paratetrapedia rufa sp. nov.
(Bolivia, Santa Cruz, Santa Cruz de La Sierra); 191–192: Paratetrapedia flaveola sp. nov. (Brazil, Minas Gerais, Três
Marias); 193–194: Paratetrapedia camargoi sp. nov. (Brazil, Acre, Rio Branco); 195–196: Paratetrapedia atlantica
sp. nov. (Brazil, Espírito Santo, Conceição da Barra); scale bar = 0.1 mm.
䉳

flagellomeres pale brown (Fig. 44). Wing membrane
dark brown infumate; apical third of fore wing mem-
brane slightly pale; microtrichia dark brown; veins
and pterostigma dark brown. Tibial spurs white;
mediotarsus, distitarsus, and claws pale yellow.
Pubescence. Mostly dark brown, almost black. Scape
with long dark brown hairs on inner face, about
0.01 mm in length. Mesoscutum and scutellum with
short dark brown plumose pubescence; metapostno-
tum with short plumose hairs, about 0.06 mm in
length. T4–T6 with marginal band of dense plumose
dark brown hairs throughout its margin; margin of S2
with continuous row of simple erect black setae; S3
deeply concave in ‘U’ shape, covered by dense short
plumose pubescence (Fig. 21); S4 and S5 typical of
genus; S6 with sparse pubescence on disc and two
conspicuous rows of stout plumose hairs on margins
laterally, apex bordered by short plumose hairs.
Pubescence of legs mostly dark brown; mid basitarsus
with dense dark brown pubescence. Integument
sculpture. Clypeus with dense coarse punctures
(1 pd); supraclypeal area with coarse punctures
(1 pd), with smooth and shiny integument amongst
punctures; frons with sparse coarse punctures
(2–3 pd); antennal scrobe completely smooth. Mesos-
cutum with fine dense punctures (< 0.5 pd) inter-
mingled amongst sparse coarse punctures (> 2 pd);
metapostnotum with sparse fine punctures (2 pd);
transition zone between metapostnotum and propo-
deum smooth and shiny; mesepisternum with sparse
fine punctures (3 pd). Structure. Lamella of pronotal
collar acute throughout its length. Disc of scutellum
weakly convex, almost flat. Hind basitarsus with
acute tooth on anterior margin. Head about 1.17¥
broader than long (2.75 : 2.35); ratio of lower to upper
interocular distance: 0.87 (1.26 : 1.44); clypeus about
1.7¥ broader than long (1.15 : 0.65); scape, length 0.6;
maximum width 0.2; length of F1–F3: 0.2, 0.17, 0.25;
diameter of F2: 0.22.
Paratype female: Body length: 8.7; maximum head
width: 2.9; fore wing length (including tegula): 8.5.
Colour. Similar to male, except for absence of pale
yellow marks on face, with faint yellow mark on
lower paraocular area; tarsomeres 2–4 orange
yellow. Pubescence. Similar to male; except for con-
spicuously longer simple setae on lateral portions of
scutellum, c. 0.42 mm in length; metapostnotum
with short plumose setae, c. 0.16 mm in length;
T4–T6 with marginal hair band occupying whole
margin. Integument sculpture. Similar to male
except for frons with sparse fine punctures (2 pd);
metapostnotum with sparse fine punctures (2 pd)
intermingled amongst sparse minute punctures
(2 pd). Structure. Lamella of pronotal collar with
lateral portions conspicuous obtuse. Scutellum disc
weakly convex, almost flat. Head about 1.2¥ broader
than long (2.9 : 2.25); ratio of lower to upper
interocular distance: 0.84 (1.37 : 1.62); clypeus about
2.1¥ broader than long (1.27 : 0.6); scape: length 0.8,
maximum width 0.17; length of F1–F3: 0.25, 0.12,
0.17; diameter of F2: 0.22.
Variation: The wing membrane can be completely
brown infumate or with apical portion pale infumate.
Etymology: The species name refers to the country of
origin, Mexico.
Distribution: Mexico: Jalisco, Morelos (Fig. 71).
Type material: Holotype male (SEMC), ‘MEXICO,
Morelos\ 3.8mi. W Yautepec\ 17August 1962 3800’\
(E. Ordway)’. Paratypes: MEXICO: Jalisco: 1 male
(BLCU), ‘MEXICO Jalisco\ Chamela Res. Sta\ 6
August 1986\ M. Sanchez-M.T.’ ‘NativeBeeSurvey\
USDA Logan Utah\ BBSL517252’; 1 female (BLCU),
‘MEXICO Jalisco\ Chamela 1/8-X-\ 85 F.D. Parker\
T.L. Griswold’ ‘NativeBeeSurvey\ USDA Logan
Utah\ BBSL517251’; 1 female (BLCU), ‘MEXICO
Jalisco\ Chamela (Est.\ Biol.) 10/14 July\ 1989 T.
Griswold’ ‘NativeBeeSurvey\ USDA Logan Utah\
BBSL517250’; 1 female (SEMC), ‘Mex. Jal. Chamela\
7-IX-1985\ R. Ayala\ RA200’ ‘Paratetrapedia\ F sp3\
det. Ayala 87’; 1 female (SEMC), ‘MEXICO, Jalisco,
Chamela\ Fecha 13-IX-1986\ Col. R. Ayala A178’
‘Paratetrapedia\ F sp3\ det. Ayala 87’; 1 female
(SEMC), ‘MEXICO Guerrero\ 12.7mi. NE. Iguala\ 1
August 1969 5200’\ Dwight Kamm’; 1 female (CUIC),
‘MEXICO. Estado de Jalisco\ Reserva Biosfera
Chamela-Cuixmala\ Estación de Biologia Chamela\
Vereda Ardila. 06/ix/2004, – 10:30h\ 19°30.280′N,
105°02.224′W, 43 m.\ E.A.B. Almeida,leg.’ ‘Paratetra-
pedia\ det. J.S. Ascher’; Morelos: 1 female (SEMC),
‘MEXICO Morelos\ 7mi. N. Yautepec\ 4000’, 18
August 1962\ U. Kans. MexExped.’; 1 female

© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
426 A. J. C. AGUIAR and G. A. R. MELO

© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
 SYSTEMATICS OF THE GENUS PARATETRAPEDIA 427

Figures 197–208. Male genital capsule, ventral and dorsal views; 197–198: Paratetrapedia scaposa sp. nov. (Brazil,
Acre, Cruzeiro do Sul); 199–200: Paratetrapedia vogeli sp. nov. (Peru, Cuzco, Quincemil); 201–202: Paratetrapedia
chocoensis sp. nov. (Panama, Barro Colorado Island); 203–204: Paratetrapedia albopilosa sp. nov. (Brazil, Mara-
nhão, São Luís); 205–206: Paratetrapedia albilabris (Costa Rica, Montezuma); 207–208: Paratetrapedia andina sp.
nov. (Bolivia, Yungas, Puerto Villa); scale bar = 0.1 mm.
䉳

(SEMC), ‘MEXICO Morelos\ 6.7 mi. S Yautepec\ 29
July 1963\ Naumann & Willis’.
PARATETRAPEDIA FLAVESCENS SP. NOV.
(FIGS 42, 71, 113, 129, 156)
Comments and diagnosis: Paratetrapedia flavescens is
remarkable for its predominantly bright yellow integu-
ment. It is the only species of Paratetrapedia with
yellow integument found in Mexico. Its male has a
unique combination of characters that does not fit in
any of the species groups here recognized. The male
possesses the margin of S2 with row of setae continu-
ous, S3 with wide concave area on posterior margin
covered by dense short plumose pubescence, and hind
basitarsus without tooth on anterior margin. The
female is characterized by a yellow integument, mostly
smooth and convex supraclypeal area, acute lamella of
pronotal collar, and metapostnotum with minute punc-
tures intermingled amongst coarser punctures.
Description
Holotype male: Body length: 8.6; maximum head
width: 3.1; fore wing length (including tegula): 9.3.
Colour. Integument mostly yellow; upper paraocular
area, lateral portion of frons and vertex black; scape
yellow, pedicel and flagellomeres orange yellow
(Fig. 42). Pronotum with small dark brown mark on
mid portion of anterior surface; mesoscutum black
with two yellow stripes on disc and lateral margins;
axilla black laterally. Terga yellow with marginal zone
black; S2 with mid portion black. Wing membrane
yellow infumate, with yellow microtrichia; apical
portion of fore wing with brown microtrichia; veins
yellow and pterostigma orange yellow; tibial spurs
pale yellow. Pubescence. Mostly golden yellow, except
margins of T3–T6 with dark brown hairs. Scape with
pale yellow long hairs on inner surface, about
0.14 mm in length. Mesoscutum and scutellum with
short plumose golden yellow pubescence; metapostno-
tum with short plumose yellow hairs, about 0.1 mm in
length. T4–T6 with marginal bands of dark brown
plumose hairs throughout its margins; T6 with
slightly pale brown hairs. S2 with row of setae con-
tinuous; S3 deeply concave in ‘U’ shape, covered by
dense short plumose pubescence; S4 and S5 typical of
the genus; S6 with two small rows of plumose hairs
on margins laterally and short plumose hairs on
margin of apex. Integument sculpture. Clypeus with
shallow coarse punctures (0.5–2 pd), intermingled
amongst sparse minute punctures (5 pd); supracly-
peal area with sparse coarse punctures (0.5–3 pd);
frons with coarse punctures, mostly sparse (1–4 pd);
mid line of frons weakly sulcated. Mesoscutum with
dense fine punctures (< 0.5 pd), intermingled amongst
sparse coarse punctures (> 2 pd); scutellum with
dense fine punctures (1 pd), intermingled amongst
sparse coarse punctures (> 2 pd); metapostnotum
with sparse fine punctures (> 2 pd); mesepisternum
with sparse fine punctures (3 pd). Structure. Disc of
frons slightly biconvex; lamella of pronotal collar
acute throughout its length. Scutellum disc weakly
convex, almost flat, mid line slightly sulcate. Head
about 1.3¥ broader than long (3.1 : 2.4); ratio of lower
to upper interocular distance: 0.8 (1.3 : 1.6); clypeus
about 1.6¥ broader than long (1.25 : 0.75); scape:
length 0.8, maximum width 0.22; length of F1–F3:
0.22: 0.16: 0.2; diameter of F2: 0.2.
Paratype female: Body length: 10.1; maximum head
width: 3.3; fore wing length (including tegula): 9.7;
head about 1.3¥ broader than long; ratio of lower to
upper interocular distance: 1.16; clypeus about 2.1¥
broader than long; scape: length 0.95, maximum
width: 0.25; length of F1–F3: 0.28; 0.1; 0.2; diameter
of F2: 0.24. Colour. Similar to male, except for hind
leg weakly dark brown. Pubescence. Similar to male;
mesoscutum and scutellum with sparse simple erect
setae, about 0.06 and 0.2 mm in length, respectively.
Terga pubescence similar to male, except for marginal
band on T4–T6 with simple hairs. Integument sculp-
ture. Similar to male, except for clypeus with only
coarse punctures; supraclypeal area with disc smooth,
with sparse punctures on lateral portions; antennal
scrobe with dense minute punctures (1 pd); metapost-
notum with sparse fine and coarse punctures
intermingled (> 2 pd); mesepisternum with coarse
punctures, mostly sparse (2 pd). Structure. Similar to
male. Disc of scutellum weakly convex, almost flat.
Head about 1.3¥ broader than long (3.3: 2.5); ratio of
lower to upper interocular distance: 1.16 (1.55 : 1.8);
clypeus about 2.1¥ broader than long (1.4 : 0.65);
scape: length 0.95; maximum width 0.25; length of
F1–F3: 0.28: 0.1: 0.2; diameter of F2: 0.24.
Variation: There is some variation in the extension of
the black maculae, mainly on frons and tergal
margins. The female from Costa Rica differs from

© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
428 A. J. C. AGUIAR and G. A. R. MELO
those from Mexico in having a mostly orange yellow
integument, frons with thin yellow stripe on mid line,
and tergal margins darker. It might represent a sepa-
rate, closely related species.
Etymology: The species name refers to its yellow
integument.
Distribution: Costa Rica; Mexico (Fig. 71).
Type material: Holotype male (AMNH), ‘MEXICO:
State of San\ Luis, Potosi, Hotel\ Covadonga c.
Valles\ 6-VI-1974, C. Porter\ & C. Calmbacher’;
paratypes: COSTA RICA: 1 female (SEMC), ‘COSTA
RICA San José Prov.\ Rio Damitas, 14.5 km.\ N.
Puerto Quepos\ 200 m., August 1962\ (A. Wille &
C.D. Michener)’; MEXICO: 2 males (SEMC),
‘MEXICO: San Luis\ Potosi, Xilitla, 1km E\ Xilitla,
at river, 600 m\ 7 July 1990, I. Yarom’; 3 males
(SEMC), ‘MEXICO: San\ Luis Potosi, Xilitla\ 400 m,
7 July 1990\ R.L. Minckley\ ex., on Malvaceae’; 1
male (SEMC), ‘MEXICO: Hidalgo, 700 m\ 25 km S
Tamazunchale\ 22-V-1989. Yanega’ ‘Paratetrapedia\
det. Douglas Yanega’; 1 female (SEMC), ‘MEXICO:\
Xilitla, Tlamaya\ Falls, 1000 m\ 15August 1987\ D.
Yanega’ ‘San Luis Potosí’; 1 female (SEMC),
‘MEXICO: Tamps, 80 km\ S. CD. Victoria, Ruta\ 85,
CA. La Gloria\ 23,25-VI-81.B.Miller\ C. Porter, L.
Stange’; 1 female (DZUP), ‘MEXICO: Tam\ El
Ensino, 250 m\ 4-13.viii.1988\ V.O. Beker col.’; 1
female (SEMC), ‘25-Ago-1990\ Xicotepec\ Puebla\ R.
Paz R.’.
NASUTOPEDIA GEN. NOV.
Type species: Tetrapedia nigripes Friese, 1899
Comments and diagnosis: The recognition of the
genus Nasutopedia is supported by at least four char-
acters (Figs 209, 210), by the distinct morphology of
male terminalia (Figs 214–218), and by its biogeo-
graphical distribution. In Nasutopedia, as in Paratet-
rapedia, Tropidopedia, and Lophopedia, the male
pygidial plate is reduced to a pygidial process, the
pronotal collar is lamellate, and the female pygidial
plate is differentiated into a basal and an apical
portion. Nasutopedia, however, exhibits a set of
unique characters, which allows its easy identifica-
tion. In addition, the male terminalia of Nasutopedia
nigripes and Nasutopedia micheneri are quite similar
to each other but very distinct from the remaining
species of Tapinotaspidini.
The following characters are common to the species
of Nasutopedia gen. nov. here recognized: females
with two large preapical teeth in the mandible (char-
acter 4:1); males with post-ocellar carina acute and
© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
very short, restricted to post-ocellar portion of vertex
(character 12:0); females with post-ocular carina nar-
rowly rounded (Fig. 210; character 11:0); basitibial
plate of female long and slightly reniform (character
31:2); metanotum weakly convex, and uniformly
broad along its entire length; notaulus inconspicuous
[completely absent in Nasutopedia rasmusseni; char-
acter 16:0)].
The shape of the preapical teeth in the female
mandible is very similar to the condition exhibited by
species of the lugubris group in Paratetrapedia, with
two large preapical teeth widely separate from the
apical tooth. Tridentate mandibles are also present in
females of Lophopedia pygmaea, but the shape of its
teeth is quite distinct. In L. pygmaea, the second
preapical tooth is smaller than the first tooth, and
they are placed very close to each other; in the
remaining species of Lophopedia, only a single and
large preapical tooth, whose apex is rectangular, is
present (Aguiar, 2009).
Nasutopedia has in common with Paratetrapedia
the female fore basitarsus without an apical projec-
tion; an acute male pygidial process; a reduced male
basitibial plate, with only a short carina on its apical
one-third; margins of basal portion of the female
pygidial plate almost carinate and converging in an
obtuse angle; setae of brush on female mid tibia with
an acute apex and flat surface; margin of male S6
with tufts of plumose setae laterally; notaulus very
weak, almost absent; row of setae on margin of male
S2 not differentiated.
Description: Body length about 7.0 to 10.0 mm;
integument varying from dark brown to black,
usually with white or yellow marks on face; wing
membrane dark brown or pale yellow infumate; terga
orange brown or black. Pubescence mainly sparse and
short; paraocular area with pubescence mostly
simple; vertex and upper paraocular area with erect
stout setae; mesoscutum and scutellum with dense
short velvet pubescence, intermingled with numerous
long setae. Terga smooth and shiny or with dense fine
punctures. Posterolateral margins of terga with
bands of hairs; in females, composed mainly of simple
hairs, in males, composed of plumose hairs. In males,
T1–T4 with marginal hair band occupying less than
one third of margin laterally; T5 with marginal band
almost complete to complete; T6 with marginal hair
band usually complete, except in N. rasmusseni in
which the margin is glabrous. In females, marginal
hair band absent on T1 (except N. nigripes); on
T2–T4, marginal hair bands occupying less than one
third of margin laterally, almost absent; on T5–T6,
complete. Males with pregradular area of S2–S6
covered by dense short plumose pubescence (except
for N. micheneri, in which the pregradular surfaces of
 SYSTEMATICS OF THE GENUS PARATETRAPEDIA 429

Figure 209. Strict consensus of four most parsimonious cladograms with equal weighting, for species of Paratetrapedia
(fast optimization; length: 126 steps; consistency index: 54; retention index: 83); solid circles represent unique transfor-
mations and open circles multiple transformations.

© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
430 A. J. C. AGUIAR and G. A. R. MELO

Figure 210. Most parsimonious cladogram with implied weighting for species of Paratetrapedia (k, concavity con-
stant = 3), fast optimization (total fitness: 455.2; length: 123; consistency index: 56; retention index: 84). Solid circles
represent unique transformations and open circles multiple transformations. Main biogeographical components indicated
following species name; components according to Camargo & Pedro (2003), except for the Cerrado and Choco. Abbrevia-
tions: Atl, Atlantic; CA, Central America; NAm, northern Amazon; Neot, Neotropical region; SEAm, south-eastern
Amazon; SWAm, south-western Amazon. An asterisk indicates that the species also has extralimital records.

S2–S3 are glabrous); margin of S2 and S3 with erect
simple setae forming a single continuous row; S4 with
band of decumbent long plumose hairs, forming long
converging fringes laterally; S5 with mid portion
mostly glabrous with erect stout plumose hairs later-
ally; S6 with short stout plumose hairs on apical
portion, with small tufts of setae on margin laterally.
Males and females with differentiated hairs for oil
and pollen collecting: fore basitarsus with comb of
simple setae on outer margin and plumose hairs on

© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
 SYSTEMATICS OF THE GENUS PARATETRAPEDIA
Figures 211–213. Head; 211: Nasutopedia nigripes (Friese), female, dorsal view; 212: Nasutopedia nigripes, male
(Colombia, Risaralda); 213: Nasutopedia micheneri, male, holotype; scale bar = 0.5 mm.
inner concave surface. Females with a comb of
stout spatulate setae with acute apex on inner
surface of mid tibia; basitarsus of mid and hind
legs with numerous short spatulate setae on inner
surface. Mandible with two preapical teeth in
females; males with only one preapical tooth.
Post-ocellar carina narrowly rounded, almost incon-
spicuous (Fig. 210), not reaching the post-ocular
portion of vertex. Pronotal collar lamellate dorsally,
with lateral sections low, confluent with lateral,
vertical portions (Fig. 210), or high and closed as a
gutter on N. rasmusseni; lamella of pronotal collar
sharp, with lateral portions obtuse. Notaulus incon-
spicuous, not sulcated and indicated by a narrow
smooth area; on N. rasmusseni completely absent.
Pygidial plate of females differentiated in apical and
basal portions; basal portion with margins almost
carinate and converging in obtuse angle; male
without pygidial plate, with only an acute pygidial
process. Basitibial plate of females with smooth gla-
brous margins and central portion convex, curved,
weakly reniform; basitibial plate of males mostly
inconspicuous with only a short carina along apical
one-third; inner hind tibial spur serrate, slightly dis-
tinct from outer spur. Male head about 1.2¥ wider
than long; eyes weakly divergent toward vertex, lower
interocular distance about 0.8¥ upper interocular dis-
tance. Male terminalia: lamellate apex of S7 densely
pilose without a conspicuous inner lobe; lamellate
apex of S8 long and narrow; dorsal bridge
of gonocoxite long and broad; parapenial lobe with
stout setae and weakly curved toward mid portion of
gonocoxite.
Etymology: The generic name refers to the strongly
convex clypeus of the males.
© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
431
NASUTOPEDIA NIGRIPES (FRIESE, 1899) COMB. NOV.
(FIGS 211, 212, 219)
Tetrapedia nigripes Friese 1899: 287; holotype male,
Venezuela (ZMB, examined).
Tetrapedia nigripes; Ducke (1910b): 369. Strand
(1910): 460. Friese (1916): 335. Friese (1923): 4.
Chalepogenus nigripes; Cockerell (1923a): 450.
Chalepogenoides nigripes; Michener (1942): 281.
Paratetrapedia nigripes; Aguiar (2007): 625.
Paratetrapedia (Paratetrapedia) nigripes; Rasmussen
& Ascher (2008): 77.
Comments and diagnosis: Nasutopedia nigripes
differs from the remaining species by the dorsal
surface of pronotal collar with opened, not raised
lateral portions. Males of N. nigripes are distin-
guished by the integument mostly black, except for
white labrum; surface of S2–S3 with dense fine short
simple setae; T5 with almost complete marginal hair
band of black plumose hairs; T6 with complete mar-
ginal hair band of black plumose hairs throughout its
margin; T1–T3 with marginal hair band of plumose
hairs occupying less than one third of margin later-
ally; T4 with marginal band of plumose hairs occu-
pying about one third of margin laterally; T1 with
dense minute punctures with very short pubescence;
marginal zone of T2–T3 with sparse minute punc-
tures, tergal disc smooth; lamella of pronotal collar
with lateral portions low, obtuse, almost closed;
sulcus on notauli thin, conspicuous.
The female is mainly identified by integument com-
pletely black to reddish brown, only terga and tarsi
weakly pale brown; notaulus very weakly sulcated;
lamella of pronotal collar with lateral portions low,
opened, without signs of lateral lamella; frons with
coarser punctures on disc (1 pd) and dense minute
432 A. J. C. AGUIAR and G. A. R. MELO
punctures (< 1 pd); terga with numerous minute
punctures on entire marginal zone, with dense
minute hairs; hind tibial scopa mostly dark brown
with distal portion pale yellow, almost white, and
hind basitarsus with pale brown pubescence.
Redescription of holotype (male): Body length: 7.9;
wing length (with tegula): 8.2; head: width, 2.56,
length, 2.12; clypeus: length, 0.68, width, 1.14; lower
interocular distance: 1.18; upper interocular distance:
1.41; scape: length, 0.61, width, 0.19; F1–F3, length:
0.25, 0.13, 0.21; F2 diameter: 0.23. Colour. Mostly
black; terga, legs, and scape reddish brown; mandible
pale yellow with apex black; labrum pale yellow;
clypeus with lower margin pale yellow; S4–S5 pale
yellow, almost white; mediotarsi and distitarsi of mid
and hind legs pale yellow; tibial spurs dark brown.
Wing membrane brown infumate; veins and
pterostigma yellow; veins and membrane with dense
dark brown microtrichiae. Integument sculpture.
Clypeus and supraclypeal area with dense coarse
punctures (< 0.5 pd); disc of supraclypeal area
smooth. Frons with coarse punctures (< 0.5–1 pd),
more sparse on upper portion of disc of frons.
Paraocular area with sparse fine punctures (> 2 pd);
antennal scrobe with dense fine punctures (c. 1 pd);
vertex with dense minute punctures. Mesoscutum
with minute punctures (1–3 pd), intermingled
amongst few coarser punctures on disc (> 4 pd).
Scutellum with dense fine punctures (1–2 pd); disc
with two smooth areas on disc laterally. Mesepister-
num with sparse fine punctures (> 3 pd); metapostno-
tum with sparse fine punctures (> 4 pd); propodeum
with sparse fine punctures (> 3 pd); terga mostly
smooth, with sparse fine punctures laterally (> 3 pd).
S1–S3 with fine punctures (c. 1 pd); on S1–S2, dense
(c. 1 pd); on S3, sparse (> 2 pd). Pubescence. Mostly
dark brown; except lower margin of mandible,
labrum, lower margin of clypeus, genal area, and
short plumose hairs on mesepisternum ventrally
mostly pale yellow. Hind tibial scopa mostly dark
brown on proximal third and white on distal third.
Lower margin of proximal portion of mandible with
long simple setae, longer as the width of mandibular
base; labrum with scarce pubescence, with short
simple and plumose hairs intermingled; clypeus with
decumbent short simple setae on lower portion and
erect simple sparse setae on lateral and upper por-
tions; paraocular area and vertex with sparse erect
simple setae (c. 0.8 ¥ F2 diameter); vertex and genal
area with dense fine plumose pale brown hairs; post-
ocellar portion of vertex with erect simple and
plumose setae, c. 0.8–1.2 ¥ F2 diameter. Mesoscutum
with dense short plumose pale brown hairs, inter-
mingled amongst sparse long simple stout setae, c.
0.7 ¥ F2 diameter; scutellum with numerous stout
© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
simple long setae (c. 0.5–1.8 ¥ F2 diameter) inter-
mingled amongst plumose hairs on posterior margins.
Mesepisternum laterally with numerous dark brown
long simple hairs, c. 1.2 ¥ F2 diameter; on lower
portion with short plumose pale white hairs inter-
mingled. Metapostnotum with numerous short
plumose hairs, c. 0.5 ¥ F2 diameter. Marginal hair
band of plumose hairs on terga: T1, almost incon-
spicuous, with few hairs on margin laterally; T2–T3
with dense band of pale brown plumose hairs, occu-
pying less than one-quarter of margin laterally; T4
with marginal hair band of dark brown hairs, occu-
pying about one third of margin laterally; T5 with
marginal hair band almost complete, with one short
interval glabrous on mid portion; T6 with marginal
hair band complete. S1–S3 with dense short decum-
bent simple hairs covering the surface; S2–S3 with
long simple setae on margin forming a continuous
row; S4 with dense band of long plumose hairs,
forming long curved fringes laterally; S5 mostly gla-
brous, with few short plumose hairs laterally; S6 with
numerous simple setae on mid line of apical portion,
with apex curved posteriorly, and two small rows of
setae oblique to margin laterally. Structure. Mandible
with one pre-apical tooth; clypeus strongly convex;
lower margin of supraclypeal area with short sulcus;
mid line of frons weakly sulcate, very narrow; upper
paraocular area weakly convex, almost flat; post-
ocellar carina short and weak, almost absent,
restricted to the post-ocellar portion of vertex; genal
area declivous, weakly convex; pronotal collar with
upper portion conspicuous lamellate, forming an
acute convex lamella; dorsal lamella of pronotal collar
with mid portion sinuous; dorsal surface of pronotal
collar concave, with lateral portions opened, not
raised; notauli weakly sulcate on mesoscutum, almost
inconspicuous; scutellum biconvex, with mid line con-
spicuously concave; T7 with only a short thin lamel-
late pygidial process, pygidial plate absent; basitibial
plate mostly inconspicuous, marked only on lower
third portion.
Female: (COLOMBIA: Prov. Valle\summit. W. of
Cali). Body length: 10.0; wing length (with tegula):
10.0; head: width, 2.63, length, 1.93; clypeus: length,
6.69, width, 1.20; lower interocular distance: 1.18;
upper interocular distance: 1.51; scape: length, 0.74,
width, 0.15; F1–F3, length: 0.19, 0.11, 0.19; F2 diam-
eter: 2.16. Colour. Mostly black; terga, legs, and scape
reddish brown; basitarsus, mediotarsi, and distitarsi
of hind legs pale yellow; tibial spurs dark brown.
Wing membrane dark brown infumate; veins and
pterostigma yellow; veins and membrane with dense
dark brown microtrichiae. Integumental sculpture.
Clypeus and supraclypeal area with dense coarse
punctures (< 0.5 pd). Disc of frons with dense fine
 SYSTEMATICS OF THE GENUS PARATETRAPEDIA
punctures (< 0.5 pd), intermingled amongst sparse
coarser punctures (c. 1–3 pd). Paraocular area with
sparse fine punctures (> 2 pd); antennal scrobe with
dense fine punctures (c. 1 pd); vertex with dense
minute punctures. Mesoscutum with dense minute
punctures (0.5–1 pd), intermingled amongst few
coarser punctures on disc (> 4 pd). Scutellum with
dense fine punctures (0.5–2 pd). Mesepisternum with
sparse fine punctures (> 3 pd); metapostnotum with
numerous minute punctures (c. 3 pd); propodeum
with numerous minute punctures intermingled
amongst sparse coarse punctures. T1 with minute
punctures (c. 1–2 pd); T2–T4 with numerous minute
punctures (c. 2–3 pd), with marginal zone completely
smooth. T5 and T6 with surface mostly undercovered
by the remaining terga. S1–S2, S4–S5 with numerous
fine punctures on pre-marginal zone; S3 mostly
smooth, with sparse punctures on posterior margin.
Pubescence. Mostly dark brown; except lower margin
of mandible, labrum, and clypeus mostly reddish
yellow. Hind tibial scopa mostly dark brown on proxi-
mal third and white on distal third. Lower margin of
proximal portion of mandible with long simple setae,
longer than width of mandibular base; labrum with
numerous setae on lower margin; clypeus with erect
simple sparse setae on lateral and upper portions;
paraocular area and post-ocellar portion of vertex
with sparse erect simple setae (c. 1.0 ¥ F2 diameter);
vertex and genal area with dense fine plumose hairs.
Mesoscutum with dense short plumose dark brown
hairs, intermingled amongst sparse long plumose
stout setae, c. 0.5 ¥ F2 diameter; scutellum with stout
simple long setae (c. 1.5–2.5 ¥ F2 diameter) inter-
mingled amongst dense plumose hairs on posterolat-
eral portions. Mesepisternum laterally with
numerous dark brown long simple hairs, c. 1.3–
2.0 ¥ F2 diameter. Metapostnotum with numerous
very short hairs, < 0.2 ¥ F2 diameter. Marginal band
of hairs on terga: T1–T4, almost inconspicuous, with
few hairs on margin laterally, occupying less than
one-quarter of margin laterally; T5 with marginal
hair band of dark brown hairs, occupying whole
margin. S1–S2, S4, and S5 with dense short decum-
bent simple hairs covering the surface; S3 with sparse
simple setae; S4 with dense band of long plumose
hairs on posterior margin. Structure. Mandible with
two pre-apical teeth; clypeus weakly convex; lower
margin of supraclypeal area with narrow sulcus; frons
convex; upper paraocular area weakly convex, almost
flat; post-ocellar carina short and weak, almost
absent, restricted to the post-ocellar portion of vertex;
genal area declivous, weakly convex; pronotal collar
with upper portion conspicuous lamellate, forming an
acute convex lamella; lamella of pronotal collar with
mid portion not sinuous; lateral portions of pronotal
collar opened; notauli weakly sulcate on mesoscu-
© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
433
tum, almost inconspicuous; scutellum homogeneous
convex, without mid line concave.
Variation: The marginal hair band on male T5 can be
complete or on about one third of margin laterally.
Distribution: Colombia, Venezuela (above 1800 m alti-
tude; Fig. 219).
Type material: Tetrapedia nigripes Friese; holotype
male, ‘Venezuela\ 1896’ ‘Type’ ‘Tetrapedia\ nigripes\
씹 1904 Friese Fr. det.’.
Additional material: See Appendix.
NASUTOPEDIA MICHENERI SP. NOV. (FIGS 213–219)
Comments and diagnosis: Nasutopedia micheneri is
distinguished by its orange yellow metasoma and
yellow infumate wing membrane. Males have thin
pale yellow stripes on lateral margins of clypeus and
lower paraocular area, and are distinguished from the
other two species of the genus by surface of S2–S3
mostly smooth, with few sparse hairs. The female
terga have numerous fine punctures, similar to
N. nigripes.
Distribution: Merida Andes of Venezuela (Fig. 219).
Description
Holotype male: Body length: 7.2; wing length (with
tegula): 8.16; head: width, 2.36, length, 2.12; clypeus:
length, 0.68, width, 1.02; lower interocular distance:
1.06; upper interocular distance: 1.33; scape: length,
0.53, width, 0.17; F1–F3, length: 0.17, 0.09, 0.19; F2
diameter: 0.19. Colour. Head, mesosoma, and legs
mostly black; metasoma orange yellow; labrum pale
yellow, almost white; clypeus with thin pale yellow
stripes on lateral margins; paraocular area with thin
yellow stripes; genal area with thin pale yellow stripe
on lower third; anterior surface of antennal flagellom-
eres pale orange. Wing membrane and veins pale
brown infumate; microtrichia mostly pale brown; fore
wing membrane with blackish microtrichia on apical
margin. Integument sculpture. Disc of clypeus and
supraclypeal area mostly with dense coarse punctures
(< 0.5–1 pd); lateral portions of clypeus with sparse
minute punctures (> 2 pd). Antennal scrobe and frons
mostly with fine punctures (c. 1 pd); disc of frons with
sparse coarse punctures (< 0.5–2 pd); paraocular area
with minute punctures (> 2 pd); mesoscutum and
scutellum with dense minute punctures (1–3 pd);
mesepisternum laterally with sparse fine punctures
(> 3 pd). Metapostnotum with numerous minute
punctures (> 3 pd), and a narrow smooth area along
the mid line. Terga mostly smooth, with few sparse
434 A. J. C. AGUIAR and G. A. R. MELO

Figures 214–218. Nasutopedia micheneri, male; sterna 6–8 (S6–S8), genital capsule, ventral and dorsal view, scale
bars = 0.1 mm.

Figure 219. Distribution records for the species of Nasutopedia.

© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
 SYSTEMATICS OF THE GENUS PARATETRAPEDIA
punctures laterally (> 4 pd). Pubescence. Mostly
brown; except for genal area, and lower portion of
mesepisternum, and pronotal lobe with whitish hairs.
Hind tibia with pale brown plumose hairs on distal
portion. Mandible with simple setae on margin; setae
on proximal portion longer than mandible width.
Labrum mostly glabrous, with few short simple hairs
on lower and lateral margins; clypeus with thin short
decumbent hairs on lateral portions; paraocular area
with sparse thin decumbent hairs; upper paraocular
area with short simple erect setae, length c. 0.5 ¥ F2
diameter; genal area with short plumose hairs inter-
mingled amongst longer simple hairs. Mesoscutum
and scutellum with sparse short plumose pale brown
hairs; mesoscutum with few sparse erect simple setae
on disc; posterolateral portions of scutellum with
erect simple and plumose hairs intermingled, c. 0.15–
0.27 mm in length; mesepisternum laterally with
stout dark brown setae, c. 0.23 mm in length. Meta-
postnotum with numerous short simple pale brown
hairs; marginal bands of plumose hairs on terga
mostly dark brown, except T1 with pale yellow hairs;
T2–T4 with marginal hair band occupying less than
one third of margin laterally; T5 with marginal band
almost complete; T6 with marginal band almost com-
plete. Surface of sterna mostly smooth, with sparse
simple setae; S2–S3 with row of simple pale yellow
setae on margin, on S2 lightly longer on mid portion.
S4 with dense band of long decumbent plumose hairs;
lateral portions with long convergent fringes; S5
mostly smooth, with few coarse plumose setae on
lateral portions; S6 with two pre-apical bands of stout
setae and numerous simple short setae on apical
portion. Structure. Clypeus strongly convex; post-
ocellar carina short, not extending until the post-
ocular portion of vertex; lamella of pronotal collar
very thin acute, with lateral portions curved and
closed, as a closed gutter; mid portion of pronotal
lamella almost contacting the mesoscutum; notauli
weakly sulcate; scutellum weakly convex; pygidial
process thin acute.
Paratype female: (Venezuela, SM0334088\ KUNHM).
Body length: 6.96; wing length (with tegula): 8.16;
head: width, 2.52, length, 2.12; clypeus: length, 0.66,
width, 1.02; lower interocular distance: 1.08; upper
interocular distance: 1.36; scape: length, 0.68, width,
0.19; F1–F3, length: 0.19, 0.09, 0.17; F2 diameter:
0.19. Colour. Similar to male, except for the labrum
black, and absence of pale yellow marks on head.
Integument sculpture. Pattern of sculpture similar to
male but denser than male; clypeus and supraclypeal
area mostly with dense coarse punctures (< 0.5 pd;
almost contiguous); clypeus with few minute punc-
tures on lateral portions; disc of frons with dense
coarse punctures (0.5–2 pd) intermingled amongst
© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
435
sparse fine punctures (1 pd); paraocular area, vertex,
and antennal scrobe mostly with dense minute punc-
tures (c. 1 pd); mesoscutum and scutellum with
minute punctures (c. 1 pd), intermingled amongst few
sparse coarser punctures (> 5 pd); metapostnotum
completely covered by minute punctures (c. 2 pd).
Disc of horizontal surface of T1 and disc of pre-
marginal zone of T2–T3 with dense minute punctures
(3 pd) and sparse coarser punctures laterally (> 3 pd);
disc of pre-marginal zone of T4 with sparse minute
punctures (> 3 pd). Lateral portions of T4 and T5
completely covered with fine sparse punctures (c.
3 pd). Pubescence. Mostly dark brown; similar to
male, except for hind tibial scopa with dense whitish
plumose hairs. Mesepisternum with numerous
minute plumose hairs intermingled amongst simple
long setae. T1 without marginal band; T2–T4 with
short marginal band of dark brown plumose hairs,
occupying less than one-quarter of margin laterally;
T5–T6 with marginal band of simple setae on whole
margin. Structure. Clypeus, weakly convex; lamella of
pronotal collar conspicuously acute on most of its
length; posterolateral portions of pronotal lamella
curved, closed, forming a closed gutter and slightly
acute. Notaulus marked only by a shiny glabrous line;
scutellum weakly convex, slightly concave on mid
line.
Variation: The colour of the metasoma can vary from
orange yellow to dark brown. In addition, the wing
membrane can vary from dark brown infumate to
pale brown infumate. Most specimens have the pubes-
cence of hind tibia with apical portion composed of
white plumose hairs, whereas only a few specimens
have the hind tibia pubescence mostly brown. The
microsculpture on T1–T3 can be restricted to the
pre-marginal zone or occupy the entire tergal surface.
Distribution: Venezuela: Aragua (Fig. 219).
Etymology: The species name is dedicated to Dr
Charles D. Michener.
Type material: Holotype male (SEMC), ‘VENEZU-
ELA: Aragua\ Rancho Grande Biol. Stn., Porta-
chuelo\ Pass, 10°21′0′N,67°41′0′W, 1100 m\ 4 JUN
1998; J.Ashe, R.Brooks, R.Harley\ VEN1ABH98 186
ex: insects moving thr.\ pass against wind-migration’
‘SM0334068\ KUNHM-ENT’. Paratypes: VENEZU-
ELA, 1 female (SEMC), ‘VENEZUELA: Aragua\
Rancho Grande Biol. Stn. Portachuelo\
Pass,10°21′0′N,67°41′0′W, 1100 m\ 4 JUN 1998;
J.Ashe, R.Brooks,R.Hanley\ VEN1ABH98 186 ex:
insects moving thru\ pass against wind-migration’
‘SM0334088\ KUNHM-ENT’, 1 male and 2 females
(SEMC), idem except ‘SM0334053’, ‘SM0334070’ and
436 A. J. C. AGUIAR and G. A. R. MELO
‘SM0334088’; 2 females (MZUSP), ‘Rancho Grande,
est\ Aragua, Venezuela\ 10/3/1982, Brandão\
Campos & Sandoval’.
NASUTOPEDIA RASMUSSENI SP. NOV.
Comments and diagnosis: Nasutopedia rasmusseni is
only known from the male holotype. It is distin-
guished by the mesoscutum mostly smooth with few
sparse punctures, integument completely black,
pronotal lamella with lateral portions raised as a
closed gutter, surface of S2 with numerous simple
setae, and margins of T4–T6 with few setae on lateral
portions.
Description
Holotype male: Body length: 10.0; wing length (with
tegula): 9.3; head: width, 2.75, length, 2.75; clypeus:
length, 0.8, width, 1.2; lower interocular distance: 1.3;
upper interocular distance: 0.15; scape: length, 0.7,
width, 0.2; F1–F3, length: 0.20, 0,17, 0.20; F2 diam-
eter: 0.22. Colour. Mostly black, except tarsomeres
and hind tibia reddish brown. Wing membrane and
veins dark brown infumate; microtrichia mostly dark
brown; fore wing membrane pale brown infumate on
apical margin. Integument sculpture. Clypeus with
sparse coarse punctures, c. 1 pd.; supraclypeal area
with dense coarse punctures, < 1 pd.; frons with
sparse coarse punctures (> 2 pd); upper paraocular
area with sparse fine punctures (> 2 pd). Mesoscutum
mostly smooth and shiny, with very sparse fine punc-
tures (> 5 pd); scutellum with dense fine punctures (c.
1 pd); terga completely smooth. Mesepisternum with
sparse fine punctures (> 4 pd). Pubescence. Mostly
dark brown, except for scopa of hind tibia and basi-
tarsus with pale brown plumose pubescence, almost
pale yellow. Mandible with numerous simple stout
setae, on proximal portion, c. 1.2¥ longer than man-
dible width. Labrum mostly glabrous, with few short
simple hairs on lower and lateral margins; clypeus,
supraclypeal area, and lower paraocular area mostly
glabrous, with few sparse setae; frons, upper paraocu-
lar area, and vertex with numerous erect stout setae;
lower paraocular area mostly glabrous. Mesoscutum
mostly glabrous, with sparse short setae; scutellum
with numerous erect stout and long setae (c. 1 ¥ F2
diameter). Mesepisternum laterally with numerous
long setae (c. 1–2 ¥ F2 diameter). Terga smooth, with
marginal hair bands on T3–T5, occupying less than
one-third of posterior margin laterally; T6 with pre-
marginal band of long plumose hairs. S2 with sparse
short setae on surface, and dense marginal band of
long simple decumbent setae; S3–S4 with dense
pubescence of short setae covering most of surface,
and a marginal band of simple setae; S4 with dense
band of long decumbent plumose hairs; lateral por-
© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
tions with long convergent fringes; S5 mostly smooth,
with few coarse plumose setae on lateral portions; S6
with two pre-apical bands of stout setae and numer-
ous simple short setae on apical portion. Structure.
Clypeus strongly convex; upper margin of clypeus
strongly convex; post-ocellar carina low, not reaching
the post-ocellar portion of vertex; pronotal collar
forming a dorsal lamella with lateral portions high as
a closed gutter; notaulus inconspicuous; scutellum
strongly convex; mesepisternum weakly concave on
upper portion, posterior to omaular area; pygidial
process narrow acute; mid basitarsus less than one-
third wider than long.
Etymology: The species name is dedicated to Dr Claus
Rasmussen.
Distribution: Ecuador (Ecuadorian Cordillera)
(Fig. 219).
Type material: Holotype male (SEMC), ‘ECUADOR:
Pichincha\ Mindo, 10.6 km W, Mindo\ Road\
0°4′23′S, 78°45′14′W, 1375 m\ 28 MAR 1999; D.
Brzoska\ ECU1B99057’ ‘SM0187509\ KUNHM-
ENT’.
BIOGEOGRAPHY
The species of Paratetrapedia exhibit a biogeographi-
cal pattern similar to that observed for Lophopedia
and Tropidopedia, occurring mainly in humid forested
areas, being absent from dry (xeric) areas. Nineteen
species of Paratetrapedia occur mainly in the Amazon
Forest, four species occur only in the Atlantic Forest,
one species is endemic to the Brazilian Cerrado, one
species occurs mainly in the Choco region and six
species occur mainly in Central America. The five
