CSIRO PUBLISHING
Invertebrate Systematics, 2011, 25, 490–585
http://dx.doi.org/10.1071/IS10016
Cladistic analysis and revision of Piestus Gravenhorst
with remarks on related genera (Coleoptera : Staphylinidae :
Piestinae)
Edilson Caron A,D, Cibele S. Ribeiro-Costa B and Alfred F. Newton C
A
Campus Palotina, Universidade Federal do Paraná, Rua Pioneiro, 2153, Jardim Dallas,
85950-000, Palotina, PR, Brazil.
B
Laboratório de Sistemática e Bioecologia de Coleoptera (Insecta), Departamento de Zoologia,
Universidade Federal do Paraná, Centro Politécnico, Caixa Postal 19020, 81531-980,
Curitiba, PR, Brazil.
C
Zoology Department, Field Museum of Natural History, Roosevelt Road at Lake Shore Drive,
Chicago, IL 60605, USA.
D
Corresponding author. Email: caron@ufpr.br
Abstract. Rove beetles of the genus Piestus Gravenhorst, 1806 are commonly captured under the bark of or inside
decaying logs from Neotropical forests. Piestus belongs to the subfamily Piestinae, historically an ill-defined dumpingground for Staphylinidae defined by plesiomorphic characters, but which has gradually been restricted in concept and
currently includes only six additional extant genera worldwide. Piestinae in this restricted sense has been considered a
probably monophyletic subfamily, but its status and phylogenetic position, as a possible sister-group of Osoriinae within
the recently proposed Oxyteline group of staphylinid subfamilies, are uncertain and need confirmation. The main aim of
the present study was to provide a morphological cladistic analysis and complete taxonomic revision of Piestus, which, as
the type and most speciose genus of Piestinae, is critical for future phylogenetic studies involving the subfamily. In our
study, the monophyly of Piestus is established and phylogenetic relationships among its species are proposed based on
70 adult morphological characters. Piestus is supported by 11 synapomorphies and high branch support. All species of
Piestus are revised and the genus is redefined. The genus contains 43 species, including 13 species described here for the first
time. The previously proposed subgenera Antropiestus Bernhauer, 1917, Eccoptopiestus Scheerpeltz, 1952, Elytropiestus
Scheerpeltz, 1952, Lissopiestus Scheerpeltz, 1952, Piestus s. str., Trachypiestus Scheerpeltz, 1952 and Zirophorus Dalman,
1821 have not been confirmed, as they were found to be poly- or paraphyletic, or are here removed from Piestus, and
therefore subgenera are not used. The main taxonomic changes are as follows. Lissopiestus, syn. nov. is proposed as new
synonym of Eleusis Laporte, 1835 and its species, E. interrupta (Erichson, 1840), comb. rest., is transferred again to that
genus. Antropiestus, syn. nov. and Eccoptopiestus, syn. nov. are proposed as new synonyms of Hypotelus Erichson, 1839
and their species, H. laevis (Solsky, 1872), comb. nov. and H. andinus (Bernhauer, 1917), comb. nov., are transferred to
Hypotelus. Fourteen new synonymies are proposed (valid species listed first): P. lacordairei Laporte, 1835 = Z. furcatus
Sharp, 1887, syn. nov.; P. capricornis Laporte, 1835 = P. frontalis Sharp, 1876, syn. nov.; P. pennicornis Fauvel,
1864 = P. plagiatus Fauvel, 1864, syn. nov.; P. rectus Sharp, 1876, syn. nov.; P. pygialis Fauvel, 1902, syn. nov.;
P. surinamensis Bernhauer, 1928, syn. nov.; P. minutus Erichson, 1840 = P. nigrator Fauvel, 1902, syn. nov.; P. sulcatus
Gravenhorst, 1806 = P. sanctaecatharinae Bernhauer, 1906, syn. nov.; P. condei Wendeler, 1955, syn. nov.; P. gounellei
Fauvel, 1902 = P. wasmanni Fauvel, 1902, syn. nov.; P. mexicanus Laporte, 1835 = P. alternans Sharp, 1887, syn. nov.;
P. aper Sharp, 1876 = P. schadei Scheerpeltz, 1952, syn. nov.; P. angularis Fauvel, 1864 = P. crassicornis Sharp, 1887, syn.
nov.; H. andinus (Bernhauer, 1917) = P. strigipennis Bernhauer, 1921, syn. nov. One species is revalidated: P. fronticornis
(Dalman, 1821), stat. rev., and one synonym is restored: P. penicillatus (Dalman, 1821) = P. erythropus Erichson, 1840, syn.
rest. Neotypes are designated for P. lacordairei Laporte, 1835 and Oxytelus bicornis Olivier, 1811, and lectotypes are
designated for P. puncticollis Fauvel, 1902, P. capricornis variety muticus Fauvel, 1902, P. zischkai Scheerpeltz, 1951,
P. pennicornis Fauvel, 1864, P. plagiatus Fauvel, 1864, P. pygmaeus Laporte, 1835, P. niger Fauvel 1864, P. minutus
Erichson, 1840, P. nigratror Fauvel, 1902, P. sulcatus Gravenhorst, 1806, P. sanctaecatharinae Bernhauer, 1906,
P. sulcipennis Scheerpeltz, 1952, P. aper Sharp, 1876, P. schadei Scheerpeltz, 1952 and P. andinus Bernhauer, 1917.
Received 24 May 2010, accepted 20 December 2011, published online 7 May 2012
Journal compilation CSIRO 2011
www.publish.csiro.au/journals/is
Revision of Piestus with remarks on related genera
Introduction
The beetle family Staphylinidae, or rove beetles, is the most
diverse family of animals known, with more than 55 400
described species (Grebennikov and Newton 2009). Piestinae
Erichson, 1839 is one of the oldest of the 32 currently
recognised extant staphylinid subfamilies, but historically has
been poorly defined and used as a repository for a diverse
assortment of rove beetles that do not fit well elsewhere;
Bernhauer and Schubert (1910), for example, included nearly
30 genera in their concept of the group (as the tribe Piestini).
Gradually, many of those genera have been removed to form
groups of their own (Apateticinae Fauvel, 1895; Trigonurinae
Reiche, 1865) or added to other well-defined subfamilies
(Blackwelder 1942; Newton 1982a, 1982b), so Piestinae in the
modern sense includes only seven extant genera (see below).
Piestinae are now allocated within the Oxyteline group sensu
Lawrence and Newton (1982) and Thayer (2005) together with
another five subfamilies: Apateticinae, Osoriinae Erichson, 1839,
Oxytelinae Fleming, 1821, Scaphidiinae Latreille, 1807 and
Trigonurinae. Thayer (2005) noted that Piestinae in the current
sense is probably a monophyletic group and sister group of
Osoriinae, but no cladistic analysis has been presented for
corroborating its monophyly, as well as the relationship with
Osoriinae. Cladistic studies within the Oxyteline group sensu
Thayer (2005) have been performed only for Scaphidiinae
(Leschen and Löbl 1995, 2005), Oxytelinae (Herman 1970,
1977; Newton 1982b) and Osoriinae in part (Wu and Zhou
2007).
Invertebrate Systematics
Piestinae currently includes 110 species placed in eight
genera: Abolescus Tikhomirova, 1968, one Jurassic fossil
species found in Karatau, Kazakhstan; Eupiestus Kraatz, 1859,
21 species in the eastern Palearctic and Oriental regions;
Hypotelus Erichson, 1839, eight species in the Nearctic and
Neotropical regions; Parasiagonum Steel, 1950, one species
from New Zealand; Piestoneus Sharp, 1889, five species in the
eastern Palearctic region; Piestus Gravenhorst, 1806, 49 species
in the Nearctic and Neotropical regions; Prognathoides Steel,
1950, one species from Australia; and Siagonium Kirby &
Spence, 1815, 24 species in the Holarctic and Neotropical
regions (Herman 2001b; Zheng 2004; Naomi 2006;
Khachikov 2007).
Piestus is the most speciose genus of Piestinae and has been
divided into seven subgenera: Antropiestus Bernhauer, 1917,
two species; Eccoptopiestus Scheerpeltz, 1952, one species;
Elytropiestus Scheerpeltz, 1952, one species; Lissopiestus
Scheerpeltz, 1952, one species; P. s. str. Gravenhorst, 1806,
24 species; Trachypiestus Scheerpeltz, 1952, nine species; and
Zirophorus (Dalman, 1821), 11 species (Scheerpeltz 1952;
Wendeler 1955; Herman 2001b). A list of all previously
known valid species organised by current subgenera is given
in Table 1. Most species of the genus are saprophagous and
are commonly found associated with decaying logs or in leaf
litter in Neotropical forests.
The general purpose of this study was to initiate a modern
phylogenetic study of Piestinae and its place within Staphylinidae
by focusing on the monophyly and phylogenetic relationships
of Piestus, the type genus and by far the most speciose genus
Table 1. Subgenera and valid species of Piestus prior to the current study (Scheerpeltz 1952,
Wendeler 1955, Herman 2001b)
P. (Antropiestus) Bernhauer, 1917
P. (A.) andinus Bernhauer, 1917
P. (A.) strigipennis Bernhauer, 1921
P. (Eccoptopiestus) Scheerpeltz, 1952
P. (E.) laevis Solsky, 1872
P. (Elytropiestus) Scheerpeltz, 1952
P. (E.) paradoxus Bernhauer, 1917
P. (Lissopiestus) Scheerpeltz, 1952
P. (L.) interruptus (Erichson, 1840)
P. (Piestus) Gravenhorst, 1806
P.(P.) alternans Sharp, 1887
P.(P.) buquetii Fauvel, 1864
P.(P.) condei Wendeler, 1955
P.(P.) erythropus Erichson, 1840
P.(P.) extimus Sharp, 1887
P.(P.) filicornis Fauvel, 1902
P.(P.) fulvipes Erichson, 1840
P.(P.) gounellei Fauvel, 1902
P.(P.) heterocephalus Fauvel 1902
P.(P.) mexicanus Laporte, 1835
P.(P.) minutus Erichson, 1840
P.(P.) niger Fauvel, 1864
P.(P.) nigrator Fauvel, 1902
P.(P.) penicillatus (Dalman, 1821)
P.(P.) pennicornis Fauvel, 1864
P.(P.) plagiatus Fauvel, 1864
P.(P.) puncticollis Fauvel, 1902
P.(P.) pygialis Fauvel, 1902
491
P.(P.) pygmaeus Laporte, 1835
P.(P.) rectus Sharp, 1876
P.(P.) sanctaecatharinae Bernhauer, 1906
P.(P.) sulcatus Gravenhorst, 1806
P.(P.) Surinameensis Bernhauer, 1928
P.(P.) wasmanni Fauvel, 1902
P. (Trachypiestus) Scheerpeltz, 1952
P.(T.) angularis Fauvel, 1864
P.(T.) aper Sharp, 1876
P.(T.) chiriquensis Sharp, 1887
P.(T.) costatus Sharp, 1887
P.(T.) crassicornis Sharp, 1887
P.(T.) nevermanni Scheerpeltz, 1952
P.(T.) rugosus Sharp, 1876
P.(T.) schadei Scheerpeltz, 1952
P.(T.) sulcipennis Scheerpeltz, 1952
P. (Zirophorus) (Dalman, 1821)
P.(Z.) bicornis (Olivier, 1811)
P.(Z.) capricornis Laporte, 1835
P.(Z.) frontalis Sharp, 1876
P.(Z.) furcatus (Sharp, 1887)
P.(Z.) lacordairei Laporte, 1835
P.(Z.) longicornis (Lacordaire, 1833)
P.(Z.) longipennis Fauvel, 1864
P.(Z.) planatus (Sharp, 1887)
P.(Z.) spinosus (Fabricius, 1801)
P.(Z.) validus Sharp, 1876
P.(Z.) zischkai Scheerpeltz, 1951
492
Invertebrate Systematics
of the subfamily. Therefore, the current work provides a
morphological cladistic analysis using adults of all species of
Piestus as well as selected outgroup members of all other extant
genera of Piestinae, and a taxonomic revision of Piestus,
including the redefinition of the genus, descriptions and
illustrations of new and previously known species, and a key
to all valid species.
Material and methods
Phylogenetic methods
Terminals
In total, 45 species of Piestus were included as ingroup, of
which 32 are listed in Herman (2001b) and considered as valid
in the present revision (see ‘Results’, ‘Taxonomic revision’), plus
13 new species. Piestus interruptus, P. fulvipes and P. longicornis
were not included in analysis (for details see ‘Results’,
‘Taxonomic revision’). Given that the relationships among the
genera of Piestinae are unknown, the following species
(representing all other extant genera of Piestinae) were used as
outgroups: Eupiestus sculpticollis Kraatz, 1859 (type species of
the genus); Hypotelus pusillus Erichson, 1840 (type species of the
genus); Parasiagonum hudsoni (Cameron, 1927) (monotypic
genus); Piestoneus monticola Naomi, 1995; Prognathoides
mjobergi (Bernhauer, 1920) (monotypic genus); and
Siagonium punctatum (LeConte, 1866).
Character construction
In the cladistic analysis adult morphological characters from
the exoskeleton and some characters from male and female
genitalia were used and are listed below (see Appendix 1). All
multistate characters were considered as unordered. The matrix
was edited by the NEXUS program, version 0.5.0 (Page 2001).
Missing data were coded as ‘?’ and the inapplicable data as ‘–’
(Table 2).
Analysis
Fitch’s parsimony (Fitch 1971) was adopted for optimisation
criteria. The search for the most parsimonious cladograms was
generated in the TNT program, version 1.1 (Goloboff et al. 2008),
where the characters were treated as of equal weighting and
the following settings were used in the analysis: ‘hold 30000;’,
1000 replications, 10 trees saved per replication, and tree
bisection reconnection (TBR) as branch swapping algorithm.
The Winclada program, version 0.9.9 (Nixon 1999) was used for
representing the cladograms. The trees were rooted ‘a posteriori’
(Nixon and Carpenter 1993) in Hypotelus pusillus. Bremer
support values (Bremer 1994) were used as a measure of
branch support and computed in the TNT program, version 1.1
(Goloboff et al. 2008), using ‘trees suboptimal’ to 10 steps and the
same settings used in the equal weighting (see above).
In a subsequent analysis, the characters were treated using
implied weighting (Goloboff 1993) in the TNT program, version
1.1 (Goloboff et al. 2008), using k = 3 and the same settings
used in the equal weighting analysis. This method resolves
conflict in favour of less homoplastic characters and was used
here as an alternative estimation of phylogeny using the same
data.
E. Caron et al.
Taxonomic revision
Material examined
In the current study ~5000 adult specimens of Piestinae
were examined. The abbreviations cited below for each
institution or private collection are used in all text citations. The
name of the responsible curator or individual is given in
parentheses: AMBC, Private Collection Ayr Bello, Rio de
Janeiro, Brazil (A. Bello); AMNH, American Museum of
Natural History, New York, USA (L. H. Herman); BMNH, The
Natural History Museum, London, United Kingdom
(R. G. Booth); CZUG, Centro de Estudios en Zoología,
Entomología, Universidad de Guadalajara, Guadalajara, México
(J. L. Navarrete-Heredia); DZUP, Coleção de Entomologia
Pe. J. S. Moure, Departamento de Zoologia, Universidade
Federal do Paraná, Curitiba, Brazil (L. M. de Almeida); FMNH,
Field Museum of Natural History, Chicago, USA (A. F. Newton);
INPA – Coleção Sistemática de Entomologia, Instituto Nacional
de Pesquisas da Amazônia, Manaus, Brazil (A. Henriques);
IRSNB, Institut Royal des Sciences Naturelles de Belgique,
Brussels, Belgium (Y. Gérard); MNRJ, Museu Nacional,
Universidade do Rio Janeiro, Rio de Janeiro, Brazil
(M. A. Monné); MUSM, Museo de Historia Natural, Universidad
Nacional Mayor de San Marcos, Lima, Peru (G. Lamas); MZSP,
Museu de Zoologia da Universidade de São Paulo, São Paulo,
Brazil (S. A. Casari); NHRS, Naturhistoriska Riksmuseet,
Stockholm, Sweden (B. Viklund); NMW, Naturhistorisches
Museum Wien, Wien, Austria (H. Schillhammer); SEMC, Snow
Entomological Museum, University of Kansas, Lawrence, USA
(Z. Falin); ZMHB, Museum für Naturkunde der HumboldtUniversität, Berlin, Germany (J. Frisch); and ZMUC, Zoological
Museum, University of Copenhagen, Copenhagen, Denmark
(O. Martin).
Type material was examined for all names listed within
Piestus by Herman (2001b), except for P. bicornis (Olivier,
1811) (lost, neotype here designated), P. oxytelinus Perty,
1830 (not found, junior synonym of P. spinosus),
P. longicornis (Lacordaire, 1833) (not found), P. lacordairei
Laporte, 1835 (lost, neotype here designated), P. fulvipes
Erichson, 1840 (syntype in BMNH), P. validus Sharp, 1876
(syntype in BMNH), Trichocoryne striata Gray, 1832 (not
found, junior synonym of P. penicillatus) and Zirophorus
striatus Guérin-Méneville, 1829 (not found, junior synonym of
P. bicornis). For more details, see ‘Results’ in ‘Taxonomic
revision’.
Labels from type material are organised in sequence from
top to bottom, where the data from each label are enclosed within
double quotes (‘ ’), a forward slash (/) separates lines, and
information enclosed by square brackets ([]) provides added
details about the labels. Information from labels of additional
material is organised, when complete, as follows: country:
district/state/province, number of specimens, locality, extra
information, date, collector (abbreviation of institution). All
information from labels is listed as found, with added details
enclosed by square brackets ([]) and comments in italics.
Distribution of each species is listed in the text by country
and followed by, when possible, district/state/province enclosed
by parentheses ‘()’. Biological notes about each species are
cited from labels and also from cited literature.
Revision of Piestus with remarks on related genera
Invertebrate Systematics
493
Table 2. Data matrix for Piestus
(?) missing data, (-) inapplicable characters
Taxa
Hypotelus pusillus
Eupiestus sculpticollis
Piestus andinus
Piestus laevis
Siagonium punctatum
Parasiagonum hudsoni
Piestoneus monticola
Prognathoides mjobergi
Piestus heterocephalus
P. lacordairei
P. convexus, sp. nov.
P. similis, sp. nov.
P. capricornis
P. puncticollis
P. planatus
P. spinosus
P. longipennis
P. zischkai
P. bicornis
P. fronticornis
P. validus
P. formicinus, sp. nov.
P. termitis, sp. nov.
P. surrufus, sp. nov.
P. pennicornis
P. rossii, sp. nov.
P. pygmaeus
P. buquetii
P. extimus
P. filicornis
P. abdominalis, sp. nov.
P. minutus
P. niger
P. penicillatus
P. sulcatus
P. gounellei
P. mexicanus
P. ecuadorensis, sp. nov.
P. foveolatus, sp. nov.
P. sulcipennis
P. paradoxus
P. nevermanni
P. costatus
P. chiriquensis
P. imperfectus, sp. nov.
P. boliviensis, sp. nov.
P. angularis
P. aper
P. rugosus
P. acuminatus, sp. nov.
P. elegans, sp. nov.
0000000000
0123456789
1111111111
0123456789
2222222222
0123456789
Characters
3333333333
0123456789
4444444444
0123456789
5555555555
0123456789
6666666666
0123456789
0––0000100
0––00001–0
0––0000130
111–000100
100–000100
0––000010?
0––0000100
100–0001–0
100–000010
100–000010
110–00003?
110–00003?
110–000010
0––001002?
100–010010
100–010010
101–010020
101–010020
101–010020
101–010020
101–010020
0––0000111
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0––00102??
000000000–
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0000000011
0000000011
?000?00011
10000??011
1000000010
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1?01010111
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1001000011
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1001010111
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0111000010
000100100–
011100100–
1211100010
1??1100010
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–100–00011
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0100–00011
1100–00000
0400–00000
0400–00000
–401000000
0111011000
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01???11?00
0111011000
01???11?00
1111011000
1111011000
0211011000
0211011000
0211011000
0211011000
0211011000
–111101100
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–1???01?00
–111101100
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–1???01?00
–111101100
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–1???01?00
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–3???01?00
1001110000
1001100100
1001110000
1001110000
1111110001
1211110000
1211110001
1211110001
0101011001
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0100011001
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0111011101
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0102210101
0102010101
0101011001
0100111001
0101111001
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0101011001
0100110111
0100210111
0100210111
0100210111
0100210111
0102200111
0102200111
0102200111
0102200111
0102200111
0102200111
0102200111
0102200111
0112200111
0102200111
0102200111
0102200111
0000–00–00
0000–01–12
0000–00–00
0000–00–00
0000–02001
0000–02000
0000–02001
0000–13011
0101012100
0101012100
0101012100
0101012100
0101012101
0101012101
0101012101
0101012101
0101112100
0101112100
0101112100
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1101012100
1101012100
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0101012100
0101012100
0101012100
0101012100
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1101013112
1101013112
1101013012
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1101013112
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1101013012
1101013012
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1111013012
1111013012
0111013012
1111013012
1111013012
?111013012
0––0001000
1010101000
0––0001000
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0000011000
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0000000110
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0000000111
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0100000010
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0000000110
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0000000110
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0010100010
0010100010
0010100010
0000100010
0010100010
0010100010
0010100010
1010100010
1011100010
1010100010
1111000000
111????00?
111100000?
111100000?
0111000000
0111000000
011100000?
011100000?
0001000101
0001000101
0001100???
0001100???
0001000101
0001000???
0001000101
0001000101
0001010111
0001010111
0000011110
0000011111
0000011111
0001100100
0001100100
0001100100
0001000100
0001100???
0001100101
0001100101
0001100101
0001100101
0001100101
0001000101
0001000101
0001000101
0001100100
0001100101
0001100100
0001100100
0001100100
0001100100
0001100100
0001100100
0001100100
0001100100
0001100100
0001100?0?
0001100100
0001100100
0001100100
0001100100
000????100
The designation of lectotypes and neotypes follows ICZN
(1999), which recommends that lectotype designation ‘should
be done as part of a revisionary or other taxonomic work to
enhance the stability of nomenclature’ (Declaration 44,
Amendment of Article 74.7.3; ICZN 1999), and for neotype
‘with the express purpose of clarifying the taxonomic
status or the type locality of a nominal taxon’ (Article 75.3.1;
ICZN 1999).
494
Invertebrate Systematics
Morphology
The morphological terminology adopted is basically from
Naomi (1987–1990) and Caron et al. (2008). The numbering
of abdominal segments is according to morphological origin and
not indicating visible segments. Gusarov (2002) was used for
describing the position of the aedeagus. Measurements were
done using a micrometric ocular in a stereoscopic microscope
Wild M5 or Leica MZ12.5 and follow Caron et al. (2008), in
which the following abbreviations are used: BL, body length
(from anterior margin of head capsule to posterior margin of
tergite 8); BW, body width (across humeral region); PL,
pronotum length (maximum); PW, pronotum width
(maximum); EL, elytron length (maximum).
Most features were observed from dried pinned specimens.
Apical abdominal segments, including genitalia, were studied
for all species after dissection. Mouthparts were dissected
and studied for the great majority of the species, except
Piestus puncticollis, P. convexus, sp. nov., P. similis, sp. nov.,
P. rossii, sp. nov., P. boliviensis, sp. nov. and P. elegans, sp. nov.
because only holotypes or syntypes for these species have
been available. The specimens were dissected following
Caron et al. (2008). For type specimens, cold ammonia,
NH3(aq), was used for five minutes. The dissections were
carried out under a Zeiss Stemi SV6 or Leica MZ12.5
stereoscopic microscope. The dissected parts were put inside
micro vials with glycerin or on plastic board covered with Canada
balsam, both pinned with the specimen. Drawings were made
under a stereoscopic microscope, Zeiss Stemi SV6 or Leica
MZ12.5, and also with a compound microscope, Zeiss
Standard 20 or Wild M20, all with a drawing tube attached.
The final drawings were produced as in Caron et al. (2008).
Photographs of most of the species were taken as in Caron et al.
(2008), except for Piestus puncticollis, P. interruptus, P. laevis
and P. andinus, which were taken using a Microptics ML
Macro XLT digital imaging system. The final photographs
were edited as in Caron et al. (2008).
Phylogenetic analysis
The cladistic analysis using equal weighting generated eight
equally parsimonious cladograms (Length, L = 170;
Consistency Index, CI = 0.49; Retention Index, RI = 0.88)
(Fig. 1). The strict consensus tree is shown in Figs 2 and 3
(L = 176, CI = 0.47, RI = 0.87).
Species of Piestus form a distinct monophyletic group,
excluding P. andinus and P. laevis, both here transferred to
Hypotelus (see ‘Taxonomic revision’ below) (Fig. 2). Within
Piestus there are four major clades (A, B, C and D), and two
different hypotheses concerning the relationships among
Piestus formicinus and the clades C and D. In hypothesis 1,
P. formicinus is sister group of clade C+D (Fig. 1; H1); in
hypothesis 2, the clade C is sister group to
P. formicinus + clade D (Fig. 1; H2). The other topologies
concern the relationships among three species of the outgroup
(Piestoneus monticola, Prognathoides mjobergi and
Parasiagonum hudsoni) (Fig. 1; H1, H3) and the relationships
within clade A (Fig. 1; H1, H2) and within clade C (Fig. 1; H1,
H2). The analysis utilising implied weighting confirmed the
monophyly of Piestus, excluding P. andinus and P. laevis, and
E. Caron et al.
showed a similar topology, with a few differences among the
relationships of internal terminals (Fig. 4). This analysis
supports the hypothesis of clade C as a sister group of
P. formicinus + clade D.
According to Scheerpeltz’s classification, Piestus
contained seven subgenera (Table 1) (Scheerpeltz 1952). In
the current analysis, the species of the subgenus Antropiestus,
P. andinus and the species of the subgenus Eccoptopiestus,
P. laevis, are excluded from Piestus. The species of the
subgenera Zirophorus, Piestus s. str., Elytropiestus and
Trachypiestus are discussed below. The discussions of
characters are based on the results of the strict consensus
cladogram from equal weighting (Figs 2, 3). When pertinent,
the characters from strict consensus with implied weighting are
also discussed.
Piestus is a strongly monophyletic group supported by 13
unambiguous changes (Fig. 3) (Bremer support, >10; Fig. 2), of
which 11 are synapomorphies: two moderately sized punctures
with setae on the basal half of the dorsal margin of the eyes
(Fig. 86); antennomere 4 densely covered with microsetae
(Fig. 106); openings of glands at the external margin of
mandibles (Fig. 114); maxillary palpomere 3 as wide as long
(Fig. 131); absence of pair of conspicuous setae on apical
margin of the ligula, near the median sclerotised plate
(Fig. 133); carinate mesoventrite process (Fig. 9); groove on
ventral margin of the anterior femur (Fig. 139); very small
emargination on each posterolateral area on sternite 6, external
openings of abdominal defensive gland complex (Fig. 11)
(Caron et al. 2008); absence of fringe on apical margin of
tergite 10 (Fig. 149); one or two pairs of setae at the apex of
tergite 10 (Fig. 149); and one or two setae on the apex of the
ovipositor of females (Fig. 153).
The analysis suggested Piestus is closely related to the clade
Siagonium punctatum + (Parasiagonum hudsoni+Piestoneus
monticola + Prognathoides mjobergi) as supported by six
synapomorphies (Fig. 3) (Bremer support, 6; Fig. 2): not
emarginate anterior angles of the mentum; absence of pair of
conspicuous setae at the apex of the median sclerotised plate of
the ligula; narrowly separated gular sutures (Fig. 8); anterior
margin of the prosternum anteriorly projected at the middle
(Fig. 8); five complete longitudinal striae on each elytron
(Fig. 5); and absence of short setae on apical margin of
sternite 8 of the female. However, a specific phylogenetic
study is necessary to confirm this hypothesis.
Clade A
Within Piestus, clade A comprises 13 species that share seven
unambiguous changes, two of which are synapomorphies
(Fig. 3) (Bremer support, 4; Fig. 2): long epipharynx visible
dorsally, ~2.5 times or so the median length of labrum
(Fig. 110); and maxillary palpomere 4 shorter than 2 and 3
combined (Fig. 130). This clade is similar to the species group
proposed as the subgenus Zirophorus by Scheerpeltz (1951,
1952), except for P. heterocephalus and P. puncticollis, which
were both previously allocated in the subgenus Piestus s. str.