species groups, each one largely distributed in the
Neotropical region, agree with the proposal of
Amorim & Pires (1996), who suggested at least five
rapid replication events involved in the diversification
of the modern Neotropical biota. Most of the groups
have species with partially overlapping distributions.
Nasutopedia, the sister group of Paratetrapedia, is
found in the western slopes of the Andes, in areas
located between 1100 and 2400 m altitude, whereas
the species of Paratetrapedia occur mainly in lowland
forests. This pattern of highland/lowland disjunction
between sister groups has also been reported for some
genera of stingless bees (Camargo & Moure, 1996;
Camargo & Pedro, 2003), frogs (Lynch, 1986), and
parrots (Ribas et al., 2007). In this latter case, Ribas
et al. (2007) inferred that the most probable event
responsible for the vicariance between low and high-
land groups (divergence of habitat areas of cloud and
tropical forest) was the rapid uplift of the northern
 SYSTEMATICS OF THE GENUS PARATETRAPEDIA
portions of Andean cordillera beginning in the
Miocene. Palaeobotanical data from the eastern Cor-
dillera of Colombia suggest a rapid increase of eleva-
tions above 1400 m asl between 2 and 5 Mya
(Gregory-Wodzicki, 2000), which is coincident with
the initial divergence between low and highland
parrot species of the genus Pionus, as estimated with
molecular data (Ribas et al., 2007).
The four species of Paratetrapedia endemic to the
Atlantic Forest, P. bicolor, P. volatilis, P. fervida, and
P. atlantica, belong to different species groups and
exhibit distinct distribution patterns within the
Atlantic Forest domain. Paratetrapedia volatilis and
P. fervida are basal species within the moesta and
lineata groups, respectively. Both reach northern
Argentina, along the Paraná river basin, and in the
northern limit of their distribution, P. volatilis
reaches the mountain chains of southern Bahia,
whereas P. fervida extends its distribution to interior
plateaus with Atlantic Forest remnants (‘Brejos’) in
north-eastern Brazil. Paratetrapedia bicolor and P.
atlantica represent derived lineages within their
respective species groups. These two species have
southern distributional limits at the northern coast of
São Paulo. Whereas P. bicolor is widely distributed,
reaching north-eastern Brazil, P. atlantica is found
only in the states of São Paulo, Rio de Janeiro, and
Espírito Santo. These two distinct vicariance patterns
observed for the Atlantic component are probably
related to a nonmonophyletic Atlantic component,
divided into northern and southern portions (Amorim
& Pires, 1996).
The presence of an endemic species of Paratetrape-
dia in the Brazilian Cerrado coincides with the bio-
geographical pattern observed for the related genera
Tropidopedia (Aguiar & Melo, 2007a, b) and Lopho-
pedia (Aguiar, 2009). Paratetrapedia, Tropidopedia,
and Lophopedia have most of their species in forested
areas, with one or two derived species endemic to
open areas of the Brazilian Cerrado: Paratetrapedia
punctata, Tropidopedia flavolineata Aguiar & Melo, T.
punctifrons (Smith), and Lophopedia savanicola
Aguiar. The presence of both P. punctata and T.
flavolineata– species widely distributed in central
Brazil – in the savannahs of Amapa, in northern
Brazil, supports the hypothesis of an eastern costal
corridor connecting the northern South American
enclaves of savannahs and the Cerrado (Silva, 1995;
Aguiar & Melo, 2007a).
The pattern observed here for Paratetrapedia,
Tropidopedia, and Lophopedia suggests that the
Cerrado endemics in these genera are derived from
vicariance events with the humid Amazon and Atlan-
tic components (Aguiar & Melo, 2007b). This pattern
is different from that obtained for other tapinotaspi-
dine bees of the genera Tapinotaspoides and Caenono-
© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
437
mada whose Cerrado endemics support a closer
relationship of the Cerrado with the Chaco and the
Caatinga (Zanella, 2002; Aguiar & Melo, 2008). These
three areas compose a larger area of endemism
known as the diagonal of open and xeric areas of
South America (Vanzollini, 1974; Ab’Sáber, 1977), or
the Chacoan subregion (Morrone, 2000).
Some of the biogeographical patterns discussed
above, in which the Atlantic Forest and the Cerrado
exhibit a hybrid composition, should be taken as
additional examples to reinforce the view that a
single unique biogeographical scenario for the entire
Neotropical region is improbable (Costa, 2003).
ACKNOWLEDGEMENTS
This paper results from the PhD dissertation of the
first author. We thank the collection curators listed in
the Material and methods for the specimen loans. The
first author also thanks CAPES for the doctoral fel-
lowship; the German Academic Exchange Service
(DAAD) for providing a fellowship to visit the
Museum für Naturkunde der Humbolt-Universität zu
Berlin; the Systematics Research Fund of the System-
atics Association and the Linnean Society of London,
for the grant provided to acquire support material;
the Museum of Comparative Zoology of Harvard Uni-
versity for providing the Mayr Travel Grant to visit
the US collections; Dr Michael Ohl and Dr Frank
Koch for support during the visit to the ZMB collec-
tion; the technician Sérgio Tokunaga of Centro de
Microscopia Eletrônica of Universidade Federal do
Paraná for the scanning electron micrographs; Aline
C. Martins and the Taxonline project for colour
photos. Luana Ferreira Rodrigues is acknowledged
for preparing the line drawings. A. J. C. Aguiar also
expresses his gratitude to Professor Charles D.
Michener, Aline Martins, Claus Rasmussen, Marcel
Hermes, Rodrigo Gonçalves, and Victor Gonzalez for
their careful review and comments on early versions
of the manuscript. Professors Claudio Carvalho,
Danúncia Urban, Fernando Silveira, João M. F.
Camargo, and Luciane Marinoni are thanked for their
corrections and suggestions to the preliminary
version of the first author’s PhD dissertation. G. A. R.
M. acknowledges CNPq for a research fellowship (PQ-
2).
REFERENCES
Ab’Sáber AN. 1977. Os domínios morfoclimáticos da América
do Sul. Primeira aproximação. Geomorfologia 52: 1–21.
Aguiar AJC. 2007. Tapinotaspidini Roig-Alsina & Michener,
1993. In: Moure JS, Urban D, Melo GAR, eds. Catalogue of
bees (Hymenoptera, Apoidea) in the Neotropical region.
Curitiba: Sociedade Brasileira de Entomologia, 608–632.
438 A. J. C. AGUIAR and G. A. R. MELO
Aguiar AJC. 2009. Taxonomic review of the bee genus Lopho-
pedia Michener & Moure (Hymenoptera, Apidae, Tapinotas-
pidini). Zootaxa 2193: 1–52.
Aguiar AJC, Melo GAR. 2005. Notes on the type species of
the subgenera Paratetrapedia (Lophopedia) and P. (Amphi-
pedia) (Hymenoptera, Apidae, Tapinotaspidini). Zootaxa
1084: 31–42.
Aguiar AJC, Melo GAR. 2007a. Taxonomic revision,
phylogenetic analysis, and biogeography of the bee
genus Tropidopedia (Hymenoptera, Apidae, Tapinotaspi-
dini). Zoological Journal of the Linnean Society 151: 511–
554.
Aguiar AJC, Melo GAR. 2007b. Systematics and biogeogra-
phy of the bee genus Paratetrapedia s.l. (Hymenoptera,
Apidae, Tapinotaspidini): cerrado as a composite area. Dar-
winiana 45: 58–60.
Aguiar AJC, Melo GAR. 2008. Biogeografia de áreas abertas
da América do Sul, com ênfase em abelhas da tribo
Tapinotaspidini (Hymenoptera, Apidae). In: De Jong D,
Francoy TM, Santana WC, eds. Anais do VIII Encontro
sobre Abelhas. Ribeirão Preto: Editora FUNPEC, 218–
223.
Aguiar AJC, Melo GAR, Rozen JG, Alves-dos-Santos I.
2004. Synopsis of the nesting biology of Tapinotaspidini
bees (Apidae: Apinae). In: Hartfelder K, ed. Proceedings of
the 8th IBRA International Conference on Tropical Bees and
VI Encontro sobre Abelhas. Ribeirão Preto: Editora
FUNPEC Editora, 80–85.
Albuquerque PMC, Mendonça JAC. 1996. Anthophoridae
(Hymenoptera; Apoidea) e flora associada em uma formação
de cerrado no município de Barreirinhas, MA, Brasil. Acta
Amazônica 26: 45–54.
Albuquerque PMC, Rego MMC. 1989. Fenologia das
abelhas visitantes de murici (Byrsonima crassifolia, Mal-
pighiaceae). Boletim do Museu Paranaense Emílio Goeldi 5:
163–178.
Alves-dos-Santos I. 1999. Abelhas e plantas melíferas da
mata atlântica, restinga e dunas do litoral norte do estado
do Rio Grande do Sul, Brasil. Revista Brasileira de Ento-
mologia 43: 191–223.
Alves-dos-Santos I. 2002. Flower-visiting bees and the
breakdown of the tristylous breeding system of Eichhornia
azurea (Swartz) Kunth (Pontederiaceae). Biological Journal
of the Linnean Society 77: 499–507.
Alves-dos-Santos I. 2003. Adaptations of bee proboscides for
collecting pollen from Pontederia flowers. In: Melo GAR,
Alves-dos-Santos I, eds. Apoidea Neotropica: Homenagem
aos 90 anos de Jesus Santiago Moure. Criciúma: Editora
Unesc, 257–263.
Amorim DS, Pires MRS. 1996. Neotropical biogeography
and a method for maximum biodiversity estimation. Bicudo
CEM, Menezes NA, eds. Biodiversity in Brazil, a first
approach. São Paulo: CNPq, 183–219.
Ayala R. 1988. Abejas silvestres (Hymenoptera: Apoidea) de
Chamela, Jalisco, México. Folia Entomológica Mexicana 77:
395–493.
Ayala R, Griswold TL, Yanega D. 1996. Apoidea
(Hymenoptera. In: Llorente JB, Garcia AN, Gonzáles E, eds.
© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
Biodiversidade Taxonomia y Biogeografia de Artrópodos de
México. Ciudad de Mexico: Universidad Nacional Autónomo
de México, 423–464.
Bortoli C, Laroca S. 1990. Estudo biocenótico em Apoidea
(Hymenoptera) de uma área restrita em São José dos
Pinhais (PR, Sul do Brasil), com notas comparativas.
Dusenia 15: 1–112.
Brothers DJ. 1976. Modifications of the metapostnotum and
origin of the ‘propodeal triangle’ in Hymenoptera Aculeata.
Systematic Entomology 1: 177–182.
Buchmann SL, Buchmann MD. 1981. Anthecology of
Mouriri myrtilioides (Melastomataceae: Memecyleae), an oil
flower in Panama. Biotropica 13: 7–24.
Camargo JMF, Moure JS. 1996. Meliponini neotropicais:
o gênero Geotrigona Moure, 1943 (Apinae, Apidae,
Hymenoptera), com especial referência à filogenia e
biogegrafia. Arquivos de Zoologia 33: 95–161.
Camargo JMF, Pedro SRM. 2003. Meliponini neotropicais:
o gênero Partamona Schwarz, 1939 (Hymenoptera, Apidae,
Apinae) – bionomia e biogeografia. Revista Brasileira de
Entomologia 47: 311–372.
Camillo E, Garófalo CA, Serrano JC. 1993. Hábitos de
nidificação de Melitoma segmentaria, Centris collaris,
Centris fuscata e Paratetrapedia gigantea (Hymenoptera,
Anthophoridae). Revista Brasileira de Entomologia 37: 145–
156.
Cane JH. 1979. The hind tibio-tarsal and tibial spur articu-
lations in bees (Hymenoptera, Apidae). Journal of the
Kansas Entomological Society 52: 123–137.
Casolari C, Moreno RC. 1980. Cataloghi I-Collezione
Imenoterologica di Massimiliano Spinola. Torino: Museo
Regionale di Scienze Naturali.
Cockerell TDA. 1899. Catálogo de las Abejas de México.
Mexico: Secretaria de Fomento.
Cockerell TDA. 1905. Notes on some bees in the British
Museum. Transactions of the American Entomological
Society 31: 309–364.
Cockerell TDA. 1906. The North American bees of the
genera Anthophoridae. Transactions of the American Ento-
mological Society 32: 63–116.
Cockerell TDA. 1909. Descriptions and records of bees.
XXIII. Annals and Magazine of Natural History 4: 393–404.
Cockerell TDA. 1912a. Descriptions and records of bees.
XLV. Annals and Magazine of Natural History 10: 21–31.
Cockerell TDA. 1914. Bees from Ecuador and Perú. Journal
of the New York Entomological Society 22: 328.
Cockerell TDA. 1917. Descriptions and records of bees.
LXXVII. Annals and Magazine of Natural History 20: 298–
304.
Cockerell TDA. 1919. Bees in the collection of the United
States National Museum – 3. Proceedings of the United
States National Museum 55: 167–221.
Cockerell TDA. 1923a. Some bees from British Guianas.
XLIII. Annals and Magazine of Natural History 11: 442–
459.
Cockerell TDA. 1923b. Some bees from Victoria, México.
Proceedings of the United States National Museum 63:
1–5.
 SYSTEMATICS OF THE GENUS PARATETRAPEDIA
Cockerell TDA. 1929. Some results of a journey to Kaieteur
Falls, British Guiana. XLIX. Annals and Magazine of
Natural History 4: 439–444.
Cockerell TDA. 1931. Descriptions and records of bees.
CXXIX. Annals and Magazine of Natural History 8: 411–
418.
Cockerell TDA. 1932. Bees (Hymenoptera, Apoidea) collected
at Chichen Itzá, Yucatan, by the Harvard expeditions of
1929–1930. Bulletin of the Brooklin Entomological Society
27: 9–18.
Cockerell TDA. 1942. Descriptions and records of bees.
CLXXXIV. Annals and Magazine of Natural History 9:
56–60.
Cockerell TDA. 1946. New species and subspecies collected
in a month in Guatemala. Journal of the New York Ento-
mological Society 54: 203–206.
Cockerell WP. 1912. Collecting bees at Gualan, Guatemala.
Canadian Entomologist 44: 277–282.
Cocucci A, Sersic A, Roig-Alsina A. 2000. Oil-collecting
structures in Tapinotaspidini: their diversity, function and
probable origin (Hymenoptera: Apidae). Mitteilungen der
Münchner Entomologischen Gesellschaft 90: 51–74.
Costa LP. 2003. The historical bridge between the Amazon
and the Atlantic Forest of Brazil: a study of molecular
phylogeography with small mammals. Journal of Biogeog-
raphy 30: 71–86.
Cresson ET. 1878. Descriptions of new North American
Hymenoptera in the collection of the American Entomologi-
cal Society. Transactions of the American Entomological
Society 7: 61–136.
Cresson ET. 1879. Catalogue of North American Apidae.
Transactions of the American Entomological Society 7: 215–
232.
Cresson ET. 1916. The Cresson Types of Hymenoptera.
Memoirs of the American Entomological Society 1: 1–141.
Dalla Torre CG. 1896. Catalogus Hymenopterorum hucusque
decriptorum systematicus et synonymicus. Vol. 10: Apidae
(Anthophila). Leipzig: Engelmann.
Darwin C. 1877. Fritz Müller on flowers and insects. Letters
to the Editor. Nature 29: 78–79.
Davies KL. 2009. Food-hair form and diversification in
orchids. In: Kull T, Arditti J, Wong SM, eds. Orchid biology:
review and perspectives, X. California: Springer Science &
Business media, 159–184.
Ducke A. 1901. Beobachtungen über Blütenbesuch, Erschei-
nungszeit etc. der bei Pará vorkommenden Bienen.
Zeitschrift für Systematische Hymenopterologie und Dipter-
ologie 1: 1–18. 49–67.
Ducke A. 1902. Beobachtungen über Blütenbesuch, Erschei-
nungszeit etc. der bei Pará vorkommenden Bienen.
Zeitschrift für Systematische Hymenopterologie und Dipter-
ologie 17: 321–368.
Ducke A. 1908. Contribution à la connaissance de la faune
hyménoptérologique du nord-est du Brésil – II.
Hyménoptéres recoltés dans l’Etat de Ceara em 1908. Revue
d’Entomologie (Caen) 27: 51–81.
Ducke A. 1910a. Zur Synonymie der neotropischen Apidae.
Deutsche Entomologische Zeitschrift 6: 362–369.
© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
439
Ducke A. 1910b. Contribution à la conaissance de faune
hymenopterologique du nord-est du Brésil. III. Hyménop-
téres récoltés dans l’État de Ceara en 1909 et supplément
aux deux listes antérieures. Revue d’Entomologie (Caen) 28:
97–122.
Friese H. 1899. Monographie der Bienengattungen Exoma-
lopsis, Ptilotrix, Melitoma und Tetrapedia. Annalen des
Kaiser-Königlichen der Naturhistorischen Hofmuseums 14:
247–304.
Friese H. 1906. Resultate einer Reise des Herrn A. C.
Jensen-Haarup in die Gegend von Mendoza (Argentina).
Flora Og Fauna Silkeborg, Danmark, III Hefte: 89–102.
Friese H. 1910. Einige neue Tetrapedia – Arten. (Hym.).
Deutsche Entomologische Zeitschrift 1910: 62–65.
Friese H. 1916. Zur Bienenfauna von Costa Rica
(Hym.). Stettiner Entomologische Zeitung 77: 287–
350.
Friese H. 1921. Nachtrag zur Bienenfauna von Costa Rica.
Stettiner Entomologische Zeitung 82: 73–98.
Friese H. 1923. Wissenschaftliche Ergebnisse der schwedis-
chen entomologischen Reise des Herrn Dr. A. Roman in
Amazonas 1914–1915. Arkiv für Zoologi 15: 1–8.
Goloboff PA. 1993. Estimating character weights during tree
search. Cladistics 9: 83–91.
Goloboff PA. 1997. PIWE, version 2.6. Parsimony and
implied weights. Program and documentation.
Goloboff P. 1999. NONA (NO NAME) ver. 2. Published by the
author, Tucumán, Argentina. Tucumán: INSUE, Fundacion
y Instituto Miguel Lillo.
Gonçalves RB, Melo GAR. 2005. A comunidade de abelhas
(Hymenoptera, Apidae s.l.) em uma área restrita de campo
natural no Parque Estadual de Vila Velha, Paraná: diver-
sidade, fenologia e fontes de alimento. Revista Brasileira de
Entomologia 49: 557–571.
Gottsberger G. 1986. Some pollination strategies in Neotro-
pical savannas and forests. Plant Systematics and Evolution
152: 29–45.
Gregory-Wodzicki KM. 2000. Uplift history of the central
and northern Andes: a review. Geological Society of America
Bulletin 112: 1091–1105.
Guth P. 2003. MICRODEM 7.0. United States Naval
Academy. Available at http://www.usna.edu/Oceanography
Heithaus ER. 1979. Community structure of neotropical
flower visiting bees and wasps: diversity and phenology.
Ecology 60: 190–202.
Lucas-de-Oliveira B. 1962. Morfologia externa da larva de
Paratetrapedia (Paratetrapedia) gigantea (S, 1909) e consid-
erações filogenéticas sobre algumas larvas de Antho-
phoridae (Hymenoptera- Apoidea). Boletim da Universidade
Federal do Paraná, Zoologia 12: 1–21.
Lucas-de-Oliveira B. 1966. Descrição de estádios imaturos
de Lanthanomelissa sp. (Hymenoptera: Apoidea). Studia
Entomologica 9: 429–440.
Lutz FE, Cockerell TDA. 1920. Notes on the distribution
and bibliography of north American bees of the families,
Apidae, Meliponidae, Bombidae, Euglossidae, and Antho-
phoridae. Bulletin of the American Museum of Natural
History 42: 491–641.
440 A. J. C. AGUIAR and G. A. R. MELO
Lynch JD. 1986. Origins of the high Andean herpetological
fauna. In: Vuilleumier F, Monasterio M, eds. High altitude
tropical biogeography. New York: Oxford University Press,
478–499.
Melo GAR, Zanella FCV. 2003. The species of the parasitic
bee genus Osirinus (Hymenoptera, Apidae). Journal of
Natural History 20: 2919–2929.
Michener CD. 1942. Taxonomic observations on bees with
descriptions of new genera and species (Hymenoptera;
Apoidea). Journal of the New York Entomological Society 50:
273–282.
Michener CD. 1944. Comparative external morphology, phy-
logeny, and a classification of the bees. Bulletin of the
American Museum of Natural History 82: 151–326.
Michener CD. 1954. Bees of Panamá. Bulletin of the Ameri-
can Museum of Natural History 104: 1–175.
Michener CD. 1997. Genus-group names of bees and supple-
mental family-group names. Scientific Papers of the Natural
History Museum, University of Kansas, 1: 1–81.
Michener CD. 2000. The Bees of the World. Baltimore: Johns
Hopkins University Press, xiv+913 p.
Michener CD, Brooks RW. 1984. Comparative study of the
glossae of bees (Apoidea). Contributions of the American
Entomological Institute 22: 1–73.
Michener CD, Moure JS. 1957. A study of the classification
of the more primitive non-parasitic anthophorine bees
(Hymenoptera, Apoidea). Bulletin of the American Museum
of Natural History 112: 395–452.
Mickeliunas L, Pansarin E, Sazima M. 2006. Biologia floral,
melitofilia e influência de besouros Curculionidae no
sucesso reprodutivo de Grobya amherstiae Lindl. (Orchi-
daceae: Cyrtopodiinae). Revista Brasileira de Botânica 29:
251–258.