(Table 1). Scheerpeltz (1952) grouped the species within
P. (Zirophorus) based mainly on the presence of the frontal
process on the head and mandibles anteriorly projected. The
Revision of Piestus with remarks on related genera
Invertebrate Systematics
Fig. 1. Three of eight equally parsimonious cladograms showing relationships among species of Piestus using equal weighting of
characters. H1, H2 and H3: hypothesis 1, 2 and 3, respectively.
495
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Invertebrate Systematics
E. Caron et al.
Clade A
Clade B
Clade C
Clade D
Fig. 2. Strict consensus cladogram of eight equally parsimonious trees using equal weighting of characters. Bremer support values shown
below branches.
frontal process (Fig. 22) occurs in other groups of Piestinae,
such as Hypotelus (H. laevis, see ‘Taxonomic revision’)
(Fig. 56), Siagonium (S. punctatum) and Prognathoides
(P. mjobergi), and thus seems to be a plesiomorphic character,
and it is not found in P. puncticollis (Fig. 16). The other
character indicated by Scheerpeltz (1952) was observed in the
Revision of Piestus with remarks on related genera
Invertebrate Systematics
Clade A
Clade B
Clade C
Clade D
Fig. 3. Strict consensus cladogram of eight equally parsimonious trees using equal weighting of characters. Only
unambiguous changes are shown. Black square, synapomorphies; white square, homoplasies.
497
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Invertebrate Systematics
E. Caron et al.
Clade A
Clade B
Clade C
Clade D
Fig. 4. Strict consensus cladogram of three using implied weighting of characters (L = 174, CI = 0.48, RI = 0.87). Only unambiguous changes are
shown. Black square, synapomorphies; white square, homoplasies.
current analysis as a homoplasy for clade A, not found in
P. capricornis and P. puncticollis (Figs 63, 64). The implied
weighting scheme showed a different topology among the
terminals within clade A (compare Fig. 1, H1, H2, and Fig. 4).
Furthermore, the current characters used with equal weighting
do not produce a good resolution at the base of clade A, in
which there is a polytomy (Fig. 3). However, both weighting
schemes confirmed two different clades: clade P. convexus +
P. similis and clade P. longipennis + P. zischkai + (P. bicornis +
P. fronticornis + P. validus).
The clade P. convexus + P. similis was supported only by
homoplasies with equal weighting (Fig. 3) (Bremer support, 8,
Fig. 2). The homoplasies were: short frontal process on head
(Fig. 13); line with more than five moderately sized punctures
with setae; curved sides of pronotum (Fig. 59); and curved
apex of the median lobe of the aedeagus (Fig. 158). The clade
Revision of Piestus with remarks on related genera
P. longipennis + P. zischkai + (P. bicornis + P. fronticornis+
P. validus) was supported by three homoplasies and five
synapomorphies (Fig. 3) (Bremer support, 7; Fig. 2). The
following synapomorphies were observed: two acute teeth on
the internal border of the right mandible, not close (Fig. 121);
long stout setae at the groove on ventral margin of the anterior
femur; longitudinal set of long setae near each lateral margin
on sternite 7; emarginate apex of the median lobe of the
aedeagus of the male (Fig. 181); and emarginate external
margin of the ovipositor of the female (Fig. 154). The clade
P. capricornis + P. puncticollis+(P. planatus + P. spinosus) was
also supported in equal weighting only by homoplasies
(Fig. 3) (Bremer support, 2; Fig. 2): long antennae of male;
projected anterior angles of pronotum (Fig. 65); a slight
emargination in each side of pronotum, anterior to basal
constriction (Fig. 65); fine punctures on striae of elytra; and
sternite 3 with conspicuous transverse carina. This clade was
not confirmed in implied weighting (Fig. 4).
Clade B+(P. formicinus + clade C+D)
This clade is supported by one homoplasy and two
synapomorphies (Fig. 3) (Bremer support, 2; Fig. 2), the
synapomorphies being: strongly acute small basal tooth on
external margin of the mandibles (Fig. 113); and subquadrate
mentum (Fig. 135). The topology of the clade P. formicinus +
clades C+D is supported in equal and implied weightings
(Figs 3, 4) only by character 9 as a unique change: the
antennal length in the male conspicuously longer than in the
female. This character is not found in P. aper (Table 2). The clade
P. formicinus + clades C+D has low branch support (Bremer
support, 1; Fig. 2). In the implied weighting, clade C is sister
group of P. formicinus + clade D. The clade P. formicinus +
clade D is supported by two homoplasies: curved sides of
the pronotum (Fig. 87), and six complete longitudinal striae on
each elytron. This topology appears in one of the most
parsimonious trees estimated using equal weighting (Fig. 1; H2).
Clade B
Clade B has only two species and is strongly supported by
four homoplasies and two synapomorphies (Fig. 3), the latter
being: two lobes at each apex of the epipharynx (Fig. 111);
and undulate microstriae on integument of the interstriae of
the elytra. Clade B has a branch support value of 4 (Fig. 2).
Invertebrate Systematics
499
occurs in most of the species of clade D, except in P. aper
(Table 2). This clade also occurs in equal weighting (Fig. 1;
H2). The clade P. pygmaeus + (P. buquetii + (P.extimus +
(P. filicornis + P. abdominalis + (P. minutus + P.niger +
P. penicillatus)))) is supported by two homoplasies and one
synapomorphy (Fig. 3) (Bremer support, 2; Fig. 2): enlarged
area on basal half of exposed dorsal face of the antennal scape of
males and females. The clade P. filicornis + P. abdominalis +
(P. minutus + P.niger + P. penicillatus) is supported by two
synapomorphies (Fig. 3) (Bremer support, 3; Fig. 2): a slightly
concave region at the middle of the front; and a transverse
carina at the base of the vertex (Fig. 82).
Clade C contains most of the species grouped by Scheerpeltz
(1952) in the subgenus Piestus s. str. (Table 1). The rest of
the species grouped in Piestus s. str. occur in clade A
(P. heterocephalus and P. puncticollis) and in clade D
(P. sulcatus, P. gounellei and P. mexicanus) (Fig. 2).
Clade D
Clade D is the largest clade within Piestus and contains 17
species, which share four homoplasies and one unique
synapomorphy (Fig. 3): a small lateral tooth on the side of the
pronotum, anterior to the basal constriction (Fig. 7). However,
clade D has good branch support (Bremer support, 6; Fig. 2).
The clade P. sulcipennis + (P. paradoxus + (P. nevermanni +
(P. costatus + (P. chiriquensis + P. imperfectus)) + (P. sp. n. L.+
(P. angularis + P. aper) + (P. rugosus + P. acuminatus+
P. elegans))))) is distinct and shares three homoplasies (Fig. 3)
(Bremer support, 3; Fig. 2): enlarged area on basal half of exposed
dorsal face of the antennal scape only in males; sinuous sides of
pronotum (Fig. 100); and inconspicuous median longitudinal
sulcus on pronotum (Fig. 92). All species of this clade were
treated as the subgenera Trachypiestus and Elytropiestus by
Scheerpeltz (1952), based on large punctures on the pronotum
(Table 1). However, similar punctures occur in Eupiestus
sculpticollis, Piestus gounellei, P. mexicanus, P. ecuadorensis
and P. foveolatus.
Within clade D, the topologies remain almost the same
between the different weighting schemes (compare Figs 3 and
4). Thus, clade D is a mixture of species from the subgenera
Piestus s. str., Trachypiestus and Elytropiestus of Scheerpeltz
(1952).
Conclusions
Clade C
Clade C contains 10 species and is supported by three
homoplasies and two synapomorphies (Fig. 3) (Bremer
support, 3; Fig. 2): long setae on basal half of exposed dorsal
face of the antennal scape of the male and female, and antenna
conspicuously narrowing from antennomere 4–11 (Fig. 35).
Within clade C the topologies remain almost the same
between the different weighting schemes, except for
P. pennicornis and P. rossii (compare Figs 3 and 4).
The clade P. pennicornis + P. rossii appears in implied
weighting, being supported by one homoplasy (Fig. 4):
conspicuous ovate set of fine punctures in median region near
the apex of the prosternum of male (Fig. 8). This character
The genus Piestus is a monophyletic group, excluding the
species P. laevis and P. andinus, both here transferred to
Hypotelus (see ‘Taxonomic revision’). Therefore, the genus
Piestus contains 43 species, including 13 new species, and is
supported by 11 synapomorphies (Fig. 3) and high branch
support (Bremer support, >10; Fig. 2). In the current analysis,
the most recent and regularly adopted classification of
Scheerpeltz (1952) (Table 1) has not been entirely confirmed,
as his subgenera are either poly- or paraphyletic or, in two cases,
removed from Piestus. Thus, the subgenera established by
Scheerpeltz (1952) are not recognised here. The genus Piestus
has four evident clades in which most of the species are
distributed. The only species excluded from the clades is
P. formicinus, which appears to be related to clades C and
500
Invertebrate Systematics
D. However, from implied weighting it is possible to suggest that
P. formicinus is more closely related to clade D than C (Fig. 4).
Taxonomic revision
Genus Piestus Gravenhorst, 1806
Piestus Gravenhorst, 1806: 223 (original description); Laporte, 1835:
126 (list of species, characters, notes); Erichson, 1840: 830 (characters,
notes); Sharp, 1876: 403 (list of species, characters, notes); Sharp,
1887: 712 (notes); Bernhauer & Schubert, 1910: 6 (list of species);
Blackwelder, 1943: 43 (characters, notes); Blackwelder, 1944: 100
(list of species); Blackwelder, 1952: 309 (nomenclatural notes);
Scheerpeltz, 1952: 281 (revision); Herman, 2001b: 1787
(bibliography, distribution, notes); Navarrete-Heredia et al., 2002:
207 (characters, notes, list of species); Newton et al., 2005: 37
(list of species from Colombia); Caron et al., 2008: 6 (abdominal
defensive glands complex). Type species: Piestus sulcatus
Gravenhorst, 1806 (fixed by monotypy).
Zirophorus Dalman, 1821: 372 (original description); Laporte, 1835:
126 (synonym of Piestus). Type species: Zirophorus fronticornis
Dalman, 1821 (fixed by subsequent designation by Crotch 1870).
Piestus (Zirophorus): Scheerpeltz, 1951: 5 (key); Scheerpeltz, 1952: 282
(key); Herman, 2001b: 1787 (bibliography); Navarrete-Heredia et al.,
2002: 207 (mention).
Irenaeus Latreille, 1829: 438 (cited as a synonym of Zirophorus);
Lacordaire, 1854: 127 (synonym of Piestus); Herman, 2001b: 1787
(synonym of Piestus, bibliography). Type species: cited as a synonym
of Zirophorus, without type species. Note: nomen nudum (Herman
2001b).
Trichocoryne Gray, 1832: 306 (original description); Herman, 2001b:
1788 (synonym of Piestus, bibliography). Type species: Zirophorus
penicillatus Dalman, 1821 (fixed by subsequent designation by
Blackwelder 1943). Note: not monotypy as cited by Blackwelder
(1943, 1952) and Herman (2001b), because Gray also originally
included T. striata Gray, which he indicated as a probable synonym
of Z. penicillatus.
Tricoryna Laporte, 1835: 126 (synonym of Piestus, attributed to
Gray, but evident misspelling of Trichocoryne). Type species: Not
applicable.
Trachypiestus Scheerpeltz, 1952: 291 (as subgenus of Piestus); Herman,
2001b: 1788 (bibliography); Navarrete-Heredia et al., 2002: 207
(mention). Type species: Piestus angularis Fauvel, 1864 (fixed by
original designation).
Elytropiestus Scheerpeltz, 1952: 294 (as subgenus of Piestus); Herman,
2001b: 1788 (bibliography). Type species: Piestus paradoxus
Bernhauer, 1917 (fixed by original designation and monotypy).
Redescription
BL: 2.6–15.8 mm, BW: 0.8–2.5 mm. Body flat dorsoventrally
to somewhat convex (Fig. 6); light brown to entirely black,
sometimes with all or only apical half of abdominal
segment 7 and segments 8–10 yellowish; antennomere 11
slightly lighter than others; tarsi, maxillary and labial palpi
lighter than body; setae on body golden or black. Body
integument with undulate microstriae, easy to see on
metaventrite (Figs 9, 10); fine to large punctures uniformly
distributed, sometimes contiguous; elytral striae punctured
or not.
Male. Head transverse to subquadrate, front slightly
deflected to entirely deflected (Figs 7, 82), some species with a
pair of acute and short or long frontal processes (Fig. 22);
E. Caron et al.
V-shaped frontal sulcus generally conspicuous and complete,
curved arms joined medially (Fig. 85); anterior angles curved
and generally prominent; eyes somewhat prominent in dorsal
view, and some moderately sized punctures with setae on
basal half of dorsal margin (Fig. 7). Antennae long, reaching
apex of elytra or abdomen (Figs 18, 22); scape the longest or
subequal in length to antennomere 3 (Fig. 106); pedicel the
shortest, antennomeres 4–11 increasing in length and generally
oblong shape; scape slightly the widest, pedicel to antennomere
11 with same width or decreasing in width towards apex;
scape with long setae distributed on basal half or on apical
half of dorsal face (Fig. 106); pedicel and antennomere 3 with
long setae generally distributed on apical half of dorsal face,
and antennomeres 4–11 entirely with microsetae and some
long dispersed setae; antennomeres 3–10 with apical circular
set of long setae. Labrum transverse (Fig. 108), anterior margin
emarginate, four to eight long setae medially; each lateral
third, four long setae (two apical and two subapical).
Epipharynx short or long (Figs 108, 110), deeply emarginate
medially, long fringes on internal margin and short fringes on
external margin. Mandibles strongly projecting or not
(Figs 25, 26); internal border smooth or teethed, symmetrical
or asymmetrical; dorsal teeth absent or present, sometimes
weakly developed; prostheca developed, almost reaching
apex of ventral tooth and ending in sulcus on ventral face
(Fig. 114); mola conspicuous, and short tooth on basal half of
external margin of mandible (Fig. 113). Maxillary cardo
subquadrate and base somewhat rounded (Fig. 130); stipes
triangular; lacinia and galea of same length; lacinia with
dense short setae and spur-like structure on apex; galea
narrower than lacinia, dense short setae on apex and some
long setae on external margin near apex; maxillary palpus
with palpomere 1 the shortest, 2 and 3 subequal in length and
4 the longest, sometimes longer than 2 and 3 combined
(Fig. 131). Mentum subquadrate to 2 times wider than long
(Figs 133–135); each side with set of long setae; anterior
margin somewhat curved; prementum short and hidden
under mentum; ligula emarginate at middle and long setae on
anterior margin; labial palpus with palpomere 1 and 3 subequal
in length and longer than 2 (Fig. 133). Gular plate narrow,
gular sutures narrowly separated (Fig. 8). Pronotum wider than
long, PW/PL = 1.25 (P. zischkai; Fig. 69) to 1.56 (P. aper;
Fig. 98); anterior angles generally weakly projected; surface
sometimes irregular; longitudinal median sulcus conspicuous
or inconspicuous (Fig. 5); longitudinal sulcus of posterior
angles present in some species and reach the middle of
pronotum (Fig. 5); abrupt constriction at basal one-third or
one-quarter and, anterior to it, in some species, a short tooth
and a slight emargination of each side (Fig. 5); apical half, twothrids or three-quarters curved, sinuous or with parallel sides.
Prosternum with anterior margin anteriorly projected in middle
(Fig. 8); some species with conspicuous ovate set of fine
punctures in median region near apex (Fig. 8); protrochantin
visible externally, developed; postcoxal hypomeral projection
acute and long, almost reaching apex of procoxa in ventral view
but not reaching prosternum or closing procoxal cavities.
Scutellum visible, developed and somewhat triangular (Fig. 5).
Elytra truncate; nearly as long as wide, EL/BW = 0.90
(P. spinosus; Fig. 19) to 1.17 (P. paradoxus; Fig. 48); each
Revision of Piestus with remarks on related genera
Invertebrate Systematics
Figs 5–11. Piestus sulcatus. 5, Habitus, male, dorsal view; 6, habitus, male, lateral view; 7, head and pronotum, male,
dorsal view; 8, head and pronotum, male, ventral view; 9, meso and metaventrite, male, ventral view; 10, metaventrite
and base of the abdomen, male, ventral view; 11, apex of the abdomen, male, ventral view. Scale bar (5, 6) = 1 mm; (7–11) =
0.50 mm.
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Invertebrate Systematics
E. Caron et al.
Figs 12–23. Habitus, dorsal view. 12, Piestus convexus, sp. nov.; 13, P. similis, sp. nov.; 14, P. lacordairei; 15, P. heterocephalus;
16, P. puncticollis; 17, P. capricornis; 18, P. planatus; 19, P. spinosus; 20, P. longipennis; 21, P. zischkai; 22, P. bicornis; 23, P. bicornis.
Scale bar = 1 mm.
elytron with five complete longitudinal striae and in some species
stria 6 on basal half and/or stria 7 complete (Fig. 5); apex with
short translucent region; epipleural carina present or not (Fig. 6).
Hind wings fully developed, ~3 times longer than wide; anal area
with long setae and short setae on posterior margin. Mesoventrite
with apical process reaching middle of mesocoxae and carinate
(Fig. 9). Metaventrite developed (Fig. 10); each lateral region
with two or three long posteriorly directed setae; paracoxal suture
Revision of Piestus with remarks on related genera
Invertebrate Systematics
503
Figs 24–35. Habitus, dorsal view. 24, Piestus fronticornis, sp. rev.; 25, P. validus; 26, P. rossii, sp. nov.; 27, P. pennicornis; 28,
P. pygmaeus; 29, P. extimus; 30, P. buquetii; 31, P. filicornis; 32, P. abdominalis, sp. nov.; 33, P. niger; 34, P. minutus; 35, P. penicillatus.
Scale bar = 1 mm.
inconspicuous; posterior margin of metacoxa slightly emarginate.
Anterior legs more robust than others (Fig. 138); procoxae
transverse and contiguous; protrochanter small and
subtriangular;
profemur
robust
and
slightly
flat
anteroposteriorly, apical region of anterior margin of ventral
face with groove and short or long set of contiguous hard setae
504
Invertebrate Systematics
(Figs 138, 139); protibia with apex wider than base, longitudinal
row of robust spines on apical half of external margin and at apex,
internal margin slightly emarginate and four longitudinal rows of
robust setae, robust spine on the internal angle of apex, reaching
tarsomere 4; tarsomeres 1–4 subequal in length, long setae at
apex, tarsomere 5 longer than 1–4 combined, slightly curved;
pretarsus composed of pair of claws, unguitractor plate,
empodium and pair of empodial setae (Fig. 142).
Mesotrochantin present, visible (Fig. 140); mesocoxae
separated, each globose, and metacoxae contiguous (Fig. 10),
each transverse and subtrapezoidal; meso- and metatrochanter
subtriangular (Fig. 141); meso- and metafemur slightly flat
anteroposteriorly and somewhat curved dorsally; meso- and
metatibiae subcylindrical and thin, mesotibia notably more
curved than metatibia; each apical half with longitudinal rows
of robust setae, external rows more robust, apex with two
robust spines, internal the longest; tarsus and pretarsus similar
to anterior legs. Abdominal segments 3–6 with parallel or
slightly curved sides; tergite 1 and 2 weakly sclerotised and
hidden under elytra (Fig. 5); tergites 3–7 with pair of
paratergites; sternite 1 short and inconspicuous and sternite 2
sclerotised and fused with sternite 3 (Figs 143, 144); sternite 3
with transverse carina developed or not (Figs 10, 143, 144);
sternite 6 with very small emargination on each posterolateral
area (Fig. 11), external openings of abdominal defensive
gland complex (for details see Caron et al. 2008); tergite 8
with apical margin curved (Fig. 134); sternite 8 with apical
margin truncate to slightly curved (Fig. 146); tergite 10
separating tergite 9 in two equal parts (Fig. 149) contiguous or
weakly separate at base of tergite 10; tergite 9 with ventral
struts, short or long (Figs 149, 150); tergite 10 with one or two
pairs of long setae near apex; sternite 9 weakly pigmented
(Fig. 152), composed of a single median sclerite, symmetrical,
3 times longer than wide, basal margin acute, apical margin
rounded, with one or two pairs of short setae on apex.
Aedeagus with median lobe curved in lateral view (Fig. 214),
sometimes bulbous base in ventral view; lateral lobes developed,
in some species exceeding apex of median lobe; internal sclerites
of variety format.
Female. In general similar to male, except: apex of
sternite 8 slightly acute (Fig. 147); tergite 9 without ventral
struts; sternite 9 as ovipositor consisting of pair of very weakly
pigmented hemisternites and pair of more apical coxites
(Fig. 153), short setae and some microsetae on apex, and
without stylus. Spermathecal duct short or long, and in some
species sclerotised at base (Fig. 262); spermatheca in various
formats.
Sexual dimorphism
Some species are sexually dimorphic in shape of the frontal
processes of the head, male converging to the apex and female
not (Figs 68, 69); in length of antenna, male longer than female
(Figs 106, 107); at antennal scape, long setae and enlarged area
only in male; on internal border of mandible, male with teeth
and female with only one tooth on the right mandible (Figs 123,
124); at middle region near the apex of prosternum, ovate set of
fine punctures only in male (Fig. 8); and at elytra, length of the
apical process longer in females than males (Figs 48, 49).
E. Caron et al.
Distribution
Nearctic region: USA (only P. extimus), and Neotropical region:
Mexico, Belize, Guatemala, El Salvador, Honduras, Nicaragua,
Costa Rica, Panama, Colombia, Venezuela, Guyana, Suriname,
French Guiana, Ecuador (including Galápagos Islands), Peru,
Bolivia, Brazil, Paraguay, Argentina, Uruguay and the West
Indies (Cuba, Dominican Republic, Puerto Rico, Guadeloupe,
Dominica, Martinique, Saint Vincent and the Grenadines,
Grenada, and Trinidad and Tobago).
Piestus is nearly restricted to the Neotropical region in a
strict sense, excluding Chile and southern Argentina. Only one
species is found in the Nearctic region, in south-western USA
(Arizona).
Biological notes
The species of Piestus are commonly found associated with
organic material in a decomposition process, where they are
saprophages and/or mycophages as adults and larvae (based on
our observations of gut contents). Piestus has been found on or
under bark of decaying logs, in leaf litter, in fungi (‘fungus
covered log’), in decaying cacti and sotol (Dasylirion spp.), in
fruit and flower falls on the ground, and also on dung of mammals.
One species may be associated with termites (P. termitis) and
another with ants (P. formicinus), but these associations need to
be confirmed. Some species of Piestus have been found occurring
together in the same habitat. Piestus species have also been
collected by flight intercept and Malaise traps, Berlese and
Winkler extraction of leaf litter, and attracted to UV light.
Key to species of Piestus
1. Pronotum with lateral tooth before abrupt constriction (Fig. 7); elytra with
epipleural carina (Fig. 6)....................................................................2
Pronotum without lateral tooth before abrupt constriction; elytra without
epipleural carina ...............................................................................18
2. (1) Head with microstriae uniformly distributed (Fig. 7); pronotum with
some fine to moderately sized punctures .............. P. sulcatus (Fig. 5)
Head without microstriae uniformly distributed; pronotum with large
punctures, contiguous or not..............................................................3
3. (2) Pronotum with large punctures not contiguous, and with conspicuous
median longitudinal sulcus (Fig. 88) .................................................4
Pronotum with large punctures contiguous, and with inconspicuous median
longitudinal sulcus (Fig. 91) ..............................................................7
4. (3) Head with fine to moderately sized punctures (Fig. 87); elytra with stria 6
developed at the basal half................................ P. gounellei (Fig. 39)
Head with large punctures (Fig. 88); elytra with stria 6 absent...................5
5. (4) Head with large punctures not contiguous at the median region of front
and vertex (Fig. 88)......................................... P. mexicanus (Fig. 40)
Head with large punctures contiguous at the median region of front and vertex
(Fig. 89) .............................................................................................6
6. (5) Head with median longitudinal sulcus and a conspicuous fovea at the
base of the vertex (Fig. 90) ................ P. foveolatus sp. nov. (Fig. 42)
Head without median longitudinal sulcus and fovea at the base of the vertex
(Fig. 89) ........................................ P. ecuadorensis sp. nov. (Fig. 41)
7. (3) Pronotum with anterior angles acute and projected (Fig. 98) ..............
..................................................................................P. aper (Fig. 51)
Pronotum with anterior angles weakly or not projected ..............................8
8. (7) Mandibles with only one acute tooth on each internal border ...........9
Mandibles with two acute teeth on each internal border, basal anteriorly
projected (Fig. 127)..........................................................................12
Revision of Piestus with remarks on related genera
9. (8) Elytra with stria 6 developed at the basal half .....................................
....................................................................... P. sulcipennis (Fig. 43)
Elytra without stria 6..................................................................................10
10. (9) Elytra with apical margin emarginate when closed (Fig. 50);
metaventrite with moderately sized punctures on median
region ................................................ P. boliviensis sp. nov. (Fig. 50)
Elytra with apical margin not emarginate when closed; metaventrite with
moderately sized punctures only on lateral region ..........................11
11. (10) Aedeagus with slightly curved apex of median lobe in lateral view,
and lateral lobes not narrowing to apex (Fig. 228); spermatheca
L-shaped and spermathecal duct long (Fig. 279).................................
............................................................................ P. costatus (Fig. 45)
Aedeagus with curved apex of median lobe in lateral view, and lateral lobes
narrowing to apex (Fig. 232); spermatheca spiral shaped and
spermathecal duct short (Fig. 281)................ P. nevermanni (Fig. 47)
12. (8) Metaventrite with moderately sized punctures on median region ..13
Metaventrite with moderately sized punctures only on lateral region .......15
13. (12) Elytra with stria 6 developed at the basal half, interstriae 2 and
4 joined at the apex and weakly posteriorly extended in male and
strongly in female (Figs 48, 49)................P. paradoxus (Figs 48, 49)
Elytra without stria 6 at the basal half, interestriae 2 and 4 not joined at
the apex and posteriorly extended ...................................................14
14. (13) Head with V-shaped frontal sulcus incomplete (Fig. 92).................
.........................................................P. imperfectus sp. nov. (Fig. 44)
Head with V-shaped frontal sulcus complete (Fig. 94) .................................
......................................................................P. chiriquensis (Fig. 46)
15. (12) Striae of elytra with large punctures and microgranulate (Fig. 55)..
.........................................................................................................16
Striae of elytra with only large punctures (Fig. 52).... P. angularis (Fig. 52)
16. (15) Elytra with interstriae 1 and 4 joined at the apex and weakly
posteriorly extended (Fig. 53); tergite 3 hidden by elytra ...................
......................................................... P. acuminatus sp. nov. (Fig. 53)
Elytra without interstriae 1 and 4 joined at the apex and weakly posteriorly
extended; tergite 3 not hidden by elytra ..........................................17
17. (16) Antennomeres 4–10 oblong shape (male and female) (Fig. 54);
spermathecal duct short, about the same length as spermatheca
(Fig. 286) .............................................................P. rugosus (Fig. 54)
Antennomeres 4–10 bead-like (female) (Fig. 55); spermathecal duct long,
~2 times the length of spermatheca (Fig. 287) ....................................
............................................................... P. elegans sp. nov. (Fig. 55)
18. (1) Pronotum with a slight emargination on each side, anterior to abrupt
constriction (Fig. 18)........................................................................19
Pronotum without a slight emargination on each side...............................22
19. (18) Head with pair of frontal processes ..............................................20
Head without frontal processes (Fig. 63).................P. puncticollis (Fig. 16)
20. (19) Head with frontal processes shorter than the antennal scape
(Fig. 64); V-shaped frontal sulcus incomplete; dorsal teeth of
mandibles with the same length as ventral teeth (Fig. 119) ................