Morrone JJ. 2000. Whats is the Chacoan subregion? Neotro-
pica 46: 51–68.
Moure JS. 1941. Notas sobre abelhas do grupo Tetrapedia
Klug (Hym. Apoidea). Revista de Entomologia, Rio de
Janeiro 12: 515–521.
Moure JS. 1944a. Abejas del Perú. Boletin del Museo de
História Natural Javier Prado 8: 67–75.
Moure JS. 1944b. Abelhas de Monte Alegre (Est. S. Paulo)
(Hym.-Apoidea). Papéis Avulsos do Departamento de Zoolo-
gia 6: 103–126.
Moure JS. 1948. Notas sobre algumas abelhas de Tacanas,
Tucumán, Argentina (Hymenopt. Apoidea). Revista de Ento-
mologia 19: 313–346.
Moure JS. 1994. Lissopedia, gen.n. de Paratetrapediini para
a região Neotropical, com as descrições de três espécies
novas (Hymenoptera, Apoidea, Anthophoridae). Revista
Brasileira de Zoologia 9: 305–317.
Moure JS. 1996. Redescrição de alguns exemplares tipos
de espécies neotropicais descritos por Friese em 1899
(Apoidea, Anthophoridae). II. Espécies excluídas do gênero
Tetrapedia Klug. Revista Brasileira de Zoologia 12: 927–937.
Neff JL, Simpson BB. 1981. Oil-collecting structures in the
Anthophoridae (Hymenoptera): morphology, function, and
use in systematics. Journal of the Kansas Entomological
Society 54: 95–123.
© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
Nixon KC. 1999. Winclada (BETA). Version 0.9.9. Published
by the author, Ithaca, NY.
Olson DM, Dinerstein E. 2002. The Global 200: priority
ecoregions for global conservation. Annals of the Missouri
Botanical Garden 89: 199–224.
Pedro SRM. 1996. Lista preliminar das espécies de abelhas
(Hymenoptera, Apoidea) que ocorrem na região de Ribeirão
Preto e Cajuru, SP. In: Garofallo CA et al., ed. Anais do II
Encontro sobre Abelhas. Ribeirão Preto: Universidade de
São Paulo, Faculdade de Filosofia, Ciências e Letras de
Ribeirão Preto, 248–258.
Pedro SRM, Camargo JMF. 1999. Apoidea, Apiformes. In:
Brandão CR, Cancello CM, eds. Biodiversidade do Estado de
São Paulo, Brasil: síntese do conhecimento ao final do século
XX. Vol. 5, Invertebrados terrestres. São Paulo: FAPESP,
194–211.
Pedro SEM, Camargo JMF. 2003. Meliponini Neotropicais:
o gênero Partamona Schwarz, 1939 (Hymenoptera, Apidae).
Revista Brasileira de Entomologia 47 (1): 1–117.
Popov VV. 1939. Relationships of the genus Osiris F. Smith
and its position in the system of bees (Hymenoptera,
Apidae). Comptes Rendus (Doklady) de l’Académie des Sci-
ences de l’URSS 25: 163–166.
Rasmussen C, Ascher JS. 2008. Heinrich Friese (1860–
1948): names proposed and notes on a pioneer melittologist
(Hymenoptera, Anthophila). Zootaxa 1833: 1–118.
Rasmussen C, Cameron SA. 2010. Global stingless bee
phylogeny supports ancient divergence, vicariance, and long
distance dispersal. Biological Journal of the Linnean Society
99: 206–232.
Rebêlo JMM, Rêgo MMC, Albuquerque MCA. 2003.
Abelhas (Hymenoptera, Apoidea) da região setentrional
do Estado do Maranhão, Brasil. In: Melo GAR, Alves-
dos-Santos I, eds. Apoidea Neotropica: homenagem aos 90
anos de Jesus Santiago Moure. Criciúma: Editora Unesc,
265–278.
Rêgo MMC, Albuquerque PMC. 1989. Comportamento das
abelhas visitantes de murici, Byrsonima crassifolia (L.)
Kunth, Malpighiaceae. Boletim do Museu Paraense Emílio
Goeldi, Zoologia 5: 179–193.
Ribas CC, Moyle RG, Miyaki CY, Cracraft J. 2007. The
assembly of montane biotas: linking Andean tectonics and
climatic oscillations to independent regimes of diversifica-
tion in Pionus parrots. Proceedings of the Royal Society B
274: 2399–2408.
Roig-Alsina A. 1997. A generic study of the bees of the
tribe Tapinotaspidini, with notes on the evolution of
their oil-collecting structures (Hymenoptera, Apidae). Mit-
teilungen der Münchner Entomologischen Gesellschaft 87:
3–21.
Roig-Alsina A. 1999. Revisión de las abejas colectoras de
aceites del género Chalepogenus Holmberg (Hymenoptera,
Apidae, Tapinotaspidini). Revista del Museo Argentino Ciên-
cias Naturales 1: 67–101.
Roig-Alsina A, Michener CD. 1993. Studies of the phylogeny
and classification of long-tongued bees (Hymenoptera:
Apoidea). University of Kansas Science Bulletin 55: 123–
173.
 SYSTEMATICS OF THE GENUS PARATETRAPEDIA
Rozen JG. 1984. Comparative nesting biology of the bee tribe
Exomalopsini (Apoidea: Anthophoridae). American Museum
Novitates 2798: 1–37.
Rozen JG, Michener CD. 1988. Nests and immature stages
of the bee Paratetrapedia swainsonae (Hymenoptera: Antho-
phoridae). American Museum Novitates 2909: 1–13.
Rozen JG, Melo GAR, Aguiar AJC, Alves-dos-Santos I.
2006. Nesting biologies and immature stages of the tapino-
taspidine bee genera Monoeca and Lanthanomelissa and
of their osirine cleptoparasites Protosiris and Parepeolus
(Hymenoptera: Apidae: Apinae). American Museum Novi-
tates 3501: 1–60.
San Martin-Gajardo I, Sazima M. 2004. Non-euglossinae
bees also function as pollinators of Sinningia species (Gesne-
riaceae) in southeastern Brazil. Plant Biology 6: 506–512.
Santos FM, Carvalho CAL, Silva RF. 2004. Diversidade de
abelhas (Hymenoptera: Apoidea) em uma área de transição
Cerrado-Amazônia. Acta Amazônica 34: 319–328.
Sazima M, Sazima I. 1989. Oil-gathering bees visit flowers of
eglandular morphs of the oil-producing Malpighiaceae.
Botanica Acta 102: 106–111.
Schrottky C. 1901. Biologische Notizen solitärer Bienen von
S. Paulo (Brasilien). Allgemeine Zeitschrift für Entomologie
6: 209–216.
Schrottky C. 1902. Ensaio sobre as abelhas solitárias do
Brazil. Revista do Museu Paulista 5: 330–613.
Schrottky C. 1909. Hymenoptera nova. Anales de la
Sociedad Científica Argentina 67: 209–228.
Schrottky C. 1913. La distribución geográfica de los
himenópteros argentinos. Anales de la Sociedad Científica
Argentina 75: 115–144. 180–286.
Schwarz H. 1934. The solitary bees of Barro Colourado
Island, Canal Zone. American Museum Novitates 722: 1–24.
Shanks SS. 1986. A revision of the neotropical bee genus
Osiris (Hymenoptera: Anthophoridae). Wasmann Journal of
Biology 44: 1–56.
Sigrist MR, Sazima M. 2004. Pollination biology of twelve
species of Neotropical Malpighiaceae: stigma morphology
and its implications for the breeding system. Annals of
Botany 94: 33–41.
Silva JMC. 1995. Biogeographic analysis of the South Ameri-
can avifauna. Steenstrupia 21: 49–67.
Silveira FA. 1995. Phylogenetic relationships and classifica-
tion of Exomalopsini with a new tribe Teratognathini
(Hymenoptera: Apoidea). University of Kansas Science Bul-
letin 55: 425–454.
Silveira FA, Campos MJO. 1995. A melissofauna de Corum-
bataí (SP) e Paraopeba (MG) e uma análise da biogeografia
das abelhas do cerrado brasileiro (Hymenoptera, Apoidea).
Revista Brasileira de Entomologia 39: 371–401.
Silveira FA, Melo GAR, Almeida EAB. 2002. Abelhas
Brasileiras: Sistemática e Identificação. Belo Horizonte:
Fernando Silveira ed.
Simpson BB. 1989. Krameriaceae. Flora Neotropica 49:
1–109.
Simpson BB, Neff JL. 1981. Floral rewards: alternatives to
pollen and nectar. Annals of the Missouri Botanical Garden
68: 301–322.
© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442
441
Smith F. 1854. Catalogue of the Hymenopterous Insects in the
Collection of the British Museum. Vol. 2. London: British
Museum, 199–465.
Smith F. 1879. Descriptions of new species of Hymenoptera in
the Collection of the British Museum. London: British
Museum.
Smith EL. 1970. Evolutionary morphology of external insect
genitalia. 2. Hymenoptera. Annals of the Entomological
Society of America 63: 1–27.
Smith-Pardo A. 2003. A preliminary account of the bees of
Colombia (Hymenoptera: Apoidea): present knowledge and
future directions. Journal of the Kansas Entomological
Society 76: 335–341.
Spinola M. 1851. Hymenopteros. In: Gay, C. Historia Fisica
y Politica de Chile, Zoologia, Vol. 6. Paris: Casa del autor,
153–569.
Spinola M. 1853. Compte rendu des hyménoptères
inédits provenants du voyage entomologique de
M. Ghiliani dans le Para en 1846. Memorie della Realle
Accademia delle Scienze di Torino, Serie 2 13: 19–
94.
Steiner KE. 1985. Functional dioecism in the Malpighiaceae:
the breeding system of Spachea membranacea Cuatr.
American Journal of Botany 72: 1537–1543.
Strand E. 1909. Beitrag zur bienenfauna von Paraguay
(Hym. Deutsche Entomologische Zeitschrift 1909: 227–
235.
Strand E. 1910. Beiträge zur Kenntnis der Hymenopteren-
fauna von Paraguay auf Grund der Sammlungen und Beo-
bachtungen von Prof. J.D. Anistis. Zoologische Jahrbücher,
Abteilung für Systematik, Geographie und Biologie der Tiere
29: 455–562.
Urban D. 1967. As espécies do gênero Thygater Holmberg,
1884. Boletim da Universidade Federal do Paraná, Zoologia
2: 177–309.
Vachal J. 1909. Especes nouvelles ou litigieuses d’Apidae du
haut bassin du Paraná et des régions contiguës et délimi-
tation d’une nouvelle sous-famille Diphaglossinae. Revue
d’Entomologie 28: 5–64.
Vanzollini P. 1974. Ecological and geographical distribution
of lizards in Pernambuco, northeastern Brazil (Sauria).
Papeis Avulsos de Zoologia 28: 61–90.
Vogel S. 1974. Ölblumen und ölsammelnde Bienen. Tropische
Und Subtropische Pflanzenwelt 7: 1–267.
Vogel S. 1990. History of the Malpighiaceae in the light of
pollination ecology. Memoirs of the New York Botanical
Garden 55: 130–142.
Wittmann D, Hoffman M. 1990. Bees of Rio Grande do Sul,
southern Brazil (Insecta, Hymenoptera, Apoidea). Iheringia
70: 17–43.
Yañez-Ordoñez O, Hinojosa-Díaz I. 2004. La Coleccion
Himenopterologica (Insecta) del Museo de Zoologia ‘Alfonso
L. Herrera’de la facultad de Ciências, UNAM, México. Acta
Zoologica Mexicana 20: 167–197.
Zanella FCV. 2002. Systematics and biogeography of the bee
genus Caenonomada Ashmead, 1899 (Hymenoptera:
Apidae: Tapinotaspidini). Studies on Neotropical Fauna and
Environment 37: 249–261.
 442

APPENDIX
Appendix Table 1. Matrix of character states; the letter ‘p’ indicates polymorphic states; ‘?’ indicates missing data; ‘–’ indicates an inapplicable condition

1 2 3 4 5 6
Terminal Taxa 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1

Arhysoceble dichroopoda 0 0 0 0 0 0 1 0 0 0 0 – 0 0 1 1 0 ? 1 0 – 0 0 – 0 1 0 0 0 0 0 0 0 0 1 0 2 0 1 1 1 1 0 0 ? ? 2 0 0 1 0 0 – 0 – 0 0 0 0 0 0
Lophopedia flava 0 0 0 0 0 1 0 0 1 1 1 2 1 0 1 1 1 1 1 0 – 0 1 0 1 1 0 1 1 0 1 1 1 1 1 0 1 0 0 0 1 1 1 0 0 ? 1 1 0 1 0 1 – 0 – 0 0 0 0 0 0
Lophopedia pygmaea 0 0 0 1 0 1 0 0 1 1 1 2 1 0 1 1 1 1 0 1 0 0 1 0 1 1 0 1 1 0 1 1 1 1 1 0 0 0 0 0 1 1 1 0 0 ? 1 1 0 1 0 0 – 0 – 0 0 0 0 0 0
Trigonopedia oligotricha 0 0 0 0 0 0 1 0 0 1 1 1 1 0 1 1 1 0 0 0 – 1 0 ? 0 0 0 0 0 0 0 0 0 0 1 0 0 0 1 1 1 1 0 0 ? – 2 0 0 1 0 0 – 0 – 0 0 0 0 0 0
Tropidopedia punctifrons 0 1 0 0 0 1 0 0 1 1 1 1 1 0 1 0 1 1 1 0 – 0 1 0 1 1 0 1 0 0 1 1 0 1 1 0 P 0 0 0 1 1 2 0 ? – 1 1 0 1 0 0 – 0 – 0 0 1 0 1 0
Tropidopedia nigrita 0 1 0 0 0 1 0 0 1 1 1 1 1 0 1 0 1 1 0 0 – 0 1 0 1 1 0 1 0 0 0 1 0 0 1 0 0 0 0 0 1 1 2 0 ? – 1 1 0 1 0 0 – 0 – 0 0 0 0 1 0
Xanthopedia larocai 0 0 0 0 0 1 0 0 1 1 1 1 1 0 0 1 1 0 1 0 – 0 0 – 0 1 0 0 0 0 0 1 0 0 1 0 P 0 1 1 1 1 1 0 0 ? 2 0 0 1 0 0 – 0 – 0 0 0 0 0 0
Nasutopedia nigripes 0 0 0 1 0 1 0 0 1 1 0 0 1 0 1 0 1 0 0 0 – 0 1 0 1 0 0 0 0 0 2 1 2 1 1 0 0 0 0 0 0 1 1 0 0 ? 1 1 1 1 0 0 – 0 – 0 0 0 0 0 0
Nasutopedia micheneri 0 0 0 1 0 1 0 0 1 1 0 0 1 0 1 0 1 1 0 0 – 0 1 0 1 0 0 0 0 0 2 1 2 1 1 0 0 0 0 0 0 1 1 0 0 0 1 1 1 1 0 0 – 0 – 0 0 0 0 0 0
Nasutopedia rasmusseni 0 0 ? ? 0 1 0 0 1 1 ? 0 1 0 1 0 1 1 0 0 ? 0 1 ? 1 ? 0 0 0 0 ? 1 2 1 ? 0 1 0 ? ? 0 1 1 0 0 0 1 1 1 1 0 0 – 0 – 0 0 0 0 0 0
Paratetrapedia albilabris 1 0 0 0 1 1 0 0 1 1 1 1 1 0 1 0 1 0 1 0 – 0 1 1 1 0 1 0 0 0 2 1 2 1 0 0 0 0 1 0 0 0 1 0 1 1 3 1 1 0 1 0 – 1 2 0 0 1 1 0 1
P. albopilosa 1 0 0 0 1 1 0 0 1 1 1 1 1 0 1 0 1 0 1 0 – 0 1 1 1 0 1 0 0 0 2 1 2 1 0 0 0 0 1 0 0 0 1 0 1 1 3 1 1 0 1 0 – 1 2 0 0 1 1 0 1
P. alsinai 1 0 0 0 0 1 0 0 1 1 1 1 1 0 1 0 1 0 1 0 – 0 1 1 1 0 1 0 0 1 2 1 2 1 0 1 2 0 1 1 1 1 1 0 1 0 3 1 1 0 ? ? – ? ? ? ? ? ? ? 1
A. J. C. AGUIAR and G. A. R. MELO

P. andina 1 0 0 0 1 1 0 0 1 1 1 1 1 0 1 0 1 0 1 0 – 0 1 1 1 0 1 0 0 0 2 1 2 1 0 0 0 0 1 0 0 1 1 0 1 1 3 1 1 0 1 0 – 1 2 0 0 1 1 0 1
P. atlantica 1 0 0 0 1 1 0 0 1 1 1 1 1 0 1 0 1 0 1 0 – 0 1 1 1 0 1 0 0 0 2 1 2 1 0 0 0 0 1 0 0 0 1 0 1 1 3 1 1 0 1 0 – 1 2 0 0 1 1 0 1
P. basilaris 1 0 0 0 0 1 0 0 1 1 1 1 1 0 1 0 1 0 1 0 – 0 1 1 1 0 1 0 1 0 2 1 2 1 0 0 0 0 1 1 0 0 1 0 0 ? 1 1 1 0 1 1 0 1 2 0 0 1 1 0 1
P. bicolor 1 0 0 0 0 1 0 1 1 1 1 1 1 0 1 0 1 0 1 0 – 0 1 1 1 0 1 0 1 0 2 1 2 1 0 0 0 1 1 1 0 0 1 0 0 ? 1 1 0 0 1 1 0 1 2 0 0 1 1 0 1
P. bifrons 1 0 0 0 0 1 0 0 1 1 1 1 1 0 1 0 1 0 1 0 – 0 1 1 1 0 1 0 0 0 2 1 2 1 0 0 0 0 1 1 P 0 1 0 0 ? 1 1 1 0 1 1 0 1 2 0 0 1 1 0 1
P. calcarata 1 0 0 0 0 1 0 1 1 1 1 1 1 0 1 0 1 0 1 0 – 0 1 1 1 0 1 0 1 0 2 1 2 1 0 0 0 0 1 1 0 0 1 0 0 ? 1 1 0 0 1 1 0 1 2 0 0 1 1 0 1
P. camargoi 1 0 0 0 1 1 0 0 1 1 1 1 1 0 1 0 1 0 1 0 ? 0 1 1 1 0 1 0 0 0 2 1 2 1 0 0 2 0 1 1 1 1 1 0 1 1 3 1 1 0 1 0 – 1 2 0 0 1 1 0 1
P. chocoensis 1 0 0 0 1 1 0 0 1 1 1 1 1 0 1 0 1 0 1 0 – 0 1 1 1 0 1 0 0 0 2 1 2 1 0 0 1 0 1 1 1 1 1 0 1 1 3 1 1 0 1 0 – 1 2 0 0 1 1 0 1
P. connexa 1 0 0 1 0 1 0 0 1 1 1 1 1 0 1 0 1 0 0 1 1 0 1 1 1 0 1 0 0 1 2 1 2 1 0 0 1 1 1 0 P 0 1 0 0 ? 0 1 1 1 1 1 0 1 1 0 0 1 1 0 1
P. duckei 1 0 0 0 1 1 0 0 1 1 1 1 1 0 1 0 1 0 1 0 – 0 1 1 1 0 1 0 0 0 2 1 2 1 0 0 2 0 1 1 1 1 1 0 1 1 1 1 1 0 1 0 – 1 2 0 0 1 1 0 1
P. fervida 1 0 0 0 0 1 0 0 1 1 1 1 1 0 1 0 1 0 P 0 – 0 1 1 1 0 1 0 0 1 2 1 2 1 0 0 1 0 1 1 1 1 1 0 1 0 3 1 1 0 1 1 0 1 2 0 0 1 1 0 1
P. flaveola 1 0 0 0 1 1 0 0 1 1 1 1 1 0 1 0 1 0 1 0 – 0 1 1 1 0 1 0 0 0 2 1 2 1 0 0 2 0 1 1 1 1 1 0 1 1 3 1 1 0 1 0 – 1 2 0 0 1 1 0 1
P. flavescens 1 0 0 0 0 1 0 0 1 1 1 1 1 0 1 0 1 0 1 0 ? 0 1 1 1 0 1 0 0 0 2 1 2 1 0 0 2 0 1 1 1 1 1 0 0 ? 3 1 1 0 1 1 0 1 2 0 0 1 1 0 1
P. flavifrons 1 0 0 0 0 1 0 0 1 1 1 1 1 0 1 0 1 0 1 0 – 0 1 1 1 0 1 0 0 1 2 1 2 1 0 1 2 0 1 1 1 1 1 0 1 0 3 1 1 0 1 1 0 1 2 0 0 1 1 0 1
P. flavipennis 1 0 0 0 1 1 0 0 1 1 1 1 1 0 1 0 1 0 1 0 – 0 1 1 1 0 1 0 0 0 2 1 2 1 0 0 2 0 1 1 1 1 1 0 1 1 1 1 1 0 1 0 – 1 2 0 0 1 1 0 1
P. hyalinata 1 0 0 1 0 1 0 0 1 1 1 1 1 0 1 0 1 0 0 1 0 0 1 1 1 0 1 0 0 1 2 1 2 1 0 0 0 1 1 1 1 1 1 0 0 ? 0 1 1 1 1 1 0 1 1 0 0 1 1 0 1
P. leucostoma 1 0 1 0 0 1 1 0 1 1 1 1 1 1 1 0 1 0 0 0 – 0 1 1 1 0 1 0 0 0 2 1 2 1 0 0 0 0 1 0 1 1 1 0 0 ? 1 1 1 1 1 1 1 1 0 1 0 1 1 0 1
P. lineata 1 0 0 0 0 1 0 0 1 1 1 1 1 0 1 0 1 0 1 0 – 0 1 1 1 0 1 0 0 1 2 1 2 1 0 1 2 0 1 1 1 1 1 0 1 0 3 1 1 0 1 1 0 1 2 0 0 1 1 0 1
P. lugubris 1 0 0 1 0 1 0 0 1 1 1 1 1 0 1 0 1 0 0 1 0 0 1 1 1 0 1 0 0 1 2 1 2 1 0 0 1 1 1 0 0 0 1 0 0 ? 0 1 1 1 1 1 0 0 1 0 0 1 1 0 1
P. mexicana 1 0 0 0 0 1 0 0 1 1 1 1 1 0 1 0 1 0 0 0 – 0 1 1 1 0 1 0 0 1 2 1 2 1 0 0 0 0 1 1 1 1 1 0 0 ? 3 1 1 0 1 1 0 1 2 0 0 1 1 0 1
P. moesta 1 0 1 0 0 1 1 0 1 1 1 1 1 1 1 0 1 0 0 0 – 0 1 1 1 0 1 0 0 0 2 1 2 1 0 0 0 0 1 0 1 1 1 1 0 ? 1 1 1 1 1 1 1 1 0 1 1 1 1 0 1
P. punctata 1 0 0 0 0 1 0 0 1 1 1 1 1 0 1 0 1 0 P 0 – 0 1 1 1 0 1 0 0 1 2 1 2 1 0 0 1 0 1 1 1 1 1 0 1 0 – 1 1 0 1 1 0 1 2 0 0 1 1 0 1
P. romani 1 0 0 0 0 1 0 0 1 1 1 1 1 0 1 0 1 0 1 0 – 0 1 1 1 0 1 0 0 1 2 1 2 1 0 1 2 0 1 0 1 1 1 0 1 0 1 1 1 0 1 1 0 1 2 0 0 1 1 0 1
P. rufa 1 0 0 0 1 1 0 0 1 1 1 1 1 0 1 0 1 0 1 0 – 0 1 1 1 0 1 0 0 0 2 1 2 1 0 0 1 0 1 0 0 1 1 0 1 1 3 1 1 0 1 0 – 1 2 0 0 1 1 0 1
P. scaposa 1 0 1 0 0 1 1 0 1 1 1 1 1 1 1 0 1 0 1 0 – 0 1 1 1 0 1 0 0 0 2 1 2 1 0 0 2 0 1 0 1 1 1 0 0 ? 1 1 1 0 P 1 1 1 0 1 1 1 1 0 1
P. testacea 1 0 0 0 0 1 0 1 1 1 1 1 1 0 1 0 1 0 1 0 – 0 1 1 1 0 1 0 1 0 2 1 2 1 0 0 1 1 1 1 0 0 1 0 0 ? 1 1 0 0 1 1 0 1 2 0 0 1 1 0 1
P. tocantinensis 1 0 0 0 1 1 0 0 1 1 1 1 1 0 1 0 1 0 1 0 – 0 1 1 1 0 1 0 0 1 2 1 2 1 0 1 2 0 1 1 0 1 1 0 1 0 3 1 1 0 1 1 0 1 2 0 0 1 1 0 1
P. vogeli 1 0 0 0 0 1 0 1 1 1 1 1 1 0 1 0 1 0 1 0 – 0 1 1 1 0 1 0 1 0 2 1 2 1 0 0 0 0 1 1 0 0 1 0 0 ? 1 1 1 0 1 1 0 1 2 0 0 1 1 0 1
P. volatilis 1 0 0 0 0 1 1 0 1 1 1 1 1 0 1 0 1 0 0 0 – 0 1 1 1 0 1 0 0 1 2 1 2 1 0 0 1 0 1 0 0 1 1 1 0 ? 1 1 1 0 1 1 1 1 0 0 1 1 1 1 1

© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 351–442