.......................................................................P. capricornis (Fig. 17)
Head with frontal processes longer than antennal scape; V-shaped frontal
sulcus complete; dorsal teeth of mandibles longer than ventral ......21
21. (20) Head, pronotum and elytra reddish brown and abdomen reddish
dark brown to black, sometimes head darker than pronotum and elytra
(Fig. 19) ............................................................. P. spinosus (Fig. 19)
Body entirely black (Fig. 18)....................................... P. planatus (Fig. 18)
22. (18) Head with frontal processes..........................................................35
Head without pair of frontal processes ......................................................23
23. (22) Antennae short, reaching or exceeding a little the apex of
elytra (male and female); pronotum with abrupt constriction on basal
third ..................................................................................................24
Antennae long, almost or reaching the apex of abdomen (male longer than
female); pronotum with abrupt constriction on basal quarter ..........26
24. (23) Head with V-shaped frontal sulcus incomplete (Fig. 86); elytra
with stria 7 complete......................... P. formicinus sp. nov. (Fig. 38)
Invertebrate Systematics
505
Head with V-shaped frontal sulcus complete (Fig. 84); elytra with stria
7 only at the apical half....................................................................25
25. (24) Aedeagus with lateral lobes exceeding the apex of median lobe
(Fig. 206); spermathecal duct about the same length as spermatheca
(Fig. 269) ............................................... P. termitis sp. nov. (Fig. 36)
Aedeagus with lateral lobes reaching the apex of median lobe, not
exceeding it (Fig. 208); spermathecal duct long, ~3 times the length
of spermatheca (Fig. 270) ..................... P. surrufus sp. nov. (Fig. 37)
26. (23) Head with V-shaped sulcus complete (Fig. 80) ...........................27
Head with V-shaped sulcus incomplete (Fig. 74)......................................34
27. (26) Head with transverse carina at base of vertex (Fig. 82) ...............28
Head without transverse carina at base of vertex (Fig. 76) .......................32
28. (27) Pronotum with undulate microstriae uniformly distributed..........29
Pronotum with undulate microstriae only on lateral regions, sometimes also
near the median longitudinal sulcus.................................................31
29. (28) Head with transverse carina at base of vertex strongly interrupted
at the middle (Fig. 83) ...................................P. penicillatus (Fig. 35)
Head with transverse carina at base of vertex not interrupted or weakly
interrupted at the middle ..................................................................30
30. (29) Specimens of moderate size, ~8.0 mm long; aedeagus with curved
apex of median lobe in lateral view (Fig. 196).. P. filicornis (Fig. 31)
Specimens of small size, ~4.5 mm long; aedeagus without curved apex of
median lobe in lateral view (Fig. 202).................P. minutus (Fig. 34)
31. (29) Pronotum with conspicuous fine punctures and some moderately
sized punctures on the median longitudinal sulcus (Fig. 80); aedeagus
with curved apex of median lobe in lateral view (Fig. 198)................
........................................................P. abdominalis sp. nov. (Fig. 32)
Pronotum only with conspicuous fine punctures (Fig. 81); aedeagus
without curved apex of median lobe in lateral view (Fig. 200)...........
.................................................................................P. niger (Fig. 33)
32. (27) Pronotum with parallel sides (Fig. 76); aedeagus with lateral lobes
exceeding the apex of median lobe by one-quarter of its length in
lateral view (Fig. 1).......................................... P. pygmaeus (Fig. 28)
Pronotum with subparallel sides, very slightly curved on apical half (Fig. 78);
aedeagus with lateral lobes exceeding a little the apex of median lobe
in lateral view (Figs 192, 194).........................................................33
33. (32) Pronotum with undulate microstriae and inconspicuous fine
punctures evenly distributed (Fig. 77); aedeagus with slightly
narrowed apex of lateral lobes in lateral view (Fig. 192) ....................
............................................................................. P. extimus (Fig. 29)
Pronotum with undulate microstriae only on lateral areas and conspicuous
fine punctures evenly distributed (Fig. 78); aedeagus with broad apex
of lateral lobes in lateral view (Fig. 194)............ P. buquetii (Fig. 30)
34. (26) Pronotum with curved sides (Fig. 73); aedeagus with curved
apex of the median lobe in lateral view (Fig. 186)..............................
...................................................................P. rossii sp. nov. (Fig. 26)
Pronotum with parallel sides (Figs 74, 75); aedeagus without curved apex
of the median lobe in lateral view (Fig. 188) ......................................
...................................................................... P. pennicornis (Fig. 27)
35. (22) Head with basal distance between the frontal processes the same
or narrower than the basal width of each one (Fig. 67)...................36
Head with basal distance between the frontal processes wider than basal
width of each one (Fig. 61)..............................................................40
36. (35) Body entirely dark brown to black (Fig. 20); aedeagus without
extended region of median lobe for supporting the base of lateral
lobes (Fig. 172) ................................................................................37
Body brown to black with apical half or all the segment 7 and the segments
8–10 of abdomen yellowish to yellowish brown (Fig. 22), or head
and pronotum dark brown to black with pronotum and elytra
reddish brown, also with apical half of the segment 7 and the
segments 8–10 of abdomen yellowish brown (Fig. 23); aedeagus
with extended region of median lobe for supporting the base of
lateral lobes (Fig. 179) .....................................................................38
506
Invertebrate Systematics
37. (36) Head with frontal processes of male strongly converging towards
their apices (Fig. 68) while in female the internal margins are parallel
(Fig. 69); tergite 10 with two pairs of short setae on the apex, equal in
length; aedeagus with lateral lobes exceeding the apex of median lobe
in lateral view (Fig. 176)..................................... P. zischkai (Fig. 21)
Head with frontal processes of male and female equally and slightly
diverging towards their apices (Fig. 67); tergite 10 with two pairs of
long setae on the apex, basal the longest; aedeagus with lateral lobes
reaching the apex of median lobe in lateral view (Fig. 172) ...............
.......................................................................P. longipennis (Fig. 20)
38. (36) Body brown to black with, in general, all of segment 7 and segments
8–10 of abdomen yellowish (Fig. 24); aedeagus in lateral view very
thin and with slightly developed internal sclerite (Fig. 182) ...............
...................................................................... P. fronticornis (Fig. 24)
Body brown to black with apical half of segment 7 and segments 8–10
of abdomen yellowish to yellowish brown (Fig. 22), or head and
pronotum dark brown to black with pronotum and elytra reddish
brown, also with apical half of the segment 7 and the segments 8–10
yellowish brown (Fig. 23); aedeagus different from above.............39
39. (38) Body brown to black with apical half of segment 7 and segments
8–10 of abdomen yellowish (Fig. 25); antennal scape with many long
setae on the apical half of the dorsal face and antennomeres 3–6
with many long setae on dorsal face (Figs 25, 72)..............................
..............................................................................P. validus (Fig. 25)
Body brown to black with apical half of segment 7 and segments 8–10 of
abdomen yellowish brown (Fig. 22), or head and pronotum dark
brown to black with pronotum and elytra reddish brown, also with
apical half of segment 7 and segments 8–10 of abdomen yellowish
brown (Fig. 23); antennal scape with some long setae at the middle of
the dorsal face and antennomeres 3–6 with some long setae on
dorsal face (Figs 22, 23, 70) .........................P. bicornis (Figs 22, 23)
40. (35) Head with short frontal processes shorter than the antennal scape
(Fig. 60) ...........................................................................................41
Head with long frontal processes, longer than the antennal scape (Fig. 61) .
.........................................................................................................42
41. (40) Antennae long, almost reaching the apex of abdomen (Fig. 12), and
scape with long setae on basal half of exposed dorsal face (Fig. 59)..
.............................................................P. convexus sp. nov. (Fig. 12)
Antennae short, reaching the apex of elytra (Fig. 13), and scape with only
two long setae at the middle of the dorsal face (Fig. 60) ....................
................................................................. P. similis sp. nov. (Fig. 13)
42. (40) Body entirely brown to black, sometimes with the abdomen
slightly lighter than the body (Fig. 14); head with ventral face of
frontal processes without or with only one tooth near the
base ................................................................ P. lacordairei (Fig. 14)
Body brown to black with apical half of segment 7 and segments 8–10 of
abdomen yellowish (Fig. 15); head with ventral face of frontal
processes coarsely serrate ........................ P. heterocephalus (Fig. 15)
The species of Piestus are arranged below based on the
current phylogenetic study (see ‘Results’, ‘Cladistic analysis’)
and to facilitate the discussion.
Piestus convexus, sp. nov.
(Figs 12, 59, 156, 157)
Type material
Holotype deposited in SEMC, male: (1) ‘Ecuador: Napo, Cosanga/4.2 km S
on Baeza-Tena/Road then 1.5 km W on pipeline/access road, 2150 m/
0370 1900 S, 77500 100 W/5–7 Nov 1999, Z.H. Falin/ECU1F99 123 flight
intercept trap’ [white label, printed in black]; (2) ‘bar code/SM0352227/
KUNHM-ENT’ [white label, printed in black]; (3) ‘<’ [white label, printed in
black].
E. Caron et al.
Description
BL: 9.5 mm, BW: 2.7 mm. Body convex; entirely black (Fig. 12);
setae on body black, except antennomeres 10–11, and tarsi
golden. Dorsal integument of head and pronotum with
undulate microstriae and inconspicuous fine punctures
(Fig. 59); elytral striae unpunctured; abdominal tergites on
anterolateral regions, with moderately sized punctures.
Male. Head with pair of broad and short frontal processes,
shorter than the scape, and basal distance between processes
~2 times basal width of each one (Fig. 59); V-shaped frontal
sulcus incomplete, curved arms not joined medially; anterior
angles curved and prominent; eyes with line of five moderately
sized punctures with long setae on basal half of dorsal margin,
equal in length. Antennae almost reaching apex of abdomen
(Fig. 12); scape with long setae on basal half of exposed
dorsal face; scape shorter than antennomeres 2 and 3
combined; antennomeres 4–11 oblong shape. Mandibles
strongly projected, bifurcate apices; each with dorsal tooth
shorter than the ventral; internal border asymmetrical, with one
acute tooth at middle and, anterior to it, in right mandible,
with two areas that resemble a tooth, absent in left mandible.
Maxillary palpus with palpomere 4 shorter than 2 and 3
combined. Pronotum wider than long (PW/PL = 1.43)
(Fig. 59); anterior angles weakly projected; longitudinal
median sulcus conspicuous; longitudinal sulcus of posterior
angles slightly developed and reaching basal quarter
of pronotum; abrupt constriction at basal quarter; apical threequarters with slightly curved sides. Femur of anterior legs with
short, hard setae on groove of ventral margin. Elytra as long as
wide (EL/BW = 1.04); stria 6 at basal half and stria 7 at apical
quarter inconspicuous; striae narrower than interstriae.
Abdominal segments 3–6 parallel sides; tergite 9 with short
ventral struts; tergite 10 with two pairs of long setae on apex,
apical the longest. Median lobe of aedeagus with bulbous base in
ventral view and curved apex in lateral view (Fig. 156); lateral
lobes exceed a little apex of median lobe in lateral view; internal
sclerites as in Fig. 157.
Female. Unknown.
Distribution
Ecuador (Napo).
Biological notes
The unique specimen (holotype) was collected in a flight
intercept trap.
Remarks
Piestus convexus differs from P. similis by the broad and short
frontal processes (Fig. 59), long antennae almost reaching the
apex of the abdomen (Fig. 12), antennal scape with long setae on
the basal half of exposed dorsal face, and mandibles with
asymmetrical internal border and without external marginal
carina on basal half.
Piestus similis has acute and short frontal processes (Fig. 60),
antennae short, reaching the apex of elytra (Fig. 13), antennal
scape with two long setae at the middle of the dorsal face,
mandibles with symmetrical internal border and external
margin slightly carinate on basal half.
Revision of Piestus with remarks on related genera
Invertebrate Systematics
507
Figs 36–46. Habitus, dorsal view. 36, Piestus termitis, sp. nov.; 37, P. surrufus, sp. nov.; 38, P. formicinus, sp. nov.; 39, P. gounellei; 40, P. mexicanus; 41,
P. ecuadorensis, sp. nov.; 42, P. foveolatus, sp. nov.; 43, P. sulcipennis; 44, P. imperfectus, sp. nov.; 45, P. costatus; 46, P. chiriquensis. Scale bar = 1 mm.
508
Invertebrate Systematics
E. Caron et al.
Figs 47–58. Habitus, dorsal view. 47, Piestus nevermanni; 48, P. paradoxus, male; 49, P. paradoxus, female; 50, P. boliviensis, sp. nov.; 51, P. aper; 52,
P. angularis; 53, P. acuminatus, sp. nov.; 54, P. rugosus; 55, P. elegans, sp. nov.; 56, Hypotelus laevis, comb. nov.; 57, H. andinus, comb. nov.; 58, Eleusis
interrupta, comb. rest. Scale bar = 1 mm.
Revision of Piestus with remarks on related genera
Invertebrate Systematics
509
Figs 59–75. Head and pronotum, dorsal view. 59, Piestus convexus, sp. nov.; 60, P. similis, sp. nov.; 61, P. lacordairei; 62, P. heterocephalus; 63,
P. puncticollis; 64, P. capricornis; 65, P. planatus; 66, P. spinosus; 67, P. longipennis; 68, P. zischkai, male; 69, P. zischkai, female; 70, P. bicornis; 71,
P. fronticornis, sp. rev.; 72, P. validus; 73, P. rossii, sp. nov.; 74, P. pennicornis, male; 75, P. pennicornis, female. Scale bar = 0.50 mm.
510
Invertebrate Systematics
E. Caron et al.
Figs 76–91. Head and pronotum, dorsal view. 76, Piestus pygmaeus; 77, P. extimus; 78, P. buquetii; 79, P. filicornis; 80, P. abdominalis, sp. nov.; 81, P. niger;
82, P. minutus; 83, P. penicillatus; 84, P. termitis, sp. nov.; 85, P. surrufus, sp. nov.; 86, P. formicinus, sp. nov.; 87, P. gounellei; 88, P. mexicanus; 89,
P. ecuadorensis, sp. nov.; 90, P. foveolatus, sp. nov.; 91, P. sulcipennis. Scale bar = 0.50 mm.
Revision of Piestus with remarks on related genera
Invertebrate Systematics
511
Etymology
Remarks
The specific name refers to the body format.
Piestus similis differs from P. convexus by the characters cited
above (see remarks under P. convexus).
Piestus similis, sp. nov.
(Figs 13, 60, 158, 159)
Type material
Etymology
The name of this species refers to the similar species,
P. convexus.
Holotype deposited in SEMC, male: (1) ‘Ecuador: Napo/Sierra Azul,
2300m/0400 000 S, 77550 000 W/21 May–6 Jun 1996, P. Hibbs/ECU2H96
004D, ex: Malaise trap’ [white label, printed in black]; (2) ‘bar code/
SM0170320/KUNHM-ENT’ [white label, printed in black]; (3) ‘<’ [white
label, printed in black].
Description
BL: 9.7 mm, BW: 2.8 mm. Body convex; entirely dark brown
(Fig. 13); setae on body black, except antennomeres 10–11 and
tarsi golden. Dorsal integument of head and pronotum with
undulate microstriae and inconspicuous fine punctures
(Fig. 60); elytral striae unpunctured; metaventrite on lateral
regions, and abdominal tergites on anterolateral regions with
moderately sized punctures.
Male. Head with pair of acute and short frontal processes,
shorter than scape, and basal distance between processes ~6
times basal width of each one (Fig. 60); V-shaped frontal
sulcus incomplete, curved arms not joined medially; anterior
angles curved and prominent; eyes with one line of five
moderately sized punctures with long setae on basal half of
dorsal margin, equal in length. Antennae reaching apex of
elytra (Fig. 13); scape with two long setae at middle of dorsal
face; scape subequal to antennomere 2 and 3 combined;
antennomeres 4–11 oblong shape. Mandibles strongly
projected; bifurcate apices; each with dorsal tooth shorter than
ventral; internal border symmetrical, right mandible with one
acute tooth at middle; external margin slightly carinate at basal
half. Maxillary palpus with palpomere 4 shorter than 2 and 3
combined. Pronotum wider than long (PW/PL = 1.39) (Fig. 60);
anterior angles weakly projected; longitudinal median sulcus
conspicuous; longitudinal sulcus of posterior angles slightly
developed and reaching basal quarter of pronotum; abrupt
constriction at basal quarter; apical three-quarters with curved
sides. Femur of anterior legs with short, hard setae on groove of
ventral margin. Elytra as long as wide (EL/BW = 1.00); stria 6 at
basal half and stria 7 absent; striae narrower than interstriae.
Abdominal segments 3–6 with parallel sides; tergite 9 with short
ventral struts; tergite 10 with two pairs of long setae on apex,
apical the shortest. Median lobe of aedeagus with bulbous base in
ventral view and curved apex in lateral view (Fig. 158); lateral
lobes reaching apex of median lobe in lateral view; internal
sclerites as in Fig. 159.
Female. Unknown.
Distribution
Ecuador (Napo).
Biological notes
The unique specimen (holotype) was collected in a Malaise
trap.
Piestus lacordairei Laporte
(Figs 14, 61, 116, 160, 161, 251)
Piestus lacordairei Laporte, 1835: 129 (original description, type
locality: ‘Cayenne’); Erichson, 1840: 832 (characters, distribution);
Fauvel, 1864: 21 (characters, distribution); Blackwelder, 1944: 100
(distribution, error: Laporte, 1834: 129); Herman, 2001b: 1791
(distribution); Newton et al., 2005: 37 (distribution); Caron et al.,
2008: 5 (characters).
Piestus (Zirophorus) lacordairei: Bernhauer & Schubert, 1910: 6
(distribution, error: Laporte, 1834: 129); Scheerpeltz, 1951: 7
(characters, distribution, error: Laporte, 1834: 129); Scheerpeltz,
1952: 283 (characters, distribution, error: Laporte, 1834: 129).
Zirophorus furcatus Sharp, 1887: 712 (original description, type locality:
‘Panama, Volcan de Chiriqui 2500 to 4000 feet’). New synonym.
Piestus (Zirophorus) furcatus: Bernhauer & Schubert, 1910: 6
(distribution); Scheerpeltz, 1951: 7 (characters, distribution);
Scheerpeltz, 1952: 283 (characters, distribution).
Piestus furcatus Blackwelder, 1944: 100 (distribution); Herman, 2001b:
1791 (distribution).
Type material
Piestus lacordairei Laporte, 1835. Neotype (here designated) deposited in
ZMHB, sex undetermined, with labels: (1) ‘6804’ [white label, printed in
black]; (2) ‘Lacordairei/Lap./Cayenne Dej.’ [green label, handwritten]
(ZMHB). Note: In the original description Laporte (1835) did not specify
how many specimens were observed. The type material was not found and
probably it is lost.
Zirophorus furcatus Sharp, 1887. Syntype deposited in FMNH, sex
undetermined, with labels: (1) ‘Piestus furcatus./D.S./V. de Chiriqui.
2500-4000 ft./Champion.’ [white label, handwritten, together with the
specimen]; (2) ‘V. de Chiriqui,/25-4000 ft./Champion.’ [white label,
printed in black]; (3) ‘B.C.A. Col. I. 2./Zirophorus/furcatus,/Sharp.’ [white
label, printed in black]; (4) ‘Sharp Coll. 1905,-313.’ [white label, printed in
black]; (5) ‘Chicago Nat. Hist. Mus./(ex. D. Sharp Colln./by exchange with/
Brit. Mus. Nat. Hist.)’ [white label, printed in black]. Note: In the original
description Sharp (1887) specified eight specimens observed. There are
seven syntypes (not examined) deposited in BMNH (Dr Roger Booth,
BMNH, pers. comm.).
Additional material
See Appendix 2.
Redescription
BL: 5.1–8.0 mm, BW: 1.4–2.1 mm. Body flattened
dorsoventrally; entirely brown to black, sometimes with
abdomen lighter than body (Fig. 14). Dorsal integument of
head and pronotum with undulate microstriae and
inconspicuous fine punctures (Fig. 61); striae of elytra
unpunctured or with inconspicuous fine punctures;
512
Invertebrate Systematics
metaventrite, on lateral regions, and abdominal tergites, on
anterolateral regions, with moderately sized punctures.
Male. Head with pair of acute and long frontal processes,
longer than the scape, ventral face of each frontal process with
only one tooth near base, and basal distance between processes
~2 times basal width of each one (Fig. 61); V-shaped frontal
sulcus incomplete, curved arms not joined medially; anterior
angles curved and prominent; eyes with two moderately sized
punctures with long setae on basal half of dorsal margin of eyes,
basal the longest. Antennae almost reaching apex of elytra
(Fig. 14); scape with two long setae near apex of dorsal face;
scape subequal to antennomere 2 and 3 combined; antennomeres
4–11 oblong shape. Labrum with four setae medially, equal in
length; each lateral third, antero-internal seta with same length as
postero-external seta. Mandibles strongly projected (Fig. 116);
bifurcate apices; each with dorsal tooth shorter than ventral;
internal border asymmetrical, internal border with one acute
tooth at middle and, anterior to it, in right mandible with an
area that resembles a tooth, absent in left mandible. Maxillary
palpus with palpomere 4 shorter than 2 and 3 combined. Mentum
2 times wider than long. Pronotum wider than long (PW/
PL = 1.27) (Fig. 61); anterior angles not projected; longitudinal
median sulcus conspicuous; longitudinal sulcus of posterior
angles slightly developed and reaching basal quarter of
pronotum; abrupt constriction at basal quarter; apical threequarters with parallel sides. Femur of anterior legs with short,
hard setae on groove of ventral margin. Elytra somewhat
longer than wide (EL/BW = 1.05); stria 6 at basal half and stria
7 absent; striae narrower than interstriae. Abdominal segments
3–6 parallel sides; tergite 9 with short ventral struts; tergite 10
with two pairs of long setae on apex, apical the longest. Median
lobe of aedeagus with bulbous base in ventral view and not
curved apex in lateral view (Fig. 160); lateral lobes exceeding
apex of median lobe by one-quarter of its length in lateral view
and slightly curved apex on ventral margin; internal sclerites as
in Fig. 161.
Female. Similar to male. Ovipositor as in P. sulcatus;
spermathecal duct ~2 times length of spermatheca, with
sclerotised basal region (Fig. 251); spermatheca with basal half
suddenly narrower than apical half (Fig. 251).
Distribution
In the current study, the species is listed from Nicaragua (Rio
San Juan), Costa Rica (Guanacaste, Alajuela, Heredia, San José,
Cartago, Limón and Puntarenas), Panamá (Bocas del Toro,
Chiriquí, Veraguas and Colón), Colombia (Cundinamarca and
Huila), French Guiana (Saint-Laurent-du-Maroni and Cayenne),
Ecuador (Esmeraldas, Pichincha, Napo and Cotopaxi), Peru
(Ucayali and Cusco) and Brazil (Pará and Rondônia). Herman
(2001b) listed the species also from Venezuela, Guyana and
Suriname, and Newton et al. (2005) from Colombia (Valle del
Cauca).
E. Caron et al.
Remarks
Piestus lacordairei is similar to P. heterocephalus but differs
mainly by the body being entirely brown to black (Fig. 14), ventral
face of frontal processes with only one tooth near the base,
labrum with four setae medially and lateral lobes of aedeagus
with slightly curved apex on ventral margin (Fig. 160).
Piestus heterocephalus has body black with apical half of
abdominal segment 7 and all of segments 8–10 yellowish
(Fig. 15), ventral face of frontal processes coarsely serrate, six
setae medially on labrum and lateral lobes of aedeagus with
truncate apex on ventral margin (Fig. 162).
Piestus heterocephalus Fauvel, 1902
(Figs 15, 62, 162, 163, 252)
Piestus heterocephalus Fauvel, 1902: 22 (original description, type
locality: ‘Venezuela: Merida’); Blackwelder, 1944: 100
(distribution, error: Fauvel, 1865: 22); Herman, 2001b: 1791
(distribution); Caron et al., 2008: 3 (redescription, distribution,
defensive abdominal gland complex).
Piestus (Piestus) heterocephalus: Bernhauer & Schubert, 1910:
7 (distribution); Scheerpeltz, 1952: 285 (characters, distribution).
Type material
Holotype deposited in IRSNB, sex undetermined, with labels: (1)
‘Merida/Venezuela’ [white label, handwritten]; (2) ‘heterocephalus/Fvl.’
[white label, handwritten]; (3) ‘R.I.Sc.N.B. 17.479/Piestus/Coll. et. det.
A. Fauvel’ [white label, first and third lines printed in black, and second
line handwritten]; (4) ‘Ex-Typis’ [white label, printed in red]. Note: For
discussion about the type of P. heterocephalus see Caron et al. (2008).
Diagnosis
Body flattened dorsoventrally; black with apical half of
abdominal segment 7 and all segments 8–10 yellowish
(Fig. 15); pair of acute and long frontal processes on head,
ventral face of frontal processes coarsely serrate and basal
distance between the processes ~2 times the basal width of
each one (Fig. 62); V-shaped frontal sulcus incomplete;
antennae short, reaching the apex of elytra and scape with two
long setae near the apex of the dorsal face; labrum with six setae
medially; mandibles with bifurcate apex, internal border
asymmetrical; pronotum with inconspicuous fine punctures
and parallel lateral sides; lateral lobes of aedeagus with
truncate apex on ventral margin (Fig. 162).
Distribution
Colombia (Valle del Cauca), Venezuela (Mérida) and Ecuador
(Pichincha, Cotopaxi and El Oro) (Caron et al. 2008).
Biological notes
Unknown.
Biological notes
Remarks
This species has been found under bark, associated with
fermenting decaying logs. Some specimens were collected at
flight intercept traps or by Berlese extraction (forest litter).
This species was recently redescribed by Caron et al. (2008)
and its abdominal defensive gland complex described and
reported for the first time in Piestinae.
Revision of Piestus with remarks on related genera
Piestus heterocephalus is similar to P. lacordairei, differing
from that species by the characters cited above (see remarks
under P. lacordairei).
Piestus puncticollis Fauvel, 1902
(Figs 16, 63, 117, 118, 164, 165)
Piestus puncticollis Fauvel, 1902: 24 (original description, type locality:
‘Amazones: Ega’); Blackwelder, 1944: 101 (distribution); Herman,
2001b: 1793 (distribution).
Piestus (Piestus) puncticollis: Bernhauer & Schubert, 1910:
7 (distribution); Scheerpeltz, 1952: 286 (characters, distribution).
Type material
Piestus puncticollis Fauvel, 1902. Lectotype (here designated) deposited
in IRSNB, male, with labels: (1) ‘Ega’ [white label, handwritten]; (2)
‘puncticollis’ [white label, handwritten]; (3) ‘Ex-Typis’ [white label,
printed in red]; (4) ‘R.I.Sc.N.B. 17.479/Piestus/Coll. et det. A. Fauvel’
[white label, first and third lines printed in black, and second line
handwritten]. Three paralectotypes deposited in IRSNB, males, with
labels: (1) ‘Ega’ [white label, handwritten]; (2) ‘Ex-Typis’ [white label,
printed in red]; (3) ‘R.I.Sc.N.B. 17.479/Piestus/puncticollis Fvl./Coll. et
det. A. Fauvel’ [white label, first and fourth lines printed in black, second
and third lines handwritten]. Note: In the original description, Fauvel (1902)
did not specify how many specimens he observed. We received from IRSNB
four males with type labels.
Redescription
BL: 5.5–5.6 mm, BW: 1.4–1.5 mm. Body strongly flattened
dorsoventrally; entirely light brown to reddish dark brown,
with head and abdomen darker (Fig. 16). Dorsal integument
of head and pronotum with inconspicuous fine punctures and
some moderately sized punctures distributed on pronotal disc,
undulate microstriae only on lateral regions (Fig. 63); striae of
elytra with moderately sized punctures; metaventrite, on lateral
regions, and abdominal tergites on anterolateral regions with
moderately sized punctures.
Male. Head with front slightly deflected (Fig. 63); V-shaped
frontal sulcus complete, sometimes curved arms slightly joined
medially; anterior angles curved, prominent and with median
fovea at base; eyes with line of three moderately sized
punctures with long setae on basal half of dorsal margin, basal
the longest. Antennae reaching apex of abdomen (Fig. 16);
scape with slightly enlarged area and long setae on basal half
of exposed dorsal face; scape shorter than antennomere 2 and
3 combined; antennomeres 4–11 oblong shape. Mandibles with
bifurcate apex (Fig. 117); each with dorsal tooth shorter than
ventral, projected dorsally and forming perpendicular angle
(Fig. 118); internal border of mandibles symmetrical, each
with strong emargination near apex and two acute teeth, basal
smaller and projected anteriorly and apical more acute and
projected almost posteriorly. Maxillary palpus with palpomere
4 shorter than 2 and 3 combined. Pronotum wider than long
(PW/PL = 1.50) (Fig. 63); anterior angles projected; longitudinal
median sulcus conspicuous; longitudinal sulcus of posterior
angles developed and reaching basal quarter of pronotum;
abrupt constriction at basal quarter and, anterior to it, in each
side with slight emargination; apical three-quarters with parallel
sides. Femur of anterior legs with short, hard setae on groove of
ventral margin. Elytra as long as wide (EL/BW = 1.05); stria 6
Invertebrate Systematics
513
at basal half and stria 7 absent or slightly developed only on
apex; striae narrower than interstriae. Abdominal segments 3–6
parallel sides; sternite 3 with conspicuous transverse carina;
tergite 9 with short ventral struts; tergite 10 with two pairs of
long setae on apex, apical the longest. Median lobe of aedeagus
with bulbous base in ventral view and not curved apex in lateral
view (Fig. 164); lateral lobes exceeding apex of median lobe by
one-quarter of its length in lateral view; internal sclerites as in
Fig. 165.
Female. Unknown.
Distribution
Brazil (Amazonas).
Biological notes
Unknown.
Remarks
Piestus puncticollis is similar to P. capricornis, from which it
is easily distinguished by the head without a pair of frontal
processes (Fig. 63), antennal scape with a slightly enlarged
area and long setae on the basal half of the dorsal face, and
mandibles with bifurcate apex, each with dorsal tooth projected
dorsally and forming a perpendicular angle with the ventral tooth,
internal border symmetrical (Figs 117, 118).
Piestus capricornis has a pair of acute and short frontal
processes (Fig. 64), antennal scape with two long setae at the
middle of the dorsal face, mandibles with internal border
asymmetrical and right mandible with trifurcate apex (Fig. 119).
Fauvel (1902), when describing P. puncticollis, cited a
slight sexual dimorphism of the antennae. However, all type
specimens examined from IRSNB are males and probably
Fauvel considered a small male as female. Polymorphism in
males of Piestus, mainly related to sexual dimorphism, is
common and it was observed in other species.
The females of P. puncticollis are still unknown, as only the
type material has been studied.
Piestus capricornis Laporte, 1835
(Figs 17, 64, 109, 119, 166, 167, 253)
Piestus capricornis Laporte, 1835: 129 (original description, type
locality: ‘Cayenne’); Erichson, 1840: 833 (characters, distribution);
Fauvel, 1864: 22 (characters, distribution); Fleutiaux & Sallé, 1889:
382 (distribution); Blackwelder, 1944: 100 (distribution); Herman,
2001b: 1789 (distribution).
Piestus (Zirophorus) capricornis: Bernhauer & Schubert, 1910: 6
(distribution); Blackwelder, 1943: 50 (characters, distribution,
notes); Scheerpeltz, 1951: 8 (characters, distribution); Scheerpeltz,
1952: 284 (characters, distribution).
Piestus capricornis variety muticus Fauvel, 1902: 22 (original
description, type locality: ‘Para: Benevides, 2; Marco da Legua, 30 );
Blackwelder, 1944: 100 (distribution); Herman, 2001b: 1790
(distribution).
Piestus (Zirophorus) capricornis variety muticus: Bernhauer & Schubert,
1910: 6 (distribution); Scheerpeltz, 1951: 8 (characters, distribution);
Scheerpeltz, 1952: 284 (characters, distribution).
514
Invertebrate Systematics
Piestus frontalis Sharp, 1876: 405 (original description, type locality:
‘Ega’). Blackwelder, 1944: 100 (distribution); Herman, 2001b: 1791
(distribution). New synonym.
Piestus (Zirophorus) frontalis: Bernhauer & Schubert, 1910: 6
(distribution); Scheerpeltz, 1951: 7 (characters, distribution);
Scheerpeltz, 1952: 284 (characters, distribution).
Type material
Piestus capricornis Laporte, 1835. Syntype deposited in BMNH, sex
undetermined [damaged apex of the left antenna], with labels: (1) ‘Piestus
capricornis Lap. Type/ex coll. Chevrolat./Guadeloupe. L‘Herminier’ [white
label, together with the specimen, handwritten]; (2) ‘Type’ [circle white label
with border red, printed in black]; (3) ‘Sharp Coll./1905-313.’ [white label,
printed in black]; (4) ‘Piestus/capricornis/Er. p. 833, H. type/Guadalpia/
pointe a pitre D. Sherminier?’ [green label, handwritten]. Note: In the
original description Laporte (1835) did not specify how many specimens
were observed. We received only one specimen from BMNH with type label,
which it is labelled as coming from Guadeloupe, and not from Cayenne
(French Guiana), the original type locality. However, we are considering this
as type following Blackwelder (1943), that already commented it.
Piestus capricornis variety muticus Fauvel, 1902. Lectotype (here
designated) deposited in IRSNB, male, with labels: (1) ‘Maroc de/Legua
(Para)/30 [white label, handwritten]; (2) ‘v. muticus Fvl.’ [white label,
handwritten]; (3) ‘Coll. et det. A. Fauvel/Piestus/capricornis Cast./R I.Sc.
N.B. 17.479’ [white label, the first and fourth lines printed in black, second
and third lines handwritten]. Note: In the original description Fauvel (1902)
did not specify how many specimens he observed, but he listed two localities
(see type locality above). We received from IRSNB one male from ‘Maroc de
Legua (Para)/30 which is here considered as lectotype.
Piestus frontalis Sharp, 1876. Holotype deposited in BMNH, sex
undetermined, with labels: (1) ‘Piestus/frontalis/Amazons Type/D.S.’
[white label, together with the specimen, handwritten]; (2) ‘Holo-/type’
[circle white label with border red, printed in black]; (3) ‘Ega.’ [circle
green label, handwritten]; (4) ‘S. America:/Brazil.’ [white label, printed in
black]; (5) ‘Sharp Coll./1905-313.’ [white label, printed in black]; (6)
‘Holotype/Piestus/frontalis/Sharp, 1876/det. R.G. Booth 2008’ [white
label, the four first lines handwritten and the last one printed in black].
Note: In the original description Sharp (1876) specified only one specimen
observed.
Additional material
See Appendix 2.
E. Caron et al.
face; scape shorter than antennomere 2 and 3 combined;
antennomeres 4–11 oblong shape. Labrum with six or seven
setae medially, equal in length; each lateral third, antero-internal
seta with same length to postero-external seta (Fig. 109).
Mandibles with bifurcate apex in left and trifurcate in right
(Fig. 119); dorsal tooth a little shorter or equal to ventral;
internal border asymmetrical, each with one acute tooth at
middle and, anterior to it, in right mandible with an area that
resemble a tooth, absent in left mandible. Maxillary palpus with
palpomere 4 shorter than 2 and 3 combined. Mentum 2 times
wider than long. Pronotum wider than long (PW/PL = 1.66)
(Fig. 64); anterior angles projected; longitudinal median sulcus
conspicuous; longitudinal sulcus of posterior angles slightly
developed and reaching basal quarter of pronotum; abrupt
constriction at basal quarter and, anterior to it, in each side
with a slight emargination; apical three-quarters with parallel
sides. Femur of anterior legs with short, hard setae on groove of
ventral margin. Elytra as long as wide (EL/BW = 0.99); stria 6 at
basal half and stria 7 absent; striae narrower than interstriae.
Abdominal segments 3 and 4 slightly narrower than 5–7; sternite
3 with conspicuous transverse carina; tergite 9 with short ventral
struts; tergite 10 with two pairs of long setae on apex, apical the
longest. Median lobe of aedeagus with bulbous base in
ventral view and not curved apex in lateral view (Fig. 166);
lateral lobes exceed a little apex of median lobe in lateral view;
internal sclerites as in Fig. 167.
Female. Similar to male. Ovipositor as in P. sulcatus.
Spermathecal duct ~2 times the length of spermatheca, with
sclerotised basal region (Fig. 253); spermatheca with basal half
suddenly narrower than apical (Fig. 253).
Distribution
In the current study, the species was examined from Guadeloupe,
Peru (Loreto) and Brazil (Amazonas, Pará and Rondônia).
Herman (2001b) listed P. capricornis also from Venezuela,
Suriname and French Guiana.
Biological notes
Unknown.
Redescription
BL: 8.2–9.5 mm, BW: 2.2–2.4 mm. Body strongly flattened
dorsoventrally; entirely reddish brown, sometimes with head
and abdomen darker (Fig. 17). Dorsal integument of head and
pronotum with inconspicuous fine punctures and some
moderately sized punctures distributed on pronotal disc,
undulate microstriae only on lateral regions (Fig. 64); striae of
elytra with moderately sized punctures; metaventrite, on lateral
regions, and abdominal tergites, on anterolateral regions, with
moderately sized punctures.
Male. Head with pair of acute and short frontal processes
(Fig. 64), each shorter than scape, and basal distance between
processes ~3 times basal width of each one; V-shaped frontal
sulcus incomplete, curved arms not joined medially; anterior
angles curved and prominent, and fovea at base; eyes with
two moderately sized punctures with long setae on basal half
of dorsal margin, basal the longest. Antennae reaching apex of
abdomen (Fig. 17); scape with two long setae at middle of dorsal
Remarks
Piestus capricornis is easily distinguished from P. puncticollis
by the characters under remarks of P. puncticollis.
Piestus planatus (Sharp, 1887)
(Figs 18, 65, 168, 169, 254)
Zirophorus planatus Sharp, 1887: 711 (original description, type
localities: ‘Mexico, Cordova; British Honduras, R. Sarstoon;
Guatemala, Sinanja; Nicaragua, Chontales; Panama, Volcan de
Chiriqui 2000 to 3000 feet’).
Piestus (Zirophorus) planatus: Bernhauer & Schubert, 1910: 6
(distribution); Scheerpeltz, 1951: 7 (characters, distribution);
Scheerpeltz, 1952: 283 (characters, distribution).
Piestus planatus: Blackwelder, 1944: 101 (distribution); Herman,
2001b: 1793 (distribution); Navarrete-Heredia et al., 2002: 209
(distribution).
Revision of Piestus with remarks on related genera
Type material
Syntype deposited in FMNH, sex undetermined, labels: (1) ‘Chontales,/
Nicaragua./Janson.’ [white label, printed in black]; (2) ‘B.C.A. Col. I. 2./
Zirophorus/planatus,/Sharp.’ [white label, printed in black]; (3) ‘Sharp Coll./
1905.-313’ [white label, printed in black]; (4) ‘Chicago Nat. Hist. Mus./
(ex. D. Sharp Colln./by exchange with/Brit. Mus. Nat. Hist.)’ [white
label, printed in black]. Note: There are 11 syntypes (not examined)
deposited in BMNH: ‘5 from Chontales, 3 from Rio Sarstoon, 1 from
Cordova, 1 from Sinanja, 1 from V. de Chiriqui’ (Dr Roger Booth,
BMNH, pers. comm.).
Additional material
See Appendix 2.
Invertebrate Systematics
515
Female. Similar to male. Ovipositor as in P. sulcatus;
spermathecal duct ~2 times length of spermatheca, with
sclerotised basal region (Fig. 255); spermatheca with basal half
suddenly narrower than apical (Fig. 255).
Distribution
In the current study the species was examined from Mexico
(Chiapas, Veracruz, Guerrero and Oaxaca), Guatemala (Alta
Verapaz), Nicaragua (Río San Juan), Costa Rica (Alajuela and
Limón), Panama (Colón and Darién) and Ecuador (Esmeraldas).
Navarrete-Heredia et al. (2002) listed P. planatus also from
Belize.
Biological notes
Redescription
BL: 8.2–10.8 mm, BW: 2.4–2.9 mm. Body strongly flattened
dorsoventrally; entirely black (Fig. 18). Dorsal integument of
head and pronotum with undulate microstriae and inconspicuous
fine punctures, some moderately sized punctures distributed on
pronotal disc (Fig. 65); striae of elytra with moderately sized
punctures; metaventrite, on lateral regions, and abdominal
tergites, on anterolateral regions, with moderately sized
punctures.
Male. Head with pair of acute and long frontal processes,
each longer than scape, weakly divergent to apex, and basal
distance between processes ~3 times basal width of each one
(Fig. 65); frontal anterior margin with two prominent areas at
middle; V-shaped frontal sulcus complete; anterior angles
curved and prominent, short median longitudinal sulcus with
fovea at base; eyes with two moderately sized punctures with
long setae on basal half of dorsal margin of eyes, basal the
longest. Antennae almost reaching apex of abdomen (Fig. 18);
scape with two long setae at middle of dorsal face; scape
shorter than antennomere 2 and 3 combined; antennomeres
4–11 oblong shape. Labrum with eight or nine setae medially,
equal in length; each lateral third, antero-internal seta with same
length to postero-external seta. Mandibles strongly projected
anteriorly; with bifurcate apices; each with dorsal tooth longer
than ventral; internal border symmetrical, each with one
acute tooth at middle. Maxillary palpus with palpomere 4
shorter than 2 and 3 combined. Mentum 2 times wider than
long. Pronotum wider than long (PW/PL = 1.54) (Fig. 65);
anterior angles projected; longitudinal median sulcus
conspicuous; longitudinal sulcus of posterior angles slightly
developed and reaching basal quarter of pronotum; abrupt
constriction at basal quarter and, anterior to it, in each side
with a slight emargination; apical three-quarters with parallel
sides. Femur of anterior legs with short, hard setae on groove of
ventral margin. Elytra somewhat shorter than wide (EL/
BW = 0.93); stria 6 at basal half and stria 7 absent; striae
narrower than interstriae. Abdominal segments 3 and 4 slightly
narrower than 5–7; sternite 3 with conspicuous transverse carina;
tergite 9 with short ventral struts; tergite 10 with two pairs of
long setae on apex, apical the longest. Median lobe of aedeagus
with bulbous base in ventral view and not curved apex in lateral
view (Fig. 168); lateral lobes exceeding apex of median lobe by
one-quarter of its length in lateral view; internal sclerites as in
Fig. 169.
Piestus planatus has been observed only under bark or associated
with decaying logs.
Remarks
This species is very similar to P. spinosus mainly by the dorsal
teeth of mandibles longer than the ventral teeth (Figs 65, 66, 120).
However, P. planatus differs by the body being entirely black
(Fig. 18), frontal processes divergent to the apex (Fig. 65) and
front with two conspicuously prominent areas at the middle
(Fig. 65).
Piestus spinosus has head, pronotum and elytra reddish
brown and abdomen reddish dark brown to black (Fig. 19),
frontal processes not divergent to the apex (Fig. 66), and
front with two inconspicuous prominent areas at the middle
(Fig. 66).
Piestus spinosus (Fabricius, 1801)
(Figs 19, 69, 120, 170, 171, 254)
Cucujus spinosus Fabricius, 1801: 93 (original description, type locality:
‘America meridionali’).
Piestus spinosus: Laporte, 1835: 128 (characters, distribution); Erichson,
1840: 832 (characters, distribution); Fauvel, 1864: 22 (characters,
distribution); Sharp, 1876: 405 (distribution); Blackwelder, 1944:
101 (distribution); Herman, 2001b: 1794 (distribution); Newton
et al., 2005: 37 (distribution).
Piestus (Zirophorus) spinosus: Bernhauer & Schubert, 1910: 6
(distribution); Scheerpeltz, 1933: 993 (mention); Scheerpeltz, 1951:
5 (mention), 7 (characters, distribution); Scheerpeltz, 1952: 283
(characters, distribution).
Piestus oxytelinus Perty, 1830: 33 (original description, type locality:
‘Provincia Piauhiensi’). - As synonym of P. spinosus: Erichson, 1840:
832; Fauvel, 1864: 22; Bernhauer & Schubert, 1910: 6; Blackwelder,
1944: 101 (error: Perty, 1834: 33); Herman, 2001b: 1794; Newton et al.,
2005: 69. Note: Scheerpeltz (1952: 283) and Scheerpeltz (1960: 66) list
this name as valid and attributed it to Laporte, 1835 (error: Laporte,
1834: 129). However, the name oxytelinus sensu Laporte, 1835 is a
misidentification of P. bicornis (Herman 2001b). For more details see
note under P. bicornis.
Type material
Cucujus spinosus Fabricius, 1801. Syntype deposited in ZMUC, sex
undetermined, with labels: (1) ‘TYPE’ [red label, printed in black]; (2) ‘C?:
spinosus/illegible/Cucujus’ [white label, handwritten]. Note: In the original
516
Invertebrate Systematics
description Fabricius (1801) did not specify how many specimens were
observed. We received from ZMUC only one specimen with type label
which is here considered as a syntype.
Piestus oxytelinus Perty, 1830. Not found.
Additional material
See Appendix 2.
Redescription
BL: 7.8–10.5 mm, BW: 2.2–3.0 mm. Body strongly flattened
dorsoventrally; head, pronotum and elytra reddish brown and
abdomen reddish dark brown to black, sometimes head darker
than pronotum and elytra (Fig. 19). Dorsal integument of head
and pronotum with undulate microstriae and inconspicuous
fine punctures, some moderately sized punctures distributed on
pronotal disc (Fig. 66); striae of elytra with moderately sized
punctures; metaventrite, on lateral regions, and abdominal
tergites, on anterolateral regions, with moderately sized
punctures.
Male. Head with pair of acute and long frontal processes,
each longer than scape, and basal distance between processes
~3 times basal width of each one (Fig. 66); frontal anterior
margin with two prominent regions at middle, sometimes
inconspicuous; V-shaped frontal sulcus complete; anterior
angles curved and prominent, and short median longitudinal
sulcus with fovea at base; eyes with two moderately sized
punctures with long setae on basal half of dorsal margin of
eyes, basal the longest. Antennae almost reaching apex of
abdomen (Fig. 19); scape with two long setae at middle of
dorsal face; scape shorter than antennomere 2 and 3 combined;
antennomeres 4–11 oblong shape. Labrum with six, seven or
eight setae medially, equal in length; each lateral third, anterointernal seta with same length to postero-external seta.
Mandibles strongly projected; bifurcate apex (Fig. 120);
each with dorsal tooth longer than ventral; internal border
symmetrical, each with one acute tooth at middle. Maxillary
palpus with palpomere 4 shorter than 2 and 3 combined. Mentum
2 times as wide as long. Pronotum wider than long (PW/
PL = 1.50) (Fig. 66); anterior angles projected; longitudinal
median sulcus conspicuous; longitudinal sulcus of posterior
angles slightly developed and reaching basal quarter of
pronotum; abrupt constriction at basal quarter and, anterior to
it, in each side with slight emargination; apical three-quarters
with parallel sides. Femur of anterior legs with short, hard setae
on groove of ventral margin. Elytra somewhat shorter than
wide (EL/BW = 0.90); stria 6 at basal half and stria 7 absent;
striae narrower than interstriae. Abdominal segments 3 and 4
slightly narrower than 5–7; sternite 3 with conspicuous transverse
carina; tergite 9 with short ventral struts; tergite 10 with two
pairs of long setae on apex, apical the longest. Median lobe of
aedeagus with bulbous base in ventral view and not curved apex
in lateral view (Fig. 170); lateral lobes exceeding the apex of
median lobe by one-quarter of its length in lateral view; internal
sclerites as in Fig. 171.
Female. Similar to male. Ovipositor as in P. sulcatus;
spermathecal duct ~2 times length of spermatheca, with
sclerotised basal region (Fig. 255); spermatheca with basal half
suddenly narrower than apical (Fig. 255).
E. Caron et al.
Distribution
In the current study, the species was examined from Venezuela
(Bolívar), French Guiana (Saint-Laurent-du Maroni and
Cayenne), Ecuador (Napo and Pastaza), Peru (Loreto,
Huánuco, Lima, Junín, Ucayali and Cusco), Bolivia
(Cochabamba and Santa Cruz) and Brazil (Amazonas, Pará,
Rondônia and Mato Grosso). Herman (2001b) listed
P. spinosus also from Colombia, Suriname and Guyana.
Biological notes
Piestus spinosus has been found under bark or in leaf litter near
decaying logs. One specimen was collected in a flight intercept
trap and another was labelled as ‘beating palm branches’. In
Bolivia one specimen was observed in Ficus sp.
Remarks
Piestus spinosus is very similar P. planatus but differs by
characters listed under remarks of P. planatus.
Piestus longipennis Fauvel, 1864
(Figs 20, 67, 172–175, 256)
Piestus longipennis Fauvel, 1864: 20 (original description, type locality:
‘Colombie’); Blackwelder, 1944: 100 (distribution, error: Fauvel 1865:
24); Herman, 2001b: 1792 (distribution); Newton et al., 2005: 37
(distribution).
Piestus (Zirophorus) longipennis: Bernhauer & Schubert, 1910: 6
(distribution, error Fauvel, 1865: 24); Scheerpeltz, 1951: 6
(characters, distribution, error: Fauvel, 1865: 25); Scheerpeltz, 1952:
282 (characters, distribution, error: Fauvel, 1865: 25).
Type material
Holotype deposited in IRSNB, sex undetermined, with labels: (1)
‘Colombie’ [white label, Fauvel’s handwritten]; (2) ‘longipennis/Fvl.’
[white label, Fauvel’s handwritten]; (3) ‘R.I.Sc.N.B. 17.479/Piestus/Coll.
et det. A. Fauvel’ [white label, first and third lines printed in black, and
second line handwritten]; (4) ‘Type’ [red label, printed in black]. Note: In the
original description Fauvel (1864) specified only one specimen observed.
Additional material
See Appendix 2.
Redescription
BL: 9.2–10.5 mm, BW: 2.4–2.8 mm. Body flattened
dorsoventrally; entirely dark brown to black (Fig. 20). Dorsal
integument of head and pronotum with inconspicuous fine
punctures, undulate microstriae only on lateral regions
(Fig. 67); striae of elytra unpunctured; metaventrite, on lateral
regions, and abdominal tergites, on anterolateral regions, with
moderately sized punctures.
Male. Head with pair of acute and long frontal processes
each longer than scape, and basal distance between processes
the same or narrower than basal width of each one (Fig. 67); Vshaped frontal sulcus complete; anterior angles curved and
weakly prominent; eyes with line of three moderately sized
punctures with long setae on basal half of dorsal margin of
eyes, equal in length. Antennae almost reaching apex of elytra
(Fig. 20); scape with some few long setae at middle of dorsal face;
antennomeres 3–6 with some long setae on dorsal face; scape
subequal to antennomere 2 and 3 combined; antennomeres 4–11
Revision of Piestus with remarks on related genera
oblong shape. Labrum with six setae medially, internal the
longest; on each lateral third, antero-internal seta longer than
postero-external seta. Mandibles strongly projected with
bifurcate apices; each with dorsal tooth shorter than ventral;
internal border with one acute tooth at middle and, anterior to
it, in right mandible with another acute tooth, absent in left
mandible. Maxillary palpus with palpomere 4 shorter than 2
and 3 combined. Mentum 2 times wider than long. Pronotum
wider than long (PW/PL = 1.38) (Fig. 67); anterior angles weakly
projected; longitudinal median sulcus conspicuous; longitudinal
sulcus of posterior angles not developed; abrupt constriction at
basal quarter; apical three-quarters with parallel sides. Femur of
anterior legs with long hard setae on groove of ventral margin.
Elytra somewhat longer than wide (EL/BW = 1.08); stria 6 at
basal half, sometimes inconspicuous, and stria 7 absent; striae
narrower than interstriae. Abdominal segments 3–6 with parallel
sides; sternite 7 with longitudinal set of long setae near of each
lateral margin; tergite 9 with short ventral struts; tergite 10 with
two pairs of long setae on apex, basal the longest. Median lobe of
the aedeagus with bulbous base in ventral view and not curved
apex in lateral view (Fig. 172); apex of median lobe strongly
emarginate in ventral view (Fig. 174); lateral lobes almost
reaching apex of median lobe in lateral view; internal sclerites
as in Figs 173 and 175.
Female. Similar to male. Ovipositor as in P. fronticornis;
spermathecal duct ~1.5 times length of spermatheca, with
sclerotised basal region (Fig. 256); spermatheca as in Fig. 256.
Distribution
In the current study the species was examined from Colombia
(Valle del Cauca) and Ecuador (Carchi, Pichincha, Napo,
Cotopaxi and Tungurahua). Herman (2001b) listed this species
also from Venezuela.
Biological notes
Piestus longipennis is only known from under bark of decaying
logs.
Remarks
Piestus longipennis is similar to P. zischkai and easily
separated from that by the frontal processes on head of male
not converging to the apex (Fig. 67) and lateral lobes of aedeagus
never exceeding the apex of median lobe in lateral view
(Fig. 172). Piestus zischkai has frontal processes of male
converging to the apex (Fig. 68) and the lateral lobes
exceeding the apex of median lobe (Fig. 176).
Piestus longipennis, as well as P. zischkai, differs from the
other closely related species (P. bicornis, P. fronticornis and
P. validus) mainly by the body being entirely black and the
presence of a bulbous base of the median lobe of aedeagus in
ventral view.
Piestus zischkai Scheerpeltz, 1951
(Figs 21, 68, 69, 176–178)
Piestus (Zirophorus) zischkai Scheerpeltz, 1951: 4 (original description,
type locality: ‘Yungas del Palmar’, ‘200 m’); Scheerpeltz, 1952: 282
(characters, distribution).
Piestus zischkai: Herman, 2001b: 1795 (distribution).
Invertebrate Systematics
517
Type material
Piestus (Zirophorus) zischkai Scheerpeltz, 1951. Lectotype (here
designated) deposited in NMW, male [damaged specimen: without
anterior and left posterior tarsi], with labels: (1) ‘<’ [white label, printed
in black]; (2) ‘Bolivia: 2000 m/Yungas del Palmar/15. III. 1951/leg R.
Zischka’ [white label, first (except ‘Bolivia’), second and third lines
handwritten, ‘Bolivia’ in the first line and fourth line printed in black]; (3)
‘ex coll./Scheerpeltz’ [blue label, printed in black]; (4) ‘TYPUS/Piestus/
zischkai/O. Scheerpeltz’ [red label, the first and fourth lines printed in
black, second and third handwritten]; (5) ‘Zischkai/Scheerp.’ [green label,
handwritten]. Paralectotype deposited in NMW, female [damaged specimen:
without last tarsomere of left median leg], with labels: (1) ‘,’ [white label,
printed in black]; (2) ‘Bolivia: 2000 m/Yungas del Palmar/15. III. 1951/leg R.
Zischka’ [white label, first (except ‘Bolivia’), second and third lines
handwritten, ‘Bolivia’ in the first line and fourth line printed in black]; (3)
‘ex coll./Scheerpeltz’ [blue label, printed in black]; (4) ‘TYPUS/Piestus/
zischkai/O. Scheerpeltz’ [red label, the first and fourth lines printed in black,
second and third handwritten]. Note: In the original description Scheerpeltz
(1951) cited two males and one female observed. However, we received from
NMW one male and one female with type labels.
Additional material
See Appendix 2.
Redescription
BL: 14.5–15.8 mm, BW: 2.3–2.5 mm. Body flattened
dorsoventrally; entirely dark brown to black (Fig. 21). Dorsal
integument of head and pronotum with inconspicuous fine
punctures, undulate microstriae only on lateral regions
(Fig. 68); striae of elytra unpunctured; metaventrite, on lateral
regions, and abdominal tergites, on anterolateral regions, with
moderately sized punctures.
Male. Head with pair of acute and long frontal processes
each longer than scape, strongly converge to apex, and basal
distance between processes the same or narrower than basal
width of each one (Fig. 68); V-shaped frontal sulcus complete;
anterior angles curved and weakly prominent; eyes with line of
three moderately sized punctures with long setae on basal half of
dorsal margin, equal in length. Antennae reaching apex of elytra
(Fig. 21); with some few long setae at middle of dorsal face;
antennomeres 3–6 with some long setae on dorsal face; scape
subequal to antennomere 2 and 3 combined; antennomeres 4–11
oblong shape. Labrum with six setae medially, internal the
longest; on each lateral third, antero-internal seta with same
length to postero-external seta. Mandibles with bifurcate
apices; each with ventral tooth anteriorly weakly projected and
dorsal tooth shorter than ventral; internal border with one acute
tooth at middle and, anterior to it, in right mandible with other
acute tooth, absent in left mandible. Maxillary palpus with
palpomere 4 shorter than 2 and 3 combined. Mentum 2 times
wider than long. Pronotum wider than long (PW/PL = 1.25)
(Fig. 68); anterior angles projected; longitudinal median sulcus
conspicuous; longitudinal sulcus of posterior angles not
developed; abrupt constriction at basal quarter; apical threequarters with parallel sides. Femur of anterior legs with long
hard setae on groove of ventral margin. Elytra somewhat longer
than wide (EL/BW = 1.10); stria 6 at basal half, sometimes
inconspicuous, and stria 7 absent; striae narrower than
interstriae. Abdominal segments 3–6 with parallel sides;
518
Invertebrate Systematics
sternite 7 with longitudinal set of long setae near of each lateral
margin; tergite 9 with short ventral struts; tergite 10 with two pairs
of short setae on apex, equal in length. Median lobe of aedeagus
with bulbous base in ventral view and not curved apex in lateral
view (Fig. 176); apex of median lobe emarginate in ventral view
(Fig. 178); lateral lobes exceed little the apex of median lobe in
lateral view; internal sclerites as in Fig. 177.
Female. Similar to male, except for: frontal processes of
head not converging to apex and internal margins are straight.
Ovipositor as in P. fronticornis; spermathecal duct with
sclerotised basal region (length unknown); spermatheca as in
P. longipennis.
Distribution
In the current study the species was examined from Ecuador
(Tungurahua and Zamora-Chinchipe), Peru and Bolivia.
Biological notes
E. Caron et al.
of P. bicornis (‘Cet insecte n’est peut-être qu’une variété du précédent’
Laporte 1835: 128).
Piestus oxytelinus: Laporte, 1835: 128. Scheerpeltz, 1952: 283 (error:
Laporte, 1834: 129); Scheerpeltz, 1960: 66 (error: Laporte, 1834: 129);
Newton et al., 2005: 37 (error: Laporte 1835: 129). Note: the name
P. oxytelinus by Laporte (1835) is not an available name, but rather a
misidentification of P. oxytelinus Perty, 1830, which is a junior
synonym of P. spinosus (see above, P. spinosus).
Type material
Oxytelus bicornis Olivier, 1811. Neotype (here designated) deposited in
FMNH, sex undetermined, with labels: ‘15.544’ [white label, handwritten],
‘Bras.S.Catharina/illegible VIII 10/lg. Luderwaldd.’ [white label,
handwritten], ‘bicornis Ol./Ihering./det. Bernh.’ [white label, the first two
lines handwritten, third line printed in black], ‘Chicago NHMus/M.
Bernhauer/Collection’ [white label, printed in black]. Note: In the original
description, Olivier (1811) did not specify how many specimens were
observed. The type material has not been found and probably it is lost.
Zirophorus striatus Guérin-Méneville, 1829. Not found.
There is one specimen of P. zischkai with a note that it was
collected using a Malaise trap.
Additional material
Remarks
Redescription
This species is very similar to P. longipennis, from which it is
easily separated by characters cited under remarks of
P. longipennis.
BL: 7.3–9.3 mm, BW: 2.1–2.4 mm. Body flattened
dorsoventrally; brown to black with apical half of segment 7
and segments 8–10 of abdomen yellowish brown, or head
and abdomen dark brown to black with pronotum and elytra
reddish brown, also with apical half of segment 7 and segments
8–10 of abdomen yellowish brown (Figs 22, 23). Dorsal
integument of head and pronotum with inconspicuous fine
punctures, undulate microstriae only on lateral regions; striae
of elytra unpunctured (Fig. 70); metaventrite, on lateral regions,
and abdominal tergites, on anterolateral regions, with moderately
sized punctures.
Male. Head with pair of acute and long frontal processes,
each longer than scape, and basal distance between processes
the same or narrower than basal width of each one (Fig. 70); Vshaped frontal sulcus complete; anterior angles curved and
weakly prominent; eyes with a line of three moderately sized
punctures with long setae on basal half of dorsal margin, equal in
length. Antennae almost reaching apex of elytra (Figs 22, 23);
with some long setae at middle of the dorsal face; antennomeres
3–6 with some long setae on dorsal face; scape subequal to
antennomere 2 and 3 combined; antennomeres 4–11 oblong
shape. Labrum with six setae medially, internal the longest;
each lateral third, antero-internal seta with same length to
postero-external seta. Mandibles strongly projected; bifurcate
apices; each with dorsal tooth shorter than the ventral; internal
border with one acute tooth at middle and, anterior to it, in right
mandible with other acute tooth, absent in left mandible.
Maxillary palpus with palpomere 4 shorter than 2 and 3
combined. Mentum 2 times wider than long. Pronotum wider
than long (PW/PL = 1.40) (Fig. 70); anterior angles weakly
projected; longitudinal median sulcus conspicuous;
longitudinal sulcus of posterior angles not developed; abrupt
constriction at basal quarter; apical three-quarters with parallel
sides. Femur of anterior legs with long hard setae on groove of
ventral margin. Elytra as long as wide (EL/BW = 1.04); stria 6 at
Piestus bicornis (Olivier, 1811)
(Figs 22, 23, 70, 150, 179–181, 257)
Oxytelus bicornis Olivier, 1811: 615 (original description; type locality:
not given).
Piestus bicornis: Laporte, 1835, 126 (mention), 128 (characters,
distribution); Erichson, 1840: 831 (characters, distribution); Fauvel,
1864: 19 (characters, distribution); Sharp, 1876: 405 (mention,
distribution); Fauvel, 1901: 70 (mention); Blackwelder, 1944: 100
(distribution, error: species attributed to Laporte, 1834: 128); Herman,
2001b: 1789 (distribution); Newton et al., 2005: 37, 69 (distribution,
synonyms); Caron et al. 2008: 6 (mention).
Zirophorus bicornis: Sharp, 1887: 712 (distribution); Sharp & Muir, 1912
(characters of aedeagus).
Piestus (Zirophorus) bicornis: Bernhauer & Schubert, 1910: 6
(distribution, error: species attributed to Laporte, 1834: 128);
Scheerpeltz, 1951: 6 (characters, distribution, error: species
attributed to Laporte, 1834: 128); Scheerpeltz, 1952: 282
(characters, distribution, error: species attributed to Laporte, 1834:
128).
Zirophorus striatus Guérin-Méneville, 1829: pl. 9, fig. 12 (type locality:
not given). - As synonym of Piestus bicornis: Laporte, 1835: 126, 128;
Erichson, 1840: 832; Fauvel, 1864: 20; Bernhauer & Schubert, 1910: 6;
Blackwelder, 1944: 100 (note: cited as ‘of authors’); Herman, 2001b:
1789; Newton et al., 2005: 69.
Piestus bicornis variety oxytelinus: Laporte, 1835: 128. Erichson, 1840:
832; Fauvel, 1864: 20; Sharp, 1976: 405; Sharp, 1887: 712; Bernhauer
& Schubert, 1910: 6 (error: Laporte, 1834: 129); Blackwelder, 1944:
100 (error: Laporte, 1834: 129); Scheerpeltz, 1951: 5, 6 (error: Laporte,
1834: 129); Newton et al., 2005: 37. Note: Laporte, 1835 did not
describe the species as new, he attributed it to Perty (1830), but probably
it was a misidentification of P. oxytelinus Perty (Herman 2001b).
However, Laporte (1835) cited the similarity between P. bicornis
and P. oxytelinus and suggested that P. oxytelinus could be a variety
See Appendix 2.
Revision of Piestus with remarks on related genera
basal half, sometimes inconspicuous, and stria 7 absent; striae
narrower than interstriae. Abdominal segments 3–6 with parallel
sides; sternite 7 with longitudinal set of long setae near of each
lateral margin; tergite 9 with long ventral struts (Fig. 150);
tergite 10 with two pairs of long setae on apex, basal the
longest. Median lobe of aedeagus without bulbous base in
ventral view and not curved apex in lateral view (Figs 179,
181); apex of median lobe slightly emarginate in ventral view
(Fig. 181); lateral lobes not reaching apex of median lobe in lateral
view; internal sclerites as in Fig. 180.
Female. Similar to male. Ovipositor as in P. fronticornis;
spermathecal duct ~5 times length of spermatheca, without
sclerotised basal region (Fig. 257); spermatheca as in Fig. 257.
Distribution
In the current study, the species was examined from Costa Rica
(Cartago and Limón), Panama (Panamá), Colombia
(Cundinamarca, Meta and Putumayo), Venezuela (Carabobo),
French Guiana (Saint-Laurent-du-Maroni and Cayenne),
Ecuador (Sucumbios, Napo, Pastaza and Morona-Santiago),
Peru (Loreto, San Martín, Huánuco, Junín, Cusco and Madre
de Dios), Bolivia (La Paz, Beni, Cochabamba and Santa Cruz),
Brazil (Acre, Amazonas, Pará, Goiás, Minas Gerais, Espírito
Santo, Rio de Janeiro, São Paulo, Paraná and Santa Catarina) and
Paraguay (Guairá). Herman (2001b) listed P. bicornis also from
Nicaragua, Suriname, Guyana and Argentina.
Biological notes
Piestus bicornis has been found under bark or in leaf litter near
decaying logs. Some specimens were collected in Malaise trap
and others were captured by ‘beating palm branches’. This
species also has been noticed occurring together with
P. validus, P. fronticornis, P. pygmaeus and P. minutus.
The specimens with black pronotum and elytra were only
observed in Brazil (Minas Gerais, Espirito Santo, Rio de Janeiro,
São Paulo and Paraná e Santa Catarina) and Paraguay (Guairá).
The specimens with reddish brown pronotum and elytra were
recorded from Costa Rica to Brazil (Acre, Amazonas, Pará,
Goiás, São Paulo and Paraná).
Invertebrate Systematics
519
tergite 10 with two pairs of long setae on the apex, and apex of
median lobe very thin and sclerite of internal sac inconspicuous
(Figs 182, 183).
Piestus validus has a brown to black body with apical half
of segment 7 and segments 8–10 yellowish (Fig. 25), antennal
scape with many long setae on the apical half of the dorsal face
and antennomeres 3–6 with many long setae on dorsal face,
tergite 9 of male with long ventral struts, tergite 10 with one pair of
long setae on the apex, and with a well-developed apex of
median lobe and sclerite of internal sac (Figs 184, 185).
There are two colour patterns in P. bicornis. Specimens with
black pronotum and elytra (Fig. 22) or specimens with reddish
brown pronotum and elytra (Fig. 23). These patterns were first
observed by Laporte (1835), who misidentified the specimens
with reddish brown pronotum and elytra as P. oxytelinus Perty,
1830 (junior synonym of P. spinosus) and cited its close
similarity to P. bicornis (specimens with black pronotum and
elytra), ‘Cet insecte n’est peut-être qu’une variété du précédent’
(Laporte 1835: 128). Since then, many authors have identified
the specimens with reddish brown pronotum and elytra as a
variety of P. bicornis, P. bicornis variety oxytelinus (Erichson
1840; Fauvel 1864; Sharp 1876, 1887; Scheerpeltz 1951), or as a
valid species, P. oxytelinus Laporte, 1835 (Scheerpeltz 1952,
1960; Newton et al. 2005). However, in this study no significant
morphological differences between the specimens with the
two colour patterns have been found, the morphology of the
aedeagus and spermatheca being the same. Different colour
patterns in the same species were also observed in
P. pennicornis (see below).
Piestus fronticornis (Dalman, 1821), sp. rev. (revalidated
species)
(Figs 24, 71, 110, 121, 122, 130, 133,
143, 154, 182, 183, 258)
Zirophorus fronticornis Dalman, 1821: 373 (original description, type
locality: ‘Kalkoen’). - As synonym of P. bicornis: Laporte, 1835: 126,
128; Erichson, 1840: 832; Fauvel, 1864: 20; Sharp, 1887: 712;
Bernhauer & Schubert, 1910: 6; Blackwelder, 1944: 100; Herman,
2001b: 1789; Newton et al., 2005: 69.
Remarks
Type material
This species is similar to P. fronticornis and P. validus, and can
be separated from them by having a body brown to black
with apical half of segment 7 and segments 8–10 yellowish
brown, or head and abdomen dark brown to black with
pronotum and elytra reddish brown and also with apical half
of segment 7 and segments 8–10 yellowish brown (Figs 22, 23),
antennal scape with a few long setae on the apical half of the
dorsal face and antennomeres 3–6 with a few long setae on dorsal
face, tergite 9 of male with long ventral struts (Fig. 150), tergite 10
with two pairs of long setae on the apex, and with a welldeveloped apex of median lobe and sclerite of internal sac
(Figs 179, 180).
Piestus fronticornis has the following characters: body
brown to black with all of segment 7 and segments 8–10
yellowish (Fig. 24), antennal scape with long setae on the
apical half of the dorsal face and antennomeres 3–6 with long
setae on dorsal face, tergite 9 of male with short ventral struts,
Syntype deposited in NHRS, sex undetermined [damaged specimen: left
antenna without antennomere 6–11; inverted mandibles; without right
anterior and median tarsomeres; without two last tarsomeres of the left
posterior leg; and without left median leg., with labels: (1) ‘Mus./Pay k’
[white label, printed in black]; (2) ‘fronticornis Dalm.’ [white label,
handwritten]; (3) ‘6402/E91 +’ [light blue label, printed in black].
Mouthparts (labrum, maxillae and mentum) are glued on a piece of paper
which is pinned separately and labelleds: ‘6401/E91 +’ [light blue label,
printed in black]; ‘6400/E91 +’ [light blue label, printed in black]; ‘6399/
E91 +’ [light blue label, printed in black]; ‘6398/E91 +’ [light blue label,
printed in black]; ‘buccal parts [handwritten in light green]/det. J. Ferrer
[printed in black]’ [white label]. Note: In the original description Dalman
(1821) did not specify how many specimens he observed, and he did not
know the location of the type locality ‘Kalkoen’. We received from NHRS
one specimen, regarded as a syntype.
Additional material
See Appendix 2.
520
Invertebrate Systematics
E. Caron et al.
Redescription
Biological notes
BL: 7.9–10.8 mm, BW: 1.9–2.8 mm. Body flattened
dorsoventrally; brown to black with all of segment 7 and
segments 8–10 yellowish (Fig. 24). Dorsal integument of head
and pronotum with inconspicuous fine punctures, undulate
microstriae only on lateral regions (Fig. 71); striae of elytra
unpunctured; metaventrite, on lateral regions, and abdominal
tergites on anterolateral regions with moderately sized
punctures.
Male. Head with pair of acute and long frontal processes,
each longer than scape, and basal distance between processes
same or narrower than basal width of each one (Fig. 71);
V-shaped frontal sulcus complete; anterior angles curved and
weakly prominent; eyes with line of three moderately sized
punctures with long setae on basal half of dorsal margin, equal
in length. Antennae almost reaching apex of elytra (Fig. 24);
scape with some few long setae on apical half of dorsal face;
antennomeres 3–6 with some few long setae on dorsal face; scape
subequal to antennomere 2 and 3 combined; antennomeres
4–11 oblong shape. Labrum with six setae medially, internal
the longest; each lateral third, antero-internal seta with same
length to postero-external seta. Mandibles strongly projected
(Figs 71, 121, 122); with bifurcate apices; each with dorsal
tooth shorter than ventral; internal border with one acute tooth
at middle and, anterior to it, in right mandible with other
acute tooth, absent in left mandible. Maxillary palpus with
palpomere 4 shorter than 2 and 3 combined (Fig. 130).
Mentum 2 times wider than long (Fig. 133). Pronotum wider
than long (PW/PL = 1.41) (Fig. 71); anterior angles weakly
projected; longitudinal median sulcus conspicuous;
longitudinal sulcus of posterior angles not developed; abrupt
constriction at basal quarter; apical three-quarters with parallel
sides. Femur of anterior legs with long hard setae on groove
of ventral margin. Elytra as long as wide (EL/BW = 1.01); stria 6
at basal half sometimes inconspicuous, and stria 7 absent;
striae narrower than interstriae. Abdominal segments 3–6
with parallel sides; sternite 7 with longitudinal set of long
setae near of each lateral margin; tergite 9 with short ventral
struts; tergite 10 with two pairs of long setae on apex, basal the
longest. Median lobe of aedeagus without bulbous base in
ventral view and not curved apex in lateral view (Fig. 182);
lateral lobes almost reaching apex of median lobe in lateral view;
internal sclerites as in Fig. 183; apex of median lobe slightly
emarginate in ventral view.
Female. Similar to male. Ovipositor strongly emarginate at
external margin (Fig. 154); spermathecal duct ~1.5 times length of
spermatheca, with sclerotised basal region (Fig. 258);
spermatheca as in Fig. 258.
Piestus fronticornis has been found on or under bark of decaying
logs. Some specimens were collected in flight intercept traps and
others were found on ‘decaying banana, blossoms, leaves’. This
species has been noticed occurring together with P. bicornis,
P. pygmaeus and P. minutus.
Distribution
Nicaragua (Granada and Rio San Juan), Guatemala (Guatemala),
Costa Rica (Guanacaste, Alajuela, Heredia, San José, Limón and
Puntarenas), Panama (Bocas del Toro, Chiriquí, Veraguas,
Colón, Panamá and Darién), Colombia (Cundinamarca and
Valle del Cauca), French Guiana (Saint-Laurent-du-Maroni
and Cayenne), Ecuador (Esmeraldas and Pastaza) and Brazil
(Amazonas, Amapá and Pará).
Remarks
This species is easily separated from those by characters listed in
the remarks under P. bicornis.
Piestus validus Sharp, 1876
(Figs 25, 72, 184, 185, 259)
Piestus validus Sharp, 1876: 404 (original description, type locality:
‘Pebas’); Blackwelder, 1944: 101 (distribution); Herman, 2001b:
1795 (distribution); Caron et al., 2008: 6 (mention).
Piestus (Zirophorus) validus: Bernhauer & Schubert, 1910: 6
(distribution); Scheerpeltz, 1951: 6 (characters, distribution);
Scheerpeltz, 1952: 283 (characters, distribution).
Type material
Three syntypes (not examined) deposited in BMNH (Dr Roger Booth,
BMNH, pers. comm.). Note: In the original description Sharp (1876)
specified three specimens observed.
Additional material
See Appendix 2.
Redescription
BL: 8.3–11.3 mm, BW: 2.2–3.2 mm. Body flattened
dorsoventrally; brown to black with apical half of segment 7
and segments 8–10 yellowish (Fig. 25). Dorsal integument of
dorsum of head and pronotum with inconspicuous fine
punctures, undulate microstriae only on lateral regions
(Fig. 72); striae of elytra unpunctured; metaventrite, on lateral
regions, and abdominal tergites, on anterolateral regions, with
moderately sized punctures.
Male. Head with pair of acute and long frontal processes,
each longer than scape, and basal distance between processes
the same or narrower than basal width of each one (Fig. 72); Vshaped frontal sulcus complete; anterior angles curved and
weakly prominent; eyes with a line of three moderately sized
punctures with long setae on basal half of dorsal margin, equal in
length. Antennae almost reaching apex of elytra (Fig. 25); scape
with many long setae on apical half of the dorsal face;
antennomeres 3–6 with many long setae on dorsal face; scape
subequal to antennomere 2 and 3 combined; antennomeres 4–11
oblong shape. Labrum with six setae medially, internal the
longest; on each lateral third, the antero-internal seta with
same length to postero-external seta. Mandibles strongly
projected; with bifurcate apices; each with dorsal tooth shorter
than ventral; internal border with acute tooth at middle and,
anterior to it, in right mandible with other acute tooth, absent
in left mandible. Maxillary palpus with palpomere 4 shorter
than the 2 and 3 combined. Mentum 2 times wider than long.
Pronotum wider than long (PW/PL = 1.40) (Fig. 72); anterior
angles weakly projected; longitudinal median sulcus
Revision of Piestus with remarks on related genera
conspicuous; longitudinal sulcus of posterior angles
inconspicuous; abrupt constriction at basal quarter; apical
three-quarters with parallel sides. Femur of anterior legs with
long hard setae on groove of ventral margin. Elytra somewhat
longer than wide (EL/BW = 1.05); stria 6 at basal half and
stria 7 absent; striae narrower than interstriae. Abdominal
segments 3–6 parallel sides; sternite 7 with longitudinal set of
long setae near of each lateral margin; tergite 9 with long ventral
struts; tergite 10 with pair of long setae on apex. Median lobe of
aedeagus without bulbous base in ventral view and not curved
apex in lateral view (Fig. 184); apex of median lobe slightly
emarginate in ventral view; lateral lobes not almost reaching
apex of median lobe in lateral view; internal sclerites as in
Fig. 185.
Female. Similar to male. Ovipositor as in P. fronticornis;
spermathecal duct ~2 times length of spermatheca, with
sclerotised basal region (Fig. 259); spermatheca as in Fig. 259.
Distribution
In the current study the species was examined from Colombia
(Meta), Venezuela (Carabobo, Aragua), Ecuador (Napo, Pastaza
and Loja), Peru (Junín, Huánuco, Cusco and Madre de Dios),
Bolivia (La Paz and Cochabamba) and Brazil.
Biological notes
Piestus validus has been found under bark or in litter near
decaying logs. Some specimens were collected in flight
intercept traps and others were captured by ‘beating palm
branches’. This species also has been noticed occurring
together with P. bicornis.
Remarks
This species is very similar to P. bicornis and P. fronticornis
but differs by characters cited above (see remarks under
P. bicornis).
Three syntypes deposited in BMNH were not examined.
However, at the moment, the original description (Sharp 1876)
and a reasonable quantity of specimens identified from many
institutions were considered sufficient for positive identification,
but the study of type material would be advisable for confirming
the identification of this species.
Piestus rossii, sp. nov.
(Figs 26, 73, 186, 187)
Invertebrate Systematics
521
conspicuous fine punctures equally distributed; striae and
interstriae of elytra with inconspicuous fine punctures;
metaventrite, on lateral regions, and abdominal tergites on
anterolateral regions with moderately sized punctures.
Male. Head with front slightly deflected (Fig. 73); V-shaped
frontal sulcus incomplete, curved arms not joined medially;
anterior angles curved, weakly prominent and with median
fovea at base; eyes with line of three moderately sized
punctures with long setae on basal half of dorsal margin of
eyes, equal in length, and additional set, not line, of five
moderately sized punctures with long setae, shorter than and
internal to the other ones. Antennae reaching apex of abdomen
(Fig. 26); scape with slightly enlarged area and long setae on
basal half of exposed dorsal face; scape shorter than antennomere
2 and 3 combined; antennomere 2 with many long setae on
apical half of dorsal face; antennomeres 4–11 oblong shape
and slightly narrowing from antennomere 4 to 11. Mandibles
with dorsal teeth developed, not forming bifurcate apex,
shorter than ventral; internal border asymmetrical, each with
one acute tooth at middle and, anterior to it, in right mandible
with an area that resembles a tooth, absent in left mandible;
external-dorsal margin on basal half of each mandible slightly
carinate. Maxillary palpus with palpomere 4 subequal in length
to 2 and 3 combined. Pronotum wider than long (PW/PL = 1.48)
(Fig. 73); anterior angles weakly projected; longitudinal median
sulcus conspicuous; longitudinal sulcus of posterior angles
slightly developed and reaching basal quarter of pronotum;
abrupt constriction at basal quarter; apical three-quarters with
slightly curved sides. Prosternum with conspicuous ovate set of
fine punctures in median region near apex. Femur of anterior
legs with short, hard setae on groove of ventral margin. Elytra as
long as wide (EL/BW = 0.98); stria 6 at basal half and stria 7
absent; striae narrower than interstriae. Abdominal segments 3–6
parallel sides; tergite 9 with short ventral struts; tergite 10 with
two pairs of long setae on apex. Median lobe of aedeagus with
bulbous base in ventral view and curved apex in lateral view
(Fig. 186); lateral lobes reaching apex of median lobe in lateral
view; internal sclerites as in Fig. 187.
Female. Unknown.
Distribution
Ecuador (Cotopaxi).
Biological notes
Unknown.
Type material
Remarks
Holotype deposited in DZUP, male, labels: (1) ‘Ecuador – Cotopaxi/
Canton Sigchos – Las/Pampas – Otonga Natural/Reserve – 25-28,
VII,2005/W. Rossi, leg.’ [white label, printed in black]; (2) ‘<’ [white
label, printed in black].
Piestus rossii is similar to P. pennicornis, differing from that
species by the single line of three moderately sized punctures
and an additional set, not in line, of five moderately sized
punctures with long setae on basal half of dorsal margin of
eyes, conspicuous fine punctures on pronotum and curved
lateral sides of pronotum (Fig. 73), and curved apex of median
lobe of aedeagus (Fig. 186).
Piestus pennicornis has a single line of three moderately
sized punctures with long setae, inconspicuous fine punctures
and subparallel sides (Figs 74, 75), and uncurved apex of
median lobe (Fig. 188).
Description
BL: 6.8 mm, BW: 1.8 mm. Body flattened dorsoventrally; black
with apical half of segment 7 and segments 8–10 yellowish
(Fig. 26). Dorsal integument of head with somewhat
microgranulate and inconspicuous fine punctures (Fig. 73);
pronotum with undulate microstriae only on lateral areas and
522
Invertebrate Systematics
Etymology
The species is dedicated to Walter Rossi, Italian mycologist,
who collected the holotype and made it available.
Piestus pennicornis Fauvel, 1864
(Figs 27, 74, 75, 123, 124, 188, 189, 260)
Piestus pennicornis Fauvel, 1864: 26 (original description, type locality:
‘Colombie’); Blackwelder, 1944: 101 (distribution, error: Fauvel 1865:
30); Herman, 2001b: 1793 (distribution); Newton et al., 2005: 37
(distribution).
Piestus (Piestus) pennicornis: Bernhauer & Schubert, 1910: 7
(distribution, error: Fauvel, 1865: 30); Scheerpeltz, 1952: 289
(characters, distribution, error: Fauvel 1865: 30).
Piestus plagiatus Fauvel, 1864: 26 (original description, type locality:
‘Nuovo-Friburgo, Brésil’); Blackwelder, 1944: 101 (distribution,
error: Fauvel 1865: 30); Herman, 2001b: 1793 (distribution). New
synonym.
Piestus (Piestus) plagiatus: Bernhauer & Schubert, 1910: 7 (distribution,
error: Fauvel, 1865: 30); Scheerpeltz, 1952: 289 (characters,
distribution, error: Fauvel 1865: 30).
Piestus rectus Sharp, 1876: 406 (original description, type locality:
‘Ega’); Blackwelder, 1944: 101 (distribution); Herman, 2001b: 1794
(distribution). New synonym.
Piestus (Piestus) rectus: Bernhauer & Schubert, 1910: 7 (distribution);
Scheerpeltz, 1952: 288 (characters, distribution).
Piestus pygialis Fauvel, 1902: 23 (original description, type locality:
‘Brésil: Blumenau’); Blackwelder, 1944: 101 (distribution); Herman,
2001b: 1793 (distribution). New synonym.
Piestus (Piestus) pygialis: Bernhauer & Schubert, 1910: 7 (distribution);
Scheerpeltz, 1952: 288 (characters, distribution).
Piestus surinamensis Bernhauer, 1928: 287 (original description,
type locality: ‘Suriname’); Blackwelder, 1944: 101 (distribution);
Herman, 2001b: 1795 (distribution). Note: In the original
description Bernhauer (1928) gives the information about the type
locality at the beginning of the paper for all species described in that
paper. New synonym.
Piestus (Piestus) surinamensis: Scheerpeltz 1933: 994 (distribution);
Scheerpeltz, 1952: 289 (characters, distribution).
Type material
Piestus pennicornis Fauvel, 1864. Lectotype (here designated) deposited
in IRSNB, female, with labels: (1) ‘Colombie’ [white label, handwritten]; (2)
‘pennicornis/FvL.’ [white label, handwritten]; (3) ‘R.I.Sc.N.B. 17.479/
Piestus/Coll. et det. A. Fauvel’ [white label, first and third lines printed in
black and second line handwritten]; (4) ‘Ex-Typis’ [white label, printed in
red]. Note: In the original description Fauvel (1864) specified two specimens,
male and female, studied from Colombia (‘Deux exemplaires < ,’). However,
we received from IRSNB three specimens with type label (‘Ex-Typis’): one
female from ‘Colombie’ (here considered as syntype) and two from ‘Para’,
which we consider additional material.
Piestus plagiatus Fauvel, 1864. Lectotype (here designated) deposited in
IRSNB, female, with labels: (1) ‘N Friburgo’ [white label, handwritten]; (2)
‘plagiatus/FvL.’ [white label/handwritten]; (3) ‘R.I.Sc.N.B. 17.479/Piestus/
Coll. et. det. A. Fauvel’ [white label, printed in black]; (4) ‘Ex-Typis’ [white
label, printed in red]. Note: In the original description Fauvel (1864) specified
two specimens observed. We received from IRSNB only one specimen with
type label.
Piestus rectus Sharp, 1876. Syntype deposited in FMNH, female, labels:
(1) ‘Piestus rectus/amazons./Ind. typ./D.S.’ [white label, together with the
specimen, handwritten]; (2) ‘Ega.’ [green label, handwritten]; (3) ‘S.America:
Brazil.’ [white label with transverse green line, printed in black]; (4) ‘Sharp
Coll./1905-313.’ [white label, printed in black]; (5) ‘Chicago Nat. Hist. Mus./
E. Caron et al.
(ex. D. Sharp Colln./by exchange with/Brit. Mus. Nat. Hist.)’ [white label,
printed in black]. Note: In the original description Sharp (1876) specified four
specimens, ‘probably all females’, studied from ‘Ega’ and more one male
from the same locality. However, we have only seen one syntype deposited
in FMNH. The other 4 syntypes (not examined) are deposited in BMNH
(Dr Roger Booth, BMNH, pers. comm.).
Piestus pygialis Fauvel, 1902. Holotype deposited in IRSNB, female, with
labels: (1) ‘Blumenau/Brésil’ [white label, handwritten]; (2) ‘pygialis/FvL.’
[white label, handwritten]; (3) ‘R.I.Sc.N.B. 17.479/Piestus/Coll. et det.
A. Fauvel’ [white label, first and third line printed in black and second
line handwritten]; (4) ‘Type’ [red label, printed in black]. Note: In the original
description Fauvel (1902) specified only one specimen observed.
Piestus surinamensis Bernhauer, 1928. Holotype deposited in FMNH,
male [damaged specimen: without right antenna and antennomeres 3–11 of
left; head separated of body], with labels: (1) ‘Suriname/Uyttenboogaart/
8-9 1900’ [white label, printed in black]; (2) ‘2/52’ [white label, printed in
black]; (3) ‘Surinamen-/sis Bernh./Typ. un.’ [white label, handwritten];
(4) ‘Piestus/Surinamensis/Bernh. Typus un.’ [light yellow label,
Bernhauer’s handwritten]; (5) ‘Chicago NHMus/M.Bernhauer/Collection’
[white label, printed in black]. Note: In the original description Bernhauer
(1928) specified only one specimen observed.
Additional material
See Appendix 2.
Redescription
BL: 5.0–7.1 mm, BW: 1.3–1.6 mm. Body flattened
dorsoventrally; entirely reddish brown to black with all or
apical half of segment 7 and segments 8–10 yellowish; or head
and abdomen dark brown to black with pronotum and elytra
reddish brown; or head, pronotum and abdomen dark brown to
black and elytra reddish brown with apical, internal and external
margin darker (Fig. 27). Integument of dorsum of head with
undulate microstriae and inconspicuous fine punctures (Figs 74,
75); pronotum with undulate microstriae only on lateral areas
and inconspicuous fine punctures equally distributed; striae of
elytra with inconspicuous fine punctures; metaventrite, on
lateral regions, and abdominal tergites on anterolateral regions
with moderately sized punctures.
Male. Head with front slightly deflected (Fig. 74); V-shaped
frontal sulcus incomplete, curved arms not joined medially;
anterior angles curved, prominent and with a median fovea at
base; eyes slightly prominent from above, and a line of three
moderately sized punctures with long setae on basal half of
dorsal margin, basal the longest. Antennae reaching apex of
abdomen (Fig. 27); scape with slightly enlarged area and long
setae on basal half of exposed dorsal face; scape shorter than
antennomere 2 and 3 combined; antennomere 2 with many long
setae on apical half of dorsal face; antennomeres 4–11 oblong
shape and conspicuously narrowing from antennomere 4 to 11.
Labrum with six setae medially, equal in length; each lateral third,
the antero-internal seta shorter than postero-external seta.
Mandibles with dorsal teeth developed (Fig. 123), not forming
a bifurcate apex, shorter than ventral; internal border
asymmetrical, each with one acute tooth at middle and,
anterior to it, in the right mandible with area that resemble a
tooth, absent in left mandible; external-dorsal margin on basal
half slightly carinate. Maxillary palpus with palpomere 4
subequal in length to 2 and 3 combined. Mentum ~1.5 times
as wide as long. Pronotum wider than long (PW/PL = 1.43)
Revision of Piestus with remarks on related genera
(Fig. 74); anterior angles weakly projected; longitudinal median
sulcus conspicuous; longitudinal sulcus of posterior angles
slightly developed and reaching basal quarter of pronotum;
abrupt constriction at basal quarter; apical three-quarters with
subparallel sides. Prosternum with conspicuous ovate set of fine
punctures in median region near apex. Femur of anterior legs with
short, hard setae on groove of ventral margin. Elytra somewhat
longer than wide (EL/BW = 1.06); stria 6 at basal half, sometimes
inconspicuous and stria 7 at apical half, sometimes
inconspicuous; striae narrower than interstriae. Abdominal
segments 3–6 parallel sides; tergite 9 with short ventral struts;
tergite 10 with two pairs of long setae on apex, apical the longest.
Median lobe of aedeagus with bulbous base in ventral view and
not curved apex in lateral view (Fig. 188); lateral lobes reaching
apex of median lobe in lateral view; internal sclerites as in
Fig. 189.
Female. Similar to male, except for: vertex of head with
anterior angles not prominent as on male (Fig. 75); antennae
exceeding apex of elytra, but not reaching apex of abdomen; scape
as in male, but fewer setae and without enlarged area;
antennomere 2 with few setae on dorsal face; internal border
of right mandible with only one tooth, broad (Fig. 124); ovipositor
as in P. sulcatus; spermathecal duct ~1.5 times length of
spermatheca, without sclerotised basal region (Fig. 260);
spermatheca as in Fig. 260.
Distribution
In the current study the species was examined from Nicaragua
(Matagalpa and Granada), Panama (Colón and Veraguas),
Colombia, Venezuela, Suriname, Ecuador (Sucumbios, Napo),
Peru (Cusco), Bolivia (Cochabamba), Brazil (Rondônia, Pará,
Rio de Janeiro, São Paulo, Paraná and Santa Catarina) and
Paraguay (Caazapá). Herman (2001b) listed the species also
from Guyana and French Guiana.
Biological notes
Piestus pennicornis has been found under bark of decaying logs.
Some specimens were captured by ‘beating palm branches’.
Remarks
Piestus pennicornis is similar to P. rossii and differs by some
characters listed above (see remarks under P. rossii). Specimens
of P. pennicornis have different colour patterns. Colour patterns
were also observed in P. bicornis (see above, remarks in
P. bicornis).
Piestus pygmaeus Laporte, 1835
(Figs 28, 76, 190, 191, 261)
Piestus pygmaeus Laporte, 1835: 130 (original description, type locality:
‘Colombie’); Erichson, 1840: 835 (characters, distribution); Fauvel,
1864: 29 (characters, distribution); Sharp, 1876: 407 (distribution);
Sharp, 1887: 714 (distribution); Blackwelder, 1943: 49 (characters,
distribution, notes); Blackwelder, 1944: 101 (distribution, error:
Laporte, 1834: 130); Herman, 2001b: 1793 (distribution); NavarreteHeredia et al., 2002: 209 (distribution); Newton et al., 2005: 37
(distribution).
Piestus (Piestus) pygmaeus: Bernhauer & Schubert, 1910: 7 (distribution,
error: Laporte, 1834: 130); Scheerpeltz, 1933: 994 (mention);
Invertebrate Systematics
523
Scheerpeltz, 1952: 290 (characters, distribution, error: Laporte,
1834: 130).
Piestus rufipennis Fleutiaux & Sallé, 1889: 382 (cited as synonym of
P. pygmaeus Laporte, 1835, note: manuscript name by Chevrolat). - As
synonym of P. pygmaeus: Blackwelder, 1943: 49, 562; Blackwelder,
1944: 100; Newton et al., 2005: 69. Note: Nomen nudum (Newton
et al. 2005).
Type material
Piestus pygmaeus Laporte, 1835. Lectotype (here designated) deposited
in BMNH, sex undetermined [damaged apex of the antennae and right
anterior leg], with labels: (1) ‘Type’ [circle white label with border red,
printed in black]; (2) ‘Pygmaeus/Cart? Colomb.’ [white label, handwritten];
(3) ‘Sharp Coll./1905.-313.’ [white label, printed in black]; (4) ‘Ex mus.
Castel-?/nau./probably type of/Piestus pygmaeus’ [white labels,
handwritten]; (5) ‘B.C.A. Col. I. 2./Piestus/pygmaeus,/Lap.’ [white label,
printed in black]. Paralectotype deposited in BMNH, sex undetermined
[damaged apex of the left antenna, right antenna and right anterior leg],
and with the same labels of lectotype [lectotype and paralectotype glued
on the same card]. In the original description, Laporte (1835) did not
specify how many specimens were observed. We received two specimens
with type labels.
Additional material
See Appendix 2.
Redescription
BL: 3.1–3.8 mm, BW: 0.8–1.00 mm. Body flattened
dorsoventrally; head, pronotum and abdomen light brown to
black and elytra light brown to reddish, always lighter than
body (Fig. 28). Dorsal integument of head with undulate
microstriae and inconspicuous fine punctures (Fig. 76);
pronotum with undulate microstriae, sometimes absent near
longitudinal sulcus, and conspicuous fine punctures equally
distribute, some moderately sized punctures on median
longitudinal sulcus; striae and interstriae of elytra with
inconspicuous fine punctures; metaventrite, on lateral regions,
and abdominal tergites, on anterolateral regions, with moderately
sized punctures.
Male. Head with front slightly deflected (Fig. 76); V-shaped
frontal sulcus complete; anterior angles curved and weakly
prominent; eyes with line of three moderately sized punctures
with long setae on basal half of dorsal margin, middle very
short and basal the longest. Antennae reaching apex of
abdomen (Fig. 28); scape with strongly enlarged area and long
setae on basal half of exposed dorsal face; scape shorter than
antennomere 2 and 3 combined; antennomeres 4–11 oblong
shape and slightly narrowing from antennomere 4 to 11.
Labrum with six setae medially, equal in length; each lateral
third, antero-internal seta shorter than postero-external seta.
Mandibles with dorsal teeth weakly developed; internal border
slightly asymmetrical, each with one small and acute tooth at
middle and, anterior to it, in right mandible with a very weak
area that resembles a tooth, absent in left mandible. Maxillary
palpus with palpomere 4 longer than 2 and 3 combined. Mentum
~1.5 times as wide as long. Pronotum wider than long (PW/
PL = 1.36) (Fig. 76); anterior weakly projected; longitudinal
media sulcus conspicuous; longitudinal sulcus of posterior
angles developed and reaching basal third of pronotum; abrupt
constriction at basal quarter; apical three-quarters with parallel
524
Invertebrate Systematics
sides. Femur of anterior legs with short, hard setae on groove of
ventral margin. Elytra somewhat longer than wide (EL/
BW = 1.08); stria 6 at basal half, sometimes inconspicuous,
and stria 7 absent; striae narrower than interstriae. Abdominal
segments 3–6 parallel sides; tergite 9 with short ventral struts;
tergite 10 with two pairs of long setae on apex, apical the longest.
Median lobe of the aedeagus with bulbous base in ventral view
and curved apex in lateral view (Fig. 190); lateral lobes exceeding
apex of median lobe by one-quarter of its length in lateral view,
and apical half much wider than basal; internal sclerites as in
Fig. 191.
Female. Similar to male, except for: antennae exceed a little
the apex of elytra; scape with fewer setae; ovipositor as in
P. sulcatus; spermathecal duct ~1.5 times length of
spermatheca, with sclerotised basal region (Fig. 261);
spermatheca with basal half suddenly narrower than apical
(Fig. 261).
Distribution
In the current study the species was examined from Mexico (San
Luís Potosí, Jalisco, Veracruz, México, Oaxaca, Chiapas and
Nayarit), Belize (Orange Walk, Belize and Toledo), Guatemala
(Zacapa and Escuintla), Honduras (Santa Bárbara, Colón and
Francisco Morazán), Nicaragua (Granada and Río San Juan),
Costa Rica (Guanacaste, Heredia, Cartago, Limón and
Puntarenas), Panama (Bocas del Toro, Chiriquí, Veraguas,
Colón and Panamá), Guadeloupe, Saint Vincent and the
Grenadines (Saint Vincent), Grenada, Trinidad and Tobago
(Trinidad), Colombia (Magdalena, César, Boyacá, Meta and
Putumayo), Venezuela (Bolívar), Guyana (Demerara-Mahaica
and Potaro-Siparuni), French Guiana (Saint-Laurent-du-Maroni
and Cayenne), Ecuador (Sucumbios, Pichincha, Chimborazo and
Pastaza), Peru (Loreto, Cusco and Junin), Bolivia (Beni,
Cochabamba and Santa Cruz), Brazil (Pará, Ceará,
Pernambuco, Bahia, Mato Grosso, Goiás, Espirito Santo,
Minas Gerais, Rio de Janeiro, São Paulo and Santa Catarina),
Paraguay (Asunción, Paraguarí, Guairá, Caazapá and Itapúa) and
Argentina (Salta, Tucumán and Misiones). Navarrete-Heredia
et al. (2002) list P. pygmaeus from some countries above
(note: ‘Antillas’) as well as Uruguay and Galápagos Islands
(Ecuador), and Newton and Peck (2006) also listed it from the
Galápagos Islands (Santa Cruz). In Mexico, Navarrete-Heredia
et al. (2002) cite this species also from Sinaloa and Tabasco.
Biological notes
Piestus pygmaeus has been found under bark, in leaf litter near
decaying logs, recently dead or not (‘in dry inner bark of
Lecythis’), and on fungi (‘fungus covered log’). Some
specimens were collected in flight intercept traps or UV light
traps. This species occurs together with P. fronticornis,
P. bicornis and P. minutus.
Remarks
Piestus pygmaeus is similar to P. extimus and P. buquetii but
differs mainly by slightly deflected front (Fig. 76), prominent
eyes when seen from above, undulate microstriae and
conspicuous fine punctures on pronotum, parallel sides of
E. Caron et al.
pronotum, and very broad apex of lateral lobes of aedeagus
(Fig. 190).
Piestus extimus and P. buquetii have front suddenly deflected
(Figs 77, 78), eyes slightly prominent from above, and subparallel
lateral sides of pronotum. Piestus extimus has undulate
microstriae and inconspicuous fine punctures equally
distributed on pronotum (Fig. 77), and narrower apex of lateral
lobes (Fig. 192), while Piestus buquetii has undulate microstriae
only on lateral areas on pronotum and conspicuous fine punctures
(Fig. 78), and slightly broad apex of lateral lobes (Fig. 194).
Piestus extimus Sharp, 1887
(Figs 29, 77, 192, 193, 262)
Piestus extimus Sharp, 1887: 713 (original description, type locality:
‘Mexico, Pinos Altos in Chihuahua’); Blackwelder, 1944: 100
(distribution); Newton et al. 2000: 376 (distribution, biological
notes); Herman, 2001b: 1791 (distribution); Navarrete-Heredia
et al., 2002: 208 (distribution).
Piestus (Piestus) extimus: Bernhauer & Schubert, 1910: 7 (distribution);
Scheerpeltz, 1952: 290 (characters, distribution).
Type material
Holotype deposited in BMNH, female, with labels: (1) ‘Piestus extimus/
Type D.S./Pinos altos. Chihua/hua. Buch-Hepburn’ [white label, together
with the specimen, handwritten]; (2) ‘Holo-/type’ [circle white label with
border red, printed in black]; (3) ‘Pinos Altos,/Chihuahua,/Mexico./BuchanHepburn’ [white label, printed in black]; (4) ‘Sharp Coll./1905-313.’ [white
label, printed in black]; (5) ‘Holotype/Piestus/extimus/Sharp, 1887/det.
R.G. Booth 2008’ [white label, the four first lines handwritten and the last
one printed in black]. Note: In the original description Sharp (1887) specified
only one specimen observed.
Additional material
See Appendix 2.
Redescription
BL: 3.7–5.2 mm, BW: 1.1–1.3 mm. Body flattened
dorsoventrally; head, pronotum and abdomen light brown to
black and elytra light brown to reddish, always lighter than
body (Fig. 29). Dorsal integument of head and pronotum
generally with undulate microstriae and inconspicuous fine
punctures, sometimes with moderately sized punctures on
median longitudinal sulcus (Fig. 77); striae and interstriae of
elytra with inconspicuous fine punctures; metaventrite, on lateral
regions, and abdominal tergites, on anterolateral regions, with
moderately sized punctures.
Male. Head with front suddenly deflected (Fig. 77);
V-shaped frontal sulcus complete; anterior angles curved and
weakly prominent; eyes with line of three moderately sized
punctures with long setae on basal half of dorsal margin of
eyes, basal the longest. Antennae almost reaching apex of
abdomen (Fig. 29); scape with strongly enlarged area and long
setae on the basal half of exposed dorsal face; scape shorter than
antennomere 2 and 3 combined; antennomeres 4–11 oblong
shape and slightly narrowing from antennomere 4 to 11.
Labrum with six setae medially, equal in length; on each
lateral third, antero-internal seta shorter than postero-external
seta. Mandibles with dorsal teeth weakly developed; internal
Revision of Piestus with remarks on related genera
border slightly asymmetrical, each with one small and acute
tooth at middle and, anterior to it, in right mandible with a
very weak area that resembles a tooth, absent in left mandible.
Maxillary palpus with palpomere 4 longer than 2 and 3 combined.
Mentum ~1.5 times as wide as long. Pronotum wider than
long (PW/PL = 1.41) (Fig. 77); anterior angles weakly
projected; longitudinal median sulcus conspicuous;
longitudinal sulcus of posterior angles developed and reaching
basal third of pronotum; abrupt constriction at basal quarter;
apical three-quarters with subparallel sides. Femur of anterior
legs with short, hard setae on groove of ventral margin. Elytra as
long as wide (EL/BW = 0.98); stria 6 at basal half, sometimes
inconspicuous, and stria 7 absent; striae narrower than interstriae.
Abdominal segments 3–6 with parallel sides; tergite 9 with
short ventral struts; tergite 10 with two pairs of long setae on
apex, apical the longest. Median lobe of aedeagus with bulbous
base in ventral view and curved apex in lateral view (Fig. 192);
lateral lobes exceed a little the apex of median lobe in lateral
view; internal sclerites as in Fig. 193.
Female. Similar to male, except for: antennae exceed a little
apex of elytra; scape with fewer setae; ovipositor as in P. sulcatus;
spermathecal duct ~1.5 times length of spermatheca, with
sclerotised basal region (Fig. 262); spermatheca with basal half
suddenly narrower than apical (Fig. 262).
Distribution
In the current study the species was examined from the United
States of America (Arizona) and Mexico (Durango, Nayarit,
Jalisco, México, Oaxaca and Guanajuato). In Mexico,
Navarrete-Heredia et al. (2002) list P. extimus also from
Chihuahua and Sonora.
Biological notes
Piestus extimus has been found on decaying cacti and sotol
(Dasylirion spp.) (Newton et al. 2000), under bark of decaying
logs and collected in flight intercept traps.
Remarks
Piestus extimus is very similar to P. buquetii, from which it is
easily separated by undulate microstriae and inconspicuous fine
punctures on pronotum, and narrow apex of lateral lobes of
aedeagus (in P. buquetii, undulate microstriae only on lateral
areas, conspicuous fine punctures equally distributed and slightly
broad apex of lateral lobes).
Piestus extimus is also similar to P. pygmaeus, from which it
differs by characters listed above (see remarks under
P. pygmaeus).
Piestus buquetii Fauvel, 1864
(Figs 30, 78, 194, 195, 263)
Piestus buquetii Fauvel, 1864: 28 (original description, type locality:
‘Cayenne’); Herman, 2001b: 1789 (distribution); Navarrete-Heredia
et al., 2002: 208 (distribution); Newton et al., 2005: 37 (distribution).
Piestus (Piestus) buqueti: Bernhauer & Schubert, 1910: 7 (error:
misspelling and Fauvel 1865: 32, distribution); Blackwelder, 1944:
100 (error: misspelling and Fauvel 1865: 32, distribution); Scheerpeltz,
Invertebrate Systematics
525
1952: 291 (error: misspelling and Fauvel 1865: 32, characters,
distribution).
Type material
Holotype deposited in IRSNB, male, with labels: (1) ‘Cayenne’ [white label,
handwritten]; (2) ‘Buqueti/Fvl.’ [white label, handwritten]; (3) ‘R.I.Sc.N.
B. 17.479/Piestus/Coll. et det. A. Fauvel’ [white label, first and third lines
printed in black and second line handwritten]; (4) ‘Type’ [red label, printed in
black]. Note: In the original description Fauvel (1864) specified only one
specimen observed.
Additional material
See Appendix 2.
Redescription
BL: 5.0–5.5 mm, BW: 1.3–1.4 mm. Body flattened
dorsoventrally; head, pronotum and abdomen light brown to
black and elytra light brown to reddish, always lighter than
body (Fig. 30). Dorsal integument of head with undulate
microstriae and inconspicuous fines punctures (Fig. 78);
pronotum with undulate microstriae only on lateral areas and
conspicuous fine punctures equally distributed, some
moderately sized punctures on longitudinal sulcus; striae and
interstriae of elytra with inconspicuous fine punctures.
Metaventrite, on lateral regions, and abdominal tergites, on
anterolateral regions, with moderately sized punctures.
Male. Head with front suddenly deflected (Fig. 78); Vshaped frontal sulcus complete; anterior angles curved and
weakly prominent; eyes with line of three moderately sized
punctures with long setae on basal half of dorsal margin, basal
the longest. Antennae exceeding apex of abdomen (Fig. 30);
scape with strongly enlarged area and long setae on basal half of
exposed dorsal face; scape shorter than antennomere 2 and 3
combined; antennomeres 4–11 oblong shape and slightly
narrowing from antennomere 4 to 11. Labrum with six setae
medially, equal in length; on each lateral third, antero-internal
seta a little shorter than postero-external seta. Mandibles with
dorsal teeth weakly developed; internal border slightly
asymmetrical, each with one small and acute tooth at middle
and, anterior to it in right mandible with a very weak area that
resembles a tooth, absent in left mandible. Pronotum wider than
long (PW/PL = 1.34) (Fig. 78); anterior angles weakly projected;
longitudinal median sulcus conspicuous; longitudinal sulcus of
posterior angles developed and reach basal third of pronotum;
abrupt constriction at basal quarter; apical three-quarters
with subparallel sides. Femur of anterior legs with short, hard
setae on groove of ventral margin. Elytra as long as wide (EL/
BW = 1.03); stria 6 at basal half, sometimes inconspicuous, and
stria 7 absent; striae narrower than interstriae. Abdominal
segments 3–6 with parallel sides; tergite 9 with short ventral
struts; tergite 10 with two pairs of long setae on apex, apical
the longest. Median lobe of the aedeagus with bulbous base
in ventral view and curved apex in lateral view (Fig. 194);
lateral lobes exceed a little apex of median lobe in lateral
view and weakly wide on apex; internal sclerites as in
Fig. 195.
Female. Similar to male, except for: antennae exceed a little
apex of elytra; scape with fewer setae; ovipositor as in P. sulcatus;
spermathecal duct ~2 times the length of spermatheca, with
526
Invertebrate Systematics
sclerotised basal region (Fig. 263); spermatheca with basal half
suddenly narrower than apical (Fig. 263).
Distribution
In the current study the species was examined from French
Guiana (Cayenne), Peru (Cusco) and Bolivia (La Paz and
Beni). Navarrete-Heredia et al. (2002) list P. buquetii also
from Mexico (Guanajuato), Guyana, Suriname, Brazil,
Paraguay and Argentina. However, the distribution of
P. buquetii in Mexico is doubtful, as this species is very
similar to P. extimus, which occurs in Mexico.
Biological notes
Piestus buquetii has been found under bark or on fungi on
decaying logs (‘fungus covered log’).
Remarks
Piestus buquetii is very similar to P. extimus, from which it is
easily separated by characters of the pronotum and lateral lobes
of aedeagus (see remarks under P. extimus). Piestus buquetii is
also similar to P. pygmaeus, from which it differs by characters
listed above (see remarks under P. pygmaeus).
Piestus filicornis Fauvel, 1902
(Figs 31, 79, 125, 196, 197, 264)
Piestus filicornis Fauvel, 1902: 23 (original description, type locality:
‘Bolivie: Yuracaris’); Blackwelder, 1944: 100 (distribution, error:
Fauvel, 1902: 22); Herman, 2001b: 1791 (distribution).
Piestus (Piestus) filicornis: Bernhauer & Schubert, 1910: 7 (distribution,
error: Fauvel, 1902: 22); Scheerpeltz, 1952: 287 (characters,
distribution, error: Fauvel, 1902: 22).
Type material
Holotype deposited in IRSNB, male [specimen damaged, left antenna
without antennomeres 8–11, right antenna without antennomeres 9–11],
with labels: (1) ‘Yuracaris/Bolivie’ [white label, handwritten]; (2)
‘filicornis/Fvl.’ [white label, handwritten]; (3) ‘R.I.Sc.N.B. 17.479/Piestus/
Coll. et det. A. Fauvel’ [white label, first and third lines printed in black
and second line handwritten]; (4) ‘Type’ [red label, printed in black]. Note:
In the original description Fauvel (1902) specified only one specimen
observed.
Additional material
See Appendix 2.
Redescription
BL: 7.9–8.3 mm, BW: 2.0–2.2 mm. Body flattened
dorsoventrally; entirely brown to reddish dark brown (Fig. 31).
Dorsal integument of head and pronotum with undulate
microstriae and inconspicuous fine punctures (Fig. 79); striae
and interstriae of elytra with inconspicuous fine punctures;
metaventrite, on lateral regions, and abdominal tergites, on
anterolateral regions, with moderately sized punctures.
Male. Head with front suddenly deflected (Fig. 79); Vshaped frontal sulcus complete; anterior angles curved and
prominent, and transverse carina on base, slightly emarginate
at middle; eyes with line of three moderately sized punctures
with long setae on basal half of dorsal margin, basal the longest.
E. Caron et al.
Antennae reaching apex of abdomen (Fig. 31); scape with
strongly enlarged area and long setae on basal half of exposed
dorsal face; scape shorter than antennomere 2 and 3 combined;
antennomeres 4–11 oblong shape and conspicuously narrowing
from antennomere 4 to 11. Labrum with six setae medially,
equal in length; on each lateral third, the antero-internal seta
shorter than postero-external seta. Mandibles with dorsal teeth
developed, not forming a bifurcate apex (Fig. 125), shorter than
ventral; internal border asymmetrical, each with one acute tooth at
middle (left somewhat deflected) and, anterior to it, in right
mandible with a area that resemble a tooth, absent in left
mandible; external dorsal margin on basal half carinate.
Maxillary palpus with palpomere 4 longer than 2 and 3
combined. Mentum ~1.5 times as wide as long. Pronotum
wider than long (PW/PL = 1.48) (Fig. 79); anterior angles
weakly projected; longitudinal median sulcus conspicuous;
longitudinal sulcus of posterior angles slightly developed and
reaching basal quarter of pronotum; abrupt constriction at basal
quarter; apical three-quarters with subparallel sides. Femur of
anterior legs with short, hard setae on groove of ventral margin.
Elytra as long as wide (EL/BW = 1.00); stria 6 at the basal half,
sometimes inconspicuous, and stria 7 absent; striae narrower
than interstriae. Abdominal segments 3–6 parallel sides; tergite 9
with short ventral struts; tergite 10 with two pairs of long setae on
apex, apical the longest. Median lobe of the aedeagus with
bulbous base in ventral view and curved apex in lateral view
(Fig. 196); lateral lobes exceeding apex of median lobe by onequarter of its length in lateral view, external margin, near the apex,
conspicuously curved; internal sclerites as in Fig. 197.
Female. Similar to male, except for: antennae exceed a little
the apex of elytra; scape with fewer long setae; ovipositor as in
P. sulcatus; spermathecal duct ~1.5 times length of spermatheca,
with sclerotised basal region (Fig. 264); spermathecal capsule
with apex somewhat globose and with projection on the base
(Fig. 264).
Distribution
Bolivia (Cochabamba).
Biological notes
All material of P. filicornis observed in this study (except the
holotype) were collected in flight intercept traps.
Remarks
Piestus filicornis is similar to P. penicillatus and P. minutus but
differs by transverse carina slightly emarginate at the middle on
vertex, and curved apex of median lobe of aedeagus (Fig. 79)
(transverse carina interrupted at the middle and uncurved apex of
median lobe in P. penicillatus; Fig. 83; transverse carina
continuous at the middle and uncurved apex of median lobe in
P. minutus; Fig. 82).
Piestus filicornis, as well as P. penicillatus and P. minutus, are
also similar to P. abdominalis and P. niger, with which they share
the transverse carina on base of vertex. However, these species
(Piestus filicornis, P. penicillatus and P. minutus) may be easily
separated from them by integument of pronotum with undulate
microstriae equally distributed on pronotum (Figs 79, 82, 83)
(P. abdominalis and P. niger have only undulate microstriae on
Revision of Piestus with remarks on related genera
lateral areas; Figs 80, 81). There is also a difference in
morphology of the spermatheca, in which the spermatheca
has a projection on the base of capsule (P. filicornis,
P. penicillatus and P. minutus; Figs 264, 267, 268) or lacks it
(P. abdominalis and P. niger; Figs 265, 266).
Piestus abdominalis, sp. nov.
(Figs 32, 80, 198, 199, 265)
Type material
Holotype deposited in AMNH, male, labels: (1) ‘Mexico: Chiapas/Union
Juarez, n.v. slope/Volcan Tacapa, Rio de/Finca Muxbal (Guat.) ex./pile of
nouldy wood/1430m 21 Dec 1976/H. Frania D. Proctor 6’ [white label,
printed in black]. 12 Paratypes deposited in following collections: 7 in
AMNH, 1 male, 1 female, labels: 1) ‘Mexico, Chiapas/Union Juarez, n.e.
slope/Volcan Tacana, on ground/agricultural land, ele./1720 m, 14-26 Dec
1975/115-75’ [white label, printed in black]; (2) ‘H. Frania/Collector’ [white
label, printed in black]; 1 female, labels: (1) ‘Mexico, Chiapas/Union Juarez
Barranca/Providencia, n.e. slope/Volcan Tacana, shaded/stream, leaf litter/
edge, ele. 1560 m,/16-25 Dec 1975, 119-75’ [white label, printed in black];
‘H. Frania/Collector’ [white label, printed in black]; 1 female, labels: (1)
‘Mexico, Chiapas/Union Juarez, n.e. slope/Volcan Tacana, (Guat.)/Rio de
Finca Muxba, ex./freshly cut stump, much/sap, peeling bark, ele./1430 m, 26
Dec 1975/219-75’ [white label, printed in black]; (2) ‘H. Frania/Collector’
[white label, printed in balck]; 1 female, labels: (1) ‘Mexico, Chiapas/Union
Juarez, n.e. slope/Volcan Tacana, (Guat.)/Rio de Finca Muxba, leaf/litter river
edge, ele./1430 m, 26 Dec 1975/218-75’ [white label, printed in black];
‘H. Frania/Collector’ [white label, printed in black]; 1 female, label: (1)
‘Mexico: Chiapas/Union Juarez n.v. slope/Volcan Tacana ex. lear/litter
weedy areas under/coffee 1000m 3 Jan 1977/H. Frania 28’ [white label,
printed in black]; 1 in SEMC, 1 male, labels: (1) ‘Mexico, Chiapas/Union
Juarez, n.e. slope/Volcan Tacana, on ground/agricultural land, ele./1720 m,
14-26 Dec 1975/115-75’ [white label, printed in black]; (2) ‘H. Frania/
Collector’ [white label, printed in black]; 1 in DZUP, male, labels: (1)
‘Mexico, Chiapas/Union Juarez, n.e. slope/Volcan Tacana, (Guat.)/Rio de
Finca Muxba, ex./freshly cut stump, much/sap, peeling bark, ele./1430 m, 26
Dec 1975/219-75’ [white label, printed in black]; (2) ‘H. Frania/Collector’
[white label, printed in balck]; 4 in FMNH, 1 sex undetermined, labels: (1)
‘Guatemala:/N.W. Slope V. de/Fuego. V:2:1948/5700’.R.Mitchell’ [white
label, printed in black]; (2) ‘on/fungi’ [white label, printed in black]; (3)
‘Piestus/(s. st.) spp./det. A.F. Newton 1987’ [white label, two first lines
handwritten, third printed in black]; 1 male, labels: (1) ‘Mun. Yepocapa,/
Chinal, Guat./V-2:48’ [white label, printed in black, except ‘V-2’
manuscript]; (2) ‘La Jolla Grande/(Finca Monserrat)/NW slope Volcan/
Fuego El.5700ft.’ [white label, printed in black]; (3) ‘CNHM Guatemala/
Zool. Exped. (1948)/R.L.Wenzel, leg./Lot. No.’ [white label, printed in
black]; (4) ‘ex rotting/banana/stalks’ [white label, printed in black]; 1 sex
undetermined, labels: the same two first labels; (3) ‘CNHM Guatemala/Zool.
Exped. (1948)/R.L.Wenzel, leg./Lot. No. 75’ [white label, printed in black,
except ‘75’ manuscript]; 1 sex undetermined, labels: (1) ‘Mun. Yepocapa,/
Chinal, Guat./V-3:48’ [white label, printed in black]; the same second label;
(3) ‘CNHM Guatemala/Zool. Exped. (1948)/R.L. Wenzel leg./Lot. No. 82’
[white label, printed in black, except ‘82’]; 4) ‘under/rotting/banana/stalks’
[white label, printed in black].
Description
BL: 5.8–6.7 mm, BW: 1.6–1.8 mm. Body flattened
dorsoventrally; head, pronotum and elytra dark brown to black
and abdomen light brown to reddish, always lighter than body.
Sometimes elytra lighter than head and pronotum, but darker than
abdomen (Fig. 32). Dorsal integument of head with undulate
Invertebrate Systematics
527
microstriae and inconspicuous fine punctures (Fig. 80); pronotum
with undulate microstriae only on lateral regions and conspicuous
fine punctures equally distributed and some moderate on
longitudinal sulcus; striae and interstriae of elytra with
inconspicuous fine punctures; metaventrite, on lateral regions,
and abdominal tergites, on anterolateral regions, with moderately
sized punctures.
Male. Head with front suddenly deflected (Fig. 80); Vshaped frontal sulcus complete; anterior angles curved and
prominent, and transverse carina on base, continuous at
middle; eyes with line of three moderately sized punctures
with long setae on basal half of dorsal margin, basal the
longest. Antennae reaching apex of abdomen (Fig. 32); scape
with strongly enlarged area and long setae on basal half of
exposed dorsal face; scape shorter than antennomere 2 and 3
combined; antennomeres 4–11 oblong shape and conspicuously
narrowing from antennomere 4 to 11. Labrum with six setae
medially, equal in length; each lateral third, antero-internal seta
shorter than postero-external seta. Mandibles with dorsal teeth
developed, not forming a bifurcate apex, shorter than ventral;
internal border asymmetrical, each with one acute tooth at
middle and, anterior to it, in right mandible with a area that
resemble a tooth, absent in left mandible; external dorsal
margin on basal half slightly carinate. Maxillary palpus with
palpomere 4 longer than 2 and 3 combined. Mentum ~1.5 times as
wide as long. Pronotum wider than long (PW/PL = 1.41)
(Fig. 80); anterior angles weakly projected; longitudinal
median sulcus conspicuous; longitudinal sulcus of posterior
angles reach basal third of pronotum; abrupt constriction at
basal quarter; apical three-quarters with subparallel sides.
Femur of anterior legs with short, hard setae on groove of
ventral margin. Elytra as long as wide (EL/BW = 0.97); stria 6
at basal half, sometimes inconspicuous, and stria 7 absent;
striae narrower than interstriae. Abdominal segments 3–6
parallel sides; tergite 9 with short ventral struts; tergite 10 with
two pairs of long setae on apex, apical the longest. Median lobe
of the aedeagus with bulbous base in ventral view and curved
apex in lateral view (Fig. 198); lateral lobes exceed a little apex of
median lobe in lateral view, and apex somewhat truncate on
ventral margin; internal sclerites as in Fig. 199.
Female. Similar to male, except for: antennae exceed a little
apex of elytra; scape with fewer long setae; ovipositor as in
P. sulcatus; spermathecal duct ~1.5 times length of spermatheca,
with sclerotised basal region (Fig. 265); spermatheca with basal
half suddenly narrower than apical (Fig. 265).
Distribution
Mexico (Chiapas) and Guatemala (Chimaltenango).
Biological notes
Piestus abdominalis has been found in litter, on fungi or in
association with rotting logs. One specimen was found in ‘leaf
litter river edge’.
Remarks
Piestus abdominalis is very similar to P. niger differing by some
moderately sized punctures on longitudinal sulcus of pronotum
528
Invertebrate Systematics
(Fig. 80), curved apex of median lobe of aedeagus and slightly
truncate apex of lateral lobes on ventral margin (Fig. 198).
Piestus niger has only inconspicuous fine punctures on
pronotum (Fig. 81), uncurved apex of median lobe and nontruncate apex of lateral lobes (Fig. 200).
Piestus abdominalis, as well as P. niger, is also similar to
P. filicornis, P. penicillatus and P. minutus, from which they
differ by characters discussed above (see remarks under
P. filicornis).
Etymology
The species name refers to the colour of the abdomen, which is
always lighter than the rest of the body.
Piestus niger Fauvel, 1864
(Figs 33, 81, 200, 201, 266)
Piestus niger Fauvel, 1864: 24 (original description, type locality:
‘Mexico’); Sharp, 1887: 712 (distribution); Blackwelder, 1944: 100
(distribution, error: Fauvel, 1865: 28); Herman, 2001b: 1792
(distribution); Navarrete-Heredia et al., 2002: 209 (distribution).
Piestus (Piestus) niger: Bernhauer & Schubert, 1910: 7 (distribution,
error: Fauvel, 1865: 28); Scheerpeltz, 1952: 289 (characters,
distribution, error: Fauvel, 1865: 28).
Type material
Piestus niger Fauvel, 1864. Lectotype (here designated) deposited in
IRSNB, male, with labels: (1) ‘Cordova/Mexique’ [white label, handwritten];
(2) ‘niger/Fvl.’ [white label, handwritten]; (3) ‘R.I.Sc.N.B. 17.479/Piestus/
Coll. et det. A. Fauvel’ [white label, first and third lines printed in black and
second line handwritten]; (4) ‘Ex-Typis’ [white label, printed in red]. Two
paralectotypes deposited in IRSNB, 1 female with the same labels of
lectotype, and 1 male, with the same labels of lectotype, except the second
label, absent. Note: In the original description Fauvel (1864) did not specify
how many specimens he observed. We consider all material from IRSNB with
type label ‘Ex-Typis’ as type material.
Additional material
E. Caron et al.
narrowing from antennomere 4 to 11. Labrum with six setae
medially, equal in length; each lateral third, antero-internal seta
shorter than postero-external seta. Mandibles with dorsal teeth
developed, not forming a bifurcate apex, shorter than ventral;
internal border asymmetrical, each with acute tooth at middle
and, anterior to it, in right mandible with a area that resembles a
tooth, absent in left mandible; external dorsal margin on basal
half slightly carinate. Maxillary palpus with palpomere 4 longer
than 2 and 3 combined. Mentum ~1.5 times as wide as long.
Pronotum wider than long (PW/PL = 1.43) (Fig. 81); anterior
angles weakly projected; longitudinal median sulcus
conspicuous; longitudinal sulcus of posterior angles reach
basal third of pronotum; abrupt constriction at basal quarter;
apical three-quarters with subparallel sides. Femur of anterior legs
with short, hard setae on groove of ventral margin. Elytra as long
as wide (EL/BW = 0.99); stria 6 at basal half, sometimes
inconspicuous, and stria 7 absent; striae narrower than
interstriae. Abdominal segments 3–6 parallel sides; tergite 9
with short ventral struts; tergite 10 with two pairs of long setae
on apex, apical the longest. Median lobe of aedeagus with
bulbous base in ventral view and not curved apex in lateral
view (Fig. 200); lateral lobes exceed a little apex of median
lobe in lateral view; internal sclerites as in Fig. 201.
Female. Similar to male, except for: antennae exceed a little
the apex of elytra; scape with fewer long setae; ovipositor as in
P. sulcatus; spermathecal duct about same length of spermatheca,
with sclerotised basal region (Fig. 266); spermatheca with basal
half suddenly narrower than apical half (Fig. 266).
Distribution
In the current study, the species was examined from
Mexico (Veracruz), Guatemala (Guatemala), Costa Rica
(Guanacaste) and Panama. Navarrete-Heredia et al. (2002) list
this species also from Honduras and Nicaragua. In Mexico,
Navarrete-Heredia et al. (2002) cite this species in Oaxaca,
Puebla and Veracruz.
See Appendix 2.
Biological notes
Redescription
The only information noted for Piestus niger is occurring under
bark of decaying logs.
BL: 5.0–6.0 mm, BW: 1.4–1.6 mm. Body flattened
dorsoventrally; entirely dark brown to black (Fig. 33). Dorsal
integument of head with undulate microstriae and inconspicuous
fine punctures (Fig. 81); pronotum with undulate microstriae only
on lateral regions and inconspicuous fine punctures equally
distributed; striae and interstriae of elytra with inconspicuous
fine punctures; metaventrite, on lateral regions, and abdominal
tergites, on anterolateral regions, with moderately sized
punctures.
Male. Head with front suddenly deflected (Fig. 81); Vshaped frontal sulcus complete; anterior angles curved and
prominent, and transverse carina on base, continuous at
middle; eyes with line of three moderately sized punctures
with long setae on basal half of dorsal margin, basal the
longest. Antennae reaching apex of abdomen (Fig. 33); scape
with strongly enlarged area and long setae on basal half of
exposed dorsal face; scape shorter than antennomere 2 and 3
combined; antennomeres 4–11 oblong shape and conspicuously
Remarks
Piestus niger is very similar to P. abdominalis but differs by
characters cited under remarks of P. abdominalis. It is also similar
to P. filicornis, P. penicillatus and P. minutus (see discussion
under remarks of P. filicornis).
Piestus minutus Erichson, 1840
(Figs 34, 82, 202, 203, 267)
Piestus minutus Erichson, 1840: 834 (original description, type locality:
‘Columbia’); Fauvel, 1864: 27 (characters, distribution); Sharp, 1876:
407 (note, distribution); Sharp, 1887: 713 (note, distribution); Fauvel,
1901: 71 (distribution); Blackwelder, 1944: 100 (distribution);
Herman, 2001b: 1792 (distribution); Navarrete-Heredia et al., 2002:
208 (distribution); Newton et al., 2005: 37 (distribution).
Piestus (Piestus) minutus: Bernhauer & Schubert, 1910: 7 (distribution);
Scheerpeltz, 1952: 291 (characters, distribution).
Revision of Piestus with remarks on related genera
Piestus sulcatus: Laporte, 1835: 129. - As junior synonym of P. minutus:
Erichson, 1840: 834; Fauvel, 1864: 27; Fauvel, 1901: 71; Bernhauer &
Schubert, 1910: 7 (error: Laporte, 1834: 129); Blackwelder, 1944: 100
(error: Laporte, 1834: 129); Navarrete-Heredia et al., 2002: 209;
Newton et al., 2005: 69. Note: Misidentification, Laporte did not
describe the species as new, he attributed it to Gravenhorst, 1806
(Herman 2001b).
Piestus nigrator Fauvel, 1902: 24 (original description, type locality:
‘Amazones: Pebas’); Blackwelder, 1944: 100 (distribution); Herman,
2001b: 1792 (distribution). New synonym.
Piestus (Piestus) nigrator: Bernhauer & Schubert, 1910: 7 (distribution);
Scheerpeltz, 1952: 291 (characters, distribution).
Type material
Piestus minutus Erichson, 1840. Lectotype (here designated) deposited in
ZMHB, male, with labels: (1) ‘6810’ [white label/printed in black]; (2)
‘minutus/Er./Columb. Mor.’ [green label, handwritten]; 3) ‘SYNTYPUS/
Piestus/minutus Erichson, 1840/labelled by MNHUB 2006’ [red label,
printed in black]. Four paralectotypes deposited in ZMHB, sex
undetermined, with labels: (1) ‘Hist.-Coll. (Coleoptera)/Nr. 6810/Piestus
minutus/Erichson, 1840/Columb. Moritz/Zool. Mus. Berlin’ [green label/
printed in black]; (2) ‘SYNTYPUS/Piestus/minutus Erichson, 1840/labelled
by MNHUB 2006’ [red label, printed in black]. Note: In the original
description Erichson (1840) did not specify how many specimens he
observed. We consider all material from ZMHB with type label
‘SYNTYPUS/Piestus/minutus. . .’ as type material.
Piestus nigrator Fauvel, 1902. Lectotype (here designated) deposited in
IRSNB, 1 female, with labels: (1) ‘Pebas’ [white label, handwritten];
(2) ‘nigrator/FvL.’ [white label, handwritten]; (3) ‘R.I.Sc.N.B. 17.479/
Piestus/Coll. et det. A. Fauvel’ [white label, first and third lines printed in
black and second line handwritten]; (4) ‘Ex-Typis’ [white label, printed in
red]. Three paralectotypes deposited in IRSNB, 2 males and 1 female, with
labels: (1) ‘Pebas’ [white label, handwritten]; (2) ‘Coll. et det. A. Fauvel/
Piestus/nigrator Fauv./R.I.Sc.N.B. 17.479’ [white label, first and fourth lines
printed in black, second and third lines handwritten]; (3) ‘Ex-Typis’ [white
label, printed in red]. Note: In the original description Fauvel (1902) did not
specify how many specimens he observed. We consider all material from
IRSNB with type label ‘Ex-Typis’ as type material. We designated a female as
lectotype because it is the only specimen with presumably Fauvel’s
handwritten label.
Additional material
See Appendix 2.
Redescription
BL: 4.2–5.0 mm, BW: 1.2–1.3 mm. Body flattened
dorsoventrally; light brown to reddish dark brown, sometimes
with elytra reddish and apical margin darker (Fig. 34). Dorsal
integument of head and pronotum with undulate microstriae
and inconspicuous fine punctures (Fig. 82); striae and
interstriae of elytra with inconspicuous fine punctures;
metaventrite, on lateral regions, and abdominal tergites, on
anterolateral regions, with moderately sized punctures.
Male. Head with front suddenly deflected (Fig. 82);
V-shaped frontal sulcus complete, sometimes inconspicuous at
middle; anterior angles curved and prominent, and a transverse
carina on base; eyes with line of three moderately sized
punctures with long setae on basal half of dorsal margin of
eyes, basal the longest. Antennae almost reaching apex of
abdomen (Fig. 34); scape with strongly enlarged area and long
setae on basal half of exposed dorsal face; scape shorter than
Invertebrate Systematics
529
antennomere 2 and 3 combined; antennomeres 4–11 oblong
shape and conspicuously narrowing from antennomere 4 to 11.
Labrum with six setae medially, equal in length; each lateral
third, antero-internal seta shorter than postero-external seta.
Mandibles with dorsal teeth developed, not forming a bifurcate
apex, shorter than ventral; internal border slightly
asymmetrical, each with one acute tooth at middle and,
anterior to it, in right mandible with area that resemble a tooth,
absent in left mandible. Maxillary palpus with palpomere 4
longer than 2 and 3 combined. Mentum ~1.5 times as wider as
long. Pronotum wider than long (PW/PL = 1.39) (Fig. 82);
anterior angles weakly projected; longitudinal median sulcus
conspicuous; longitudinal sulcus of posterior angles slightly
developed and reaching basal quarter of pronotum; abrupt
constriction at basal quarter; apical three-quarters with parallel
sides. Femur of anterior legs with short, hard setae on groove of
ventral margin. Elytra as long as wide (EL/BW = 1.00); stria 6 at
the basal half, sometimes inconspicuous, and stria 7 absent;
striae narrower than interstriae. Abdominal segments 3–6
parallel sides; tergite 9 with short ventral struts; tergite 10 with
two pairs of long setae on apex, apical the longest. Median lobe of
aedeagus with bulbous base in ventral view and not curved apex in
lateral view (Fig. 202); lateral lobes exceeding apex of median
lobe by one-third of its length in lateral view; internal sclerites as
in Fig. 203.
Female. Similar to male, except for: antennae reaching apex
of elytra; scape with fewer long setae; ovipositor as in P. sulcatus;
spermathecal duct ~1.5 times length of spermatheca, with
sclerotised basal region (Fig. 267); spermathecal capsule with
basal half suddenly narrower than apical and with projection on
base (Fig. 267).
Distribution
In the current study, this species was examined from Mexico
(Nayarit), Belize (Orange Walk), Guatemala (Alta Verapaz,
Izabal and Escuintla), El Salvador (Usulatán), Honduras
(Atlántida), Nicaragua (Granada and Río San Juan), Costa
Rica (Guanacaste, Heredia, San José, Cartago and Puntarenas),
Panama (Chiriquí, Veraguas, Colón, Panamá and Darién),
Dominica, Grenada, Trinidad and Tobago (Trinidad),
Colombia (Magdalena and Amazonas), Venezuela (Aragua),
Guyana, French Guiana (Saint-Laurent-du-Maroni and
Cayenne), Ecuador (Sucumbios, Pichincha, Napo and
Pastaza), Peru (Cusco and Madre de Dios), Bolivia (Beni,
Cochabamba and Santa Cruz), Brazil (Amazonas, Pará,
Rondônia, Goiás, São Paulo, Santa Catarina and Rio Grande
do Sul), Paraguay (Guairá) and Argentina (Tucumán). In
addition, Navarrete-Heredia et al. (2002) listed this species
from Mexico (Tabasco) and Cuba and Newton and Peck
(2006) listed it from the Galápagos Islands (Santa Cruz).
Biological notes
Piestus minutus has been found under bark, in leaf litter near
decaying logs, on fermenting fruits (‘berlese in fruit fall’) and on
fungi (‘fungusy log’). Some specimens were collected in flight
intercept or Malaise traps. This species also has been noticed
occurring together with P. fronticornis, P. bicornis and
P. pygmaeus.
530
Invertebrate Systematics
Remarks
This species is very similar to P. penicillatus but differs mainly
by transverse carina on base of vertex of head continuous
at the middle (Fig. 82), and long lateral lobes of aedeagus
(Fig. 202).
Piestus penicillatus has transverse carina interrupted at the
middle (Fig. 35), and short lateral lobes (Fig. 204).
Piestus minutus, as well as P. penicillatus, may be confused
with P. filicornis because they have similar integument of
pronotum with undulate microstriae and inconspicuous fine
punctures. However, they are easily separated by characters
cited under P. filicornis.
Piestus minutus is also similar to P. abdominalis and
P. niger, from which it differs by characters discussed under
P. filicornis.
Piestus penicillatus (Dalman, 1821)
(Figs 35, 83, 204, 205, 268)
Zirophorus penicillatus Dalman, 1821: 373 (original description, type
locality: ‘insula Guadeloupe’).
Piestus paenicillatus: Laporte, 1835: 129 (misspelling, as junior synonym
of P. sulcatus); Blackwelder, 1943: 46 (mention, error: Laporte, 1834:
129).
Piestus penicillatus: Erichson, 1840: 834 (characters, distribution);
Fauvel, 1864: 23 (characters, distribution); Blackwelder, 1943: 46
(characters, distribution, notes); Blackwelder, 1944: 100
(distribution); Herman, 2001b: 1793: 373 (distribution); NavarreteHeredia et al., 2002: 209 (distribution).
Piestus (Piestus) penicillatus: Bernhauer & Schubert, 1910: 7
(distribution); Scheerpeltz, 1952: 288 (characters, distribution).
Piestus erythropus Erichson, 1840: 834 (original description, type
locality: ‘Cuba’); Fauvel, 1864: 25 (characters, distribution); Sharp,
1887: 713 (note: discussion about distribution); Fauvel, 1902: 26 (note:
discussion about distribution); Herman, 2001b: 1790 (distribution).
syn. rest. (restored synonym).
Piestus (Piestus) erythropus: Bernhauer & Schubert, 1910: 7
(distribution); Scheerpeltz, 1952: 290 (characters, distribution). - As
synonym of P. penicillatus: Blackwelder, 1943: 46; Blackwelder,
1944: 100; Navarrete-Heredia et al., 2002: 209 (error: Dalman,
1822).
Trichocoryne striata Gray, 1832: 306, Fig. 5 (original description, type
locality: ‘West Indies’). - As junior synonym of P. erythropus:
Erichson, 1840: 834; Fauvel, 1864: 25; Bernhauer & Schubert,
1910: 7; Herman, 2001b: 1790; Navarrete-Heredia et al., 2002: 209.
- As junior synonym of P. penicillatus: Blackwelder, 1943: 46;
Blackwelder, 1944: 100. Note: Name preoccupied by Zirophorus
striatus Guérin-Méneville, 1829 (see P. bicornis above).
Type material
Zirophorus penicillatus Dalman, 1821. Syntype deposited in NHRS,
male, with labels: (1) ‘Guadeloupe/Forsström.’ [white label, printed in
black]. Note: In the original description Dalman (1821) did not specify
how many specimens he observed. We received from NHRS one
specimen, regarded as a syntype.
Piestus erythropus Erichson, 1840. Syntype deposited in ZMHB, male,
with labels: (1) ‘6807’ [white label, printed in black]; (2) ‘erythropus/Er.?/
Cuba Aiehr?’ [green label, handwritten]; (3) ‘SYNTYPUS/Piestus/
erythropus Erichson, 1840/labelled by MNHUB 2006’ [red label, printed
in black]. Note: In the original description Erichson (1840) did not specify
E. Caron et al.
how many specimens he observed. We received from ZMHB one specimen,
regarded as a syntype.
Trichocoryne striata Gray, 1832. Not found. Blackwelder (1943) cited the
type as ‘unknown’.
Additional material
See Appendix 2.
Redescription
BL: 4.6–6.7 mm, BW: 1.1–1.5 mm. Body flattened
dorsoventrally; light brown to reddish dark brown (Fig. 35).
Dorsal integument of head and pronotum with undulate
microstriae and inconspicuous fine punctures (Fig. 83); striae
and interstriae of elytra with inconspicuous fine punctures;
metaventrite, on lateral regions, and abdominal tergites, on
anterolateral regions, with moderately sized punctures.
Male. Head with front suddenly deflected (Fig. 83); Vshaped frontal sulcus complete; anterior angles curved and
prominent, and transverse carina on base, interrupted at
middle; eyes with line of three moderately sized punctures
with long setae on basal half of dorsal margin, basal the
longest. Antennae reaching apex of abdomen (Fig. 35); scape
with strongly enlarged area and long setae on basal half of
exposed dorsal face; scape shorter than antennomere 2 and 3
combined; antennomeres 4–11 oblong shape and conspicuously
narrowing from antennomere 4 to 11. Labrum with six setae
medially, equal in length; on each lateral third, antero-internal
seta shorter than postero-external seta. Mandibles with dorsal
teeth developed, not forming a bifurcate apex, shorter than
ventral; internal border asymmetrical, each with one acute
tooth at middle and, anterior to it, in right mandible with a
area that resemble a tooth, absent in left mandible. Maxillary
palpus with palpomere 4 longer than 2 and 3 combined. Mentum
~1.5 times as wide as long. Pronotum wider than long (PW/
PL = 1.31) (Fig. 83); anterior angles weakly projected;
longitudinal median sulcus conspicuous; longitudinal sulcus of
posterior angles slightly developed and reaching basal quarter of
pronotum; abrupt constriction at basal quarter; apical threequarters with parallel sides. Femur of anterior legs with short,
hard setae on groove of ventral margin. Elytra somewhat longer
than wide (EL/BW = 1.07); stria 6 at basal half, sometimes
inconspicuous, and stria 7 absent; striae narrower than
interstriae. Abdominal segments 3–6 parallel sides; tergite 9
with short ventral struts; tergite 10 with two pairs of long setae
on apex, apical the longest. Median lobe of aedeagus with
bulbous base in ventral view and not curved apex in lateral
view (Fig. 204); lateral lobes exceed a little apex of median
lobe in lateral view; internal sclerites as in Fig. 205.
Female. Similar to male, except for: antennae exceed a little
apex of elytra; scape with fewer long setae; ovipositor as in
P. sulcatus; spermathecal duct ~1.5 times length of spermatheca,
with sclerotised basal region (Fig. 268); spermathecal capsule
with basal half suddenly narrower than apical and with a
projection on the base (Fig. 268).
Distribution
In the current study, the species was examined from Mexico,
Cuba (Pina del Río, La Habana, Matanzas, Santiago de Cuba and
Revision of Piestus with remarks on related genera
Guantánamo), Dominican Republic, Puerto Rico and
Guadeloupe. The distribution in Mexico is doubtful (Sharp
1887; Fauvel 1902; Navarrete-Heredia et al. 2002). We
observed one old specimen, female, from ‘Sierra de Durango,
Mexique’ deposited in IRSNB.
Biological notes
Piestus penicillatus occurs on silk cotton tree, under bark of
logs, in rotten coconut husks, in old banana stalks and in rotten
cocoa pods (Blackwelder 1943).
Remarks
This species is very similar to P. minutus, from which it is easily
distinguished by characters cited above (see remarks under
P. minutus).
Piestus penicillatus, as well as Piestus minutus, may be
confused with P. filicornis because they share the integument
of pronotum with undulate microstriae and inconspicuous fine
punctures. However, they can be separated by characters cited
above (see remarks under P. filicornis).
Piestus penicillatus is also similar to P. abdominalis and
P. niger, which differ by characters discussed above (see
remarks under P. filicornis).
Piestus termitis, sp. nov.
(Figs 36, 84, 206, 207, 269)
Type material
Holotype deposited in SEMC, male, labels: (1) ‘Ecuador: Napo, mid./Rio
Tiputini, Yasumi Res./Stn. 040.50 S, 76240 W/28 July 1999. AKT#105’
[white label, printed in black]; (2) ‘Ex. Arboreal carton/termitarium TE#4/
A. Tishechkin’ [white label, printed in black]. 6 paratypes distributed: 4 in
SEMC: 1 female, the same labels of holotype; 1 sex undetermined, the same
first label, (2) ‘bar code/SM0023850/KUNHM-ENT’ [white label, printed in
black]; 1 female, labels: (1) ‘Ecuador: Sucumbios/Sacha Lodge, 0.5S,/
76.5W, 270m, 27-VIII-10-IX/1994, Hibbs, ex: malaise’ [white label,
printed in black]; (2) ‘bar code/SM0023742/KUNHM-ENT’ [white label,
printed in black]; 1 male, labels: (1) ‘Ecuador: Sucumbios/Sacha Lodge,
0.5S,/76.5W, 270m, 31-X-10-XI/1994, Hibbs, ex: malaise’ [white label,
printed in black], ‘bar code/SM0023085/KUNHM-ENT’ [white label,
printed in black]; 1 in FMNH: sex undetermined, labels: (1) ‘Peru: Madre
de Dios/Cocha Cashu Bio. Stn./Manu National Park, 350m/11530 45’S,
71240 24’W/17–19 Oct 2000; R.Brooks/PERU1B00 042/ex: flight
intercept trap’ [white label, printed in black], ‘bar code/SM0257917/
KUNHM-ENT’ [white label, printed in black]; 1 in DZUP: male, labels:
(1) ‘Ecuador: Sucumbios/Sacha Lodge, 0.5S,/76.5W, 270m, 16-29-VIII-/
1994, Hibbs, ex: malaise’ [white label, printed in black]; (2) ‘bar code/
SM0024457/KUNHM-ENT’ [white label, printed in black].
Description
BL: 4.2–4.3 mm, BW: 1.2–1.3 mm. Body somewhat convex;
head, pronotum and abdomen black and elytra reddish
(Fig. 36). Dorsal integument of head with undulate microstriae
and moderately sized punctures, not contiguous (Fig. 84);
pronotum with large punctures, sometimes contiguous, there
are some areas not punctured and undulate microstriae only on
lateral areas; striae of elytra with inconspicuous fine punctures
and interstriae with undulate microstriae; abdominal tergites
with moderately sized punctures uniformly distributed.
Invertebrate Systematics
531
Male. Head with front slightly deflected (Fig. 84); V-shaped
frontal sulcus complete; anterior angles curved and weakly
prominent; eyes with one moderately sized punctures with
long setae on basal half of dorsal margin of eyes. Antennae
exceed a little apex of elytra (Fig. 36); scape with one long setae
at middle of dorsal face; scape subequal to antennomere 2 and
3 combined in length; antennomeres 4–11 oblong shape. Labrum
with six setae medially, internal the shortest; each lateral third,
antero-internal seta shorter than postero-external seta.
Epipharynx as visible dorsally with two lobes at apex.
Mandibles without dorsal teeth developed; internal border
symmetrical, smooth. Maxillary palpus with palpomere 4
longer than 2 and 3 combined. Mentum ~1.5 times as wide as
long. Pronotum wider than long (PW/PL = 1.48) (Fig. 84);
anterior angles not projected; on basal half a slightly
transverse
depression;
longitudinal
median
sulcus
conspicuous; longitudinal sulcus of posterior angles developed
and reaching the middle of pronotum; abrupt constriction at basal
third; and, anterior to it, in each side with a slight emargination;
apical two-thirds with sinuous sides. Femur of anterior legs with
short, hard setae on groove of ventral margin. Elytra as long as
wide (EL/BW = 0.94); stria 6 absent and stria 7 incomplete, only
on apical half; striae narrower than interstriae. Metaventrite with
conspicuous median depression near apex. Abdominal segments
3 and 4 slightly narrower than 5–7; tergite 9 with long ventral
struts; tergite 10 with two pairs of long setae on apex, apical the
longest. Median lobe of aedeagus with bulbous base in ventral
view and curved apex in lateral view (Fig. 206); lateral lobes
exceeding the apex of median lobe by one-quarter of its length in
lateral view; internal sclerites as in Fig. 207.
Female. Similar to male, except for: ovipositor as in
P. sulcatus; spermathecal duct about same length of
spermatheca, without sclerotised basal region (Fig. 269);
spermatheca as in Fig. 269.
Distribution
Ecuador (Sucumbios and Napo) and Peru (Madre de Dios).
Biological notes
Piestus termitis has been collected in Malaise and flight intercept
traps. It was also cited occurring in a termitarium, but its
association with termites needs to be confirmed.
Remarks
Piestus termitis is very similar to P. surrufus because they share
some characters, such as: complete V-shaped frontal sulcus
(Figs 84, 85), prominent eyes from above, antenna short (male
and female), two lobes at the apex of epipharynx (Fig. 111),
symmetrical internal border of mandibles (Fig. 126), incomplete
stria 7 on elytra and undulate microstriae on interstriae, and
abdominal segment 3 and 4 narrower than 5–7 (Figs 36, 37).
However, P. termitis may be easily separated by having one long
setae on dorsal margin of the eyes, one long setae on scape (male
and female), large punctures on pronotum (Fig. 84), long lateral
lobes of aedeagus, exceeding the apex of median lobe (Fig. 206),
and short spermathecal duct, about the same length as
spermatheca (Fig. 269).
532
Invertebrate Systematics
Piestus surrufus has two long setae on dorsal margin of the
eyes, slightly enlarged area and long setae on scape of male and
only two long setae on female, inconspicuous fine punctures and
some moderately sized punctures dispersed on pronotum
(Fig. 85), short lateral lobes of aedeagus, reaching the apex of
median lobe (Fig. 208), and long spermathecal duct, ~3 times the
length of spermatheca (Fig. 270).
Etymology
The specific name is derived from the Latin word termes (termite),
in allusion to the presumed association of this species with
termites, as some type specimens were found in an arboreal
carton termitarium.
Piestus surrufus, sp. nov.
(Figs 37, 85, 111, 126, 208, 209, 270)
Type material
Holotype deposited in SEMC, female, labels: (1) ‘Nicaragua: Río San Juan/
Dept., 8 km SE EI Castillo/Refugio Bartole, 30m, 1058.60 N/8420.40 W,
23-31-V-2002/S.Peck 02-09, ex. flight/intercept trap, NIC1P02 002’ [white
label, printed in black]; (2) ‘bar code/SM0561263/KUNHM-ENT’ [white
label, printed in black]. 5 paratypes deposited in the following collections: in
SEMC, 1 male, the same first label of holotype, (2) ‘bar code/SM0561144/
KUNHM-ENT’ [white label, printed in black]; 1 female, the same first label,
(2) ‘bar code/SM0561230/KUNHM-ENT’ [white label, printed in black];
1 male, labels: (1) ‘Nicaragua: Río San Juan Dept./60 km SE San Carlos,
Refúgio/Bartola, 100m 1058.400 N, 8420.300 W/25-28-V-2002, R.Brooks,
Z.Falin,/S.Chatzimanolis ex. flight intercept/trap, NIC1BFC02 112’ [white
label, printed in black]; (2) ‘bar code/SM539779/KUNHM-ENT’ [white
label, printed in black]; in FMNH, 1 female, labels: (1) ‘Ecuador: Pastaza
Prov.;/Ashuara indian village on/Rio Macuma nr. Rio Morona./Leg:
B. Malkin VII.11-16.1971’ [white label, printed in black]; (2) ‘Beating/
dry/foliage’ [white label, printed in black]; in ZMHB, 1 sex undetermined,
label: (1) ‘Peru/VI’ (ZMHB); in DZUP, 1 female, the same first label of
holotype, (2) ‘bar code/SM0561230/KUNHM-ENT’ [white label, printed in
black].
Description
BL: 4.4–4.8 mm, BW: 1.1–1.2 mm. Body somewhat convex;
head, pronotum and abdomen reddish brown to reddish dark
brown and elytra reddish with external and apical margins region
darker (Fig. 37). Dorsal integument of head and pronotum with
undulate microstriae and inconspicuous fine punctures, some
moderately sized punctures on pronotum near base of
longitudinal median sulcus and on longitudinal sulcus of
posterior angles (Fig. 85); striae of elytra with inconspicuous
fine punctures and interstriae with undulate microstriae;
abdominal tergites with moderately sized punctures uniformly
distributed.
Male. Head with front slightly deflected (Fig. 85); V-shaped
frontal sulcus complete; anterior angles curved and weakly
prominent; eyes with two moderately sized punctures with
long setae on basal half of dorsal margin of eyes, basal the
longest. Antennae exceed a little apex of elytra (Fig. 37);
scape with slightly enlarged area and long setae on basal half
of exposed dorsal face; scape subequal to antennomere 2 and 3
combined in length; antennomeres 4–11 oblong shape. Labrum
with six setae medially, internal the shortest (Fig. 111); on each
E. Caron et al.
lateral third, antero-internal seta shorter than postero-external
seta. Epipharynx as visible dorsally with two lobes at apex
(Fig. 111). Mandibles without dorsal teeth (Fig. 126); internal
border symmetrical, smooth. Maxillary palpus with palpomere 4
longer than 2 and 3 combined. Mentum ~1.5 times as wide as
long. Pronotum wider than long (PW/PL = 1.43) (Fig. 85);
anterior angles not projected; on basal half a slightly
transverse
depression;
longitudinal
median
sulcus
conspicuous; longitudinal sulcus of posterior angles developed
and reaching middle of pronotum; abrupt constriction at basal
third and, anterior to it, in each side with a slight emargination;
apical two-thirds with slightly sinuous sides. Femur of anterior
legs with short, hard setae on groove of ventral margin. Elytra as
long as wide (EL/BW = 1.00); stria 6 at the basal half and stria 7
incomplete, only on apical two-thirds; striae narrower than
interstriae. Metaventrite with slightly median depression near
apex. Abdominal segments 3 and 4 slightly narrower than 5–7;
tergite 9 with long ventral struts; tergite 10 with two pairs of long
setae on apex, apical the longest. Median lobe of aedeagus with
bulbous base in ventral view and curved apex in lateral view
(Fig. 208); lateral lobes reaching apex of median lobe in lateral
view; internal sclerites as in Fig. 270.
Female. Similar to male, except for antennal scape without
enlarged area and with two long setae at middle of dorsal face;
ovipositor as in P. sulcatus; spermathecal duct long, ~3 times
length of spermatheca, without sclerotised basal region
(Fig. 270); spermatheca as in Fig. 270.
Distribution
Nicaragua (Río San Juan), Ecuador (Pastaza) and Peru.
Biological notes
Piestus surrufus has been collected in flight intercept traps. One
specimen from Ecuador was collected by beating dry foliage.
Remarks
Piestus differs from P. termitis by characters cited under remarks
of P. termitis.
Etymology
The name refers to the colour of the elytra.
Piestus formicinus, sp. nov.
(Figs 38, 86, 210, 211, 271)
Type material
Holotype deposited in SEMC, male, labels: (1) ‘Guiana: Region 8/
Iwokrama Forest, 1 km W Kurupukari/Iwokrama Field Stn., 60m/
4400 19’N, 58410 4’W, 21 May 2001, R.Brooks, Z.Falin/GUY1BF01 004/
ex: fogging fungusy logs’ [white label, printed in black]; (2) ‘bar code/
SM0226160/KUNHM-ENT’ [white label, printed in black]. 25 paratypes
distributed: 15 SEMC, 3 AMNH, 3 DZUP, 3 FMNH and 1 MZSP, which: 1
female, the same first label as holotype, (2) ‘bar code/SM0266113/KUNHMENT’ [white label, printed in black] (SEMC); 1 sex undetermined, the same
first label, (2) ‘bar code/SM0226214/KUNHM-ENT’ [white label, printed in
black] (FMNH); 1 sex undetermined, the same first label, (2) ‘bar code/
SM0226229/KUNHM-ENT’ [white label, printed in black] (DZUP); 1 sex
undetermined, the same first label, (2) ‘bar code/SM0226230/KUNHM-
Revision of Piestus with remarks on related genera
Invertebrate Systematics
533
ENT’ [white label, printed in black] (SEMC); 1 sex undetermined, the same
first label, (2) ‘bar code/SM0226231/KUNHM-ENT’ [white label, printed in
black] (SEMC); 1 sex undetermined, the same first label, (2) ‘bar code/
SM0226232/KUNHM-ENT’ [white label, printed in black] (SEMC); 1 sex
undetermined, the same first label, (2) ‘bar code/SM0226233/KUNHMENT’ [white label, printed in black] (SEMC); 1 sex undetermined, the
same first label, (2) ‘bar code/SM0226216/KUNHM-ENT’ [white label,
printed in black] (SEMC); 1 sex undetermined, the same first label, (2)
‘bar code/SM0226205/KUNHM-ENT’ [white label, printed in black]
(SEMC); 1 sex undetermined, the same first label, (2) ‘bar code/
SM0226212/KUNHM-ENT’ [white label, printed in black] (SEMC); 1 sex
undetermined, the same first label, (2) ‘bar code/SM0226224/KUNHMENT’ [white label, printed in black] (SEMC); 1 sex undetermined, the
same first label, (2) ‘bar code/SM0226223/KUNHM-ENT’ [white label,
printed in black] (SEMC); 1 sex undetermined, the same first label, (2)
‘bar code/SM0226228/KUNHM-ENT’ [white label, printed in black]
(SEMC); 1 sex undetermined, the same first label, (2) ‘bar code/
SM0226221/KUNHM-ENT’ [white label, printed in black] (SEMC); 1 sex
undetermined, the same first label, (2) ‘bar code/SM0226220/KUNHMENT’ [white label, printed in black] (MZSP); 1 sex undetermined, the
same first label, (2) ‘bar code/SM0226219/KUNHM-ENT’ [white label,
printed in black] (AMNH); 1 sex undetermined, the same first label, (2)
‘bar code/SM0226218/KUNHM-ENT’ [white label, printed in black]
(AMNH); 1 sex undetermined, the same first label, (2) ‘bar code/
SM0226213/KUNHM-ENT’ [white label, printed in black] (FMNH); 1
sex undetermined, the same first label, (2) ‘bar code/SM0226193/
KUNHM-ENT’ [white label, printed in black] (SEMC); 1 sex
undetermined, the same first label, (2) ‘bar code/SM0226210/KUNHMENT’ [white label, printed in black] (DZUP); 1 sex undetermined, the
same first label, (2) ‘bar code/SM0226209/KUNHM-ENT’ [white label,
printed in black] (DZUP); 1 sex undetermined, the same first label, (2)
‘bar code/SM0226208/KUNHM-ENT’ [white label, printed in black]
(SEMC); 1 sex undetermined, the same first label, (2) ‘bar code/
SM0226207/KUNHM-ENT’ [white label, printed in black] (SEMC); 1 sex
undetermined, the same first label, (2) ‘bar code/SM0226206/KUNHMENT’ [white label, printed in black] (FMNH); 1 sex undetermined, the
same first label, (2) ‘bar code/SM0226226/KUNHM-ENT’ [white label,
printed in black] (AMNH).
developed; internal border slightly asymmetrical, each with
one small and acute tooth at middle and, anterior to it in right
mandible with a very weak area that resembles a tooth, absent in
left mandible. Maxillary palpus with palpomere 4 longer than 2
and 3 combined. Mentum ~1.5 times as wide as long. Pronotum
wider than long (PW/PL = 1.35) (Fig. 38); anterior angles weakly
projected; longitudinal median sulcus conspicuous; longitudinal
sulcus of posterior angles gently developed and reaching the basal
third of pronotum; abrupt constriction at basal third; apical
two-thirds with curved sides. Femur of anterior legs with
short, hard setae on groove of ventral margin. Elytra as long as
wide (EL/BW = 0.94); stria 6 absent and stria 7 complete; striae
narrower than interstriae. Abdominal segments 3–6 parallel
sides; tergite 9 with short ventral struts; tergite 10 with two
pairs of long setae on apex, apical the longest. Median lobe of
the aedeagus with bulbous base in ventral view and slightly
curved apex in lateral view (Fig. 210); lateral lobes not
reaching the apex of median lobe in lateral view; internal
sclerites as in Fig. 211.
Female. Similar to male, except for: antennae not reaching
apex of elytra; scape without enlarged area and with two long
setae at middle of the dorsal face; ovipositor as in P. sulcatus;
spermathecal duct about same length of spermatheca, without
sclerotised basal region (Fig. 271); spermatheca as in Fig. 271.
Additional material
Remarks
See Appendix 2.
Piestus formicinus is easily separated from the other species of
Piestus by incomplete V-shaped frontal sulcus (Fig. 86), slightly
prominent eyes from above, slightly enlarged area with long
setae on scape only on male, undulate microstriae and moderately
sized punctures equally distributed on pronotum.
Description
BL: 2.6–3.6 mm, BW: 0.8–0.9 mm. Body somewhat convex;
light brown to reddish dark brown (Fig. 38). Dorsal
integument of head and pronotum with undulate microstriae
and moderately sized punctures (Fig. 86); striae and interstriae
of elytra with conspicuous fine punctures; metaventrite, only on
lateral regions, and abdominal tergites with moderately sized
punctures uniformly distributed; dorsal teeth of mandibles
weakly developed.
Male. Head with front slightly deflected (Fig. 86); V-shaped
frontal sulcus incomplete, curved arms not joined medially;
anterior angles curved and weakly prominent; eyes with two
moderately sized punctures with long setae on basal half of dorsal
margin of eyes, basal the longest. Antennae exceed a little apex of
elytra (Fig. 38); scape with slightly enlarged area and long setae
on the basal half of exposed dorsal face; scape subequal to
antennomere 2 and 3 combined in length; antennomeres 4–11
oblong shape. Labrum with six setae medially, internal the
longest; each lateral third, antero-internal seta shorter than
postero-external seta. Mandibles with dorsal teeth weakly
Distribution
Guyana (Upper Demerara-Berbice).
Biological notes
Piestus formicinus may be associated with ants of the genus
Acromyrmex, or at least with its refuse material (‘ex: Acromyrmex
hystrix refuse pile’). The type series was captured by fogging
fungus logs.
Etymology
The specific name is derived from the Latin word formica (ant), in
allusion the presumed association of this species with ants, as
some specimens were found in refuse piles of the genus
Acromyrmex Mayr, 1865.
Piestus sulcatus Gravenhorst, 1806
(Figs 5–10, 106–108, 113–115, 131, 134, 139, 149, 153,
212–214, 272)
Piestus sulcatus Gravenhorst, 1806: 224 (original description, type
locality: ‘Brasilia’); Erichson, 1840: 835 (characters, distribution);
Fauvel 1864: 30 (note, distribution); Sharp, 1876: 407 (note,
distribution); Blackwelder, 1943: 44 (characters, distribution, notes);
Blackwelder, 1944: 101 (distribution); Blackwelder, 1952: 309
(mention); Herman, 2001b: 1795 (distribution).
534
Invertebrate Systematics
Piestus (Piestus) sulcatus: Bernhauer & Schubert, 1910: 7 (distribution);
Scheerpeltz, 1952: 286 (distribution).
Piestus sanctae-catharinae Bernhauer, 1906: 193 (original description,
type locality: ‘Brasilia (Santa Catharina)’); Blackwelder, 1944: 101
(distribution). New synonym.
Piestus (Piestus) sanctae-catharinae: Bernhauer & Schubert, 1910: 7
(distribution); Scheerpeltz, 1952: 286 (characters, distribution).
Piestus sanctaecatharinae: Herman, 2001b: 1794 (distribution).
Piestus condei Wendeler, 1955: 187 (original description, type locality:
‘Brasilien, Espirito Santo, Sta. Theresa’); Herman, 2001b: 1790
(distribution). New synonym.
Type material
Piestus sulcatus Gravenhorst, 1806. Lectotype (here designated)
deposited in ZMHB, male, with labels: (1) ‘Para/Sieber/Nr. 6812’ [green
label, handwritten]; (2) ‘Typus’ [red label, printed in black]; (3) ‘Zool. Mus./
Berlin’ [light yellow label, printed in black]; (4) ‘SYNTYPUS/Piestus/
sulcatus Gravenhorst, 1806/labelled by MNHUB 2006’ [red label, printed
in black]. Three paralectotypes deposited in ZMHB, 2 females, with the same
labels of lectotype, and 1 male [specimen damaged, without abdomen], with
labels: (1) ‘6810’ [white label, printed in black]; (2) ‘sulcatus/Gr. x/Pará Sieb’
[green label, handwritten]; (3–5) [third to fifth labels are the same of second to
fourth of lectotype]. Note: In the original description Gravenhorst (1806)
did not specify how many specimens he used for description. We received
4 specimens with type labels ‘Typus’ from ZMHB from which we are
considering as type material.
Piestus sanctae-catharinae Bernhauer, 1906. Lectotype (here designated)
deposited in NMW, male, with labels: (1) ‘S.Catharina/Brasil. Klimsch’
[white label/printed in black]; (2) ‘Piestus Sanctae/Catharinae Bh.’ [white
label, handwritten]; (3) ‘Klimsch/donavit’ [white label, handwritten]; (4) ‘ex
coll./Scheerpeltz’ [blue label, printed in black]; (5) ‘COTYPUS/Piestus/
Sanctae Catharin/Bernhauer.’ [rose label, first line printed in black and the
others handwritten]; (6) ‘Sanctae-/catharinae Bh.’ [green label, handwritten].
Two paralectotypes, 1 female deposited in NMW with the same labels of
lectotype, except for: (2) ‘Sanctae Catharinae/Bernh./det. Bernhauer’ [white
label, two first lines handwritten and third line printed in black], and 1 female
deposited in FMNH with the same labels of lectotype, except for: (2) ‘Sanctae
Cathari’/nae Bernh./Typus.’ [light yellow label, Bernhaeur’s handwritten];
(3) ‘Chicago NHMus/M.Bernhauer/Collection’ [white label, printed in
black]. Note: In the original description Bernhauer (1906) did not specify
how many specimens he used for description. We received 3 specimens with
type labels from NMW and FMNH: we are considering all three as type
material.
Piestus condei Wendeler, 1955. Holotype deposited in ZMHB, male, with
labels: (1) ‘EspiritoSanto/Sta. Theresa/Jan.29.Conde’ [white label, printed in
black]; (2) ‘,’ [white label, printed in black]; (3) ‘Piestus s.str./condei n.sp.
Wdlr.’ [white label, Wendeler’s handwritten]; (4) ‘Holotypus’ [red label,
printed in black]; (5) ‘Sammlung/Wendeler’ [white label, printed in black]; (6)
‘condei */Wdlr.’ [green label, handwritten]. Note: In the original description
Wendeler (1955) specified one female studied; however, the type observed
(ZMHB) is male.
E. Caron et al.
metaventrite, only on lateral regions, and abdominal tergites
with moderately sized punctures uniformly distributed.
Male. Head with front slightly deflected (Fig. 7); V-shaped
frontal sulcus incomplete (gently complete in some species);
anterior angles curved and weakly prominent; eyes with two
moderately sized punctures with long setae on basal half of
dorsal margin, basal the longest. Antennae almost reaching
apex of abdomen (reach in some species) (Fig. 5); scape with
slightly enlarged area and long setae on basal half of exposed
dorsal face (Fig. 106); scape shorter than antennomere 2 and 3
combined; antennomeres 4–11 oblong shape. Labrum with six
setae medially, equal in length (Fig. 108); each lateral third,
antero-internal seta shorter than postero-external seta. Mandibles
with dorsal teeth weakly developed (Fig. 113); internal border
slightly asymmetrical, each with one small and acute tooth at
middle and, anterior to it in right mandible with weak area that
resembles a tooth, absent in left mandible. Maxillary palpus with
palpomere 4 longer than 2 and 3 combined (Fig. 130). Mentum
~1.5 times as wide as long (Fig. 134). Pronotum wider than long
(PW/PL = 1.49) (Fig. 7); anterior angles weakly projected;
longitudinal median sulcus conspicuous; longitudinal sulcus of
posterior angles developed and reaching middle of pronotum;
abrupt constriction at basal third and, anterior to it, in each side
with short tooth and slight emargination; apical two-thirds with
curved sides. Prosternum with conspicuous ovate set of fine
punctures in median region near apex (Fig. 8). Femur of
anterior legs with short, hard setae on groove of ventral
margin. Elytra as long as wide (EL/BW = 0.97) (Fig. 7); stria 6
at basal quarter; stria 7 complete; striae narrower than interstriae;
epipleural carina developed (Fig. 6). Metaventrite with slightly
median depression near apex (Fig. 9). Abdominal segments 3–6
subparallel sides, segments 3 and 4 slightly narrower than 5–7;
sternite 3 with conspicuous transverse carina (Fig. 10); tergite 9
with short ventral struts (Fig. 149); tergite 10 with two pairs of
long setae on apex, apical the longest. Median lobe of the
aedeagus with bulbous base in ventral view and curved apex
in lateral view (Figs 212, 214); lateral lobes exceeding apex of
median lobe by one-quarter of its length in lateral view; internal
sclerites as in Fig. 213.
Female. Similar to male, except for: antennae reaching apex
of elytra; scape without enlarged area and with two long setae at
middle of the dorsal face (Fig. 107); prosternum without ovate set
of fine punctures; ovipositor with curved external margin
(Fig. 153); spermathecal duct ~1.5 times length of
spermatheca, without sclerotised basal region (Fig. 272);
spermatheca somewhat elongate (Fig. 272).
Additional material
Distribution
See Appendix 2.
In the current study, the species was examined from Nicaragua
(Rio San Juan), Costa Rica (Guanacaste, Alajuela, Heredia,
Cartago, Limón and Puntarenas), Panama (Veraguas, Colón,
Panamá and Darién), Guadeloupe, Dominica (Saint David),
Martinique, Saint Vincent and the Grenadines (Saint Vincent),
Trinidad and Tobago (Trinidad), Colombia (Choco), Venezuela
(Aragua), Guyana (Upper Demerara-Berbice), Suriname
(Marowijne), French Guiana (Saint-Laurent-du-Maroni and
Cayenne), Ecuador (Esmeraldas, Sucumbios, Pichincha, Napo,
Cotopaxi and Pastaza), Peru (Loreto, San Martín, Huancavelica,
Redescription
BL: 3.8–5.8 mm, BW: 1.1–1.6 mm. Body flattened
dorsoventrally; light brown to reddish dark brown (Fig. 6).
Dorsal integument of head and pronotum with undulate
microstriae and fine to moderately sized punctures, not
contiguous (Fig. 7); striae of elytra with moderately sized
punctures and interstriae with conspicuous fine punctures;
Revision of Piestus with remarks on related genera
Cusco and Madre de Dios), Bolivia (Cochabamba and Santa
Cruz), and Brazil (Amazonas, Pará, Bahia, Goiás, Distrito
Federal, Rio de Janeiro, São Paulo, Paraná and Santa
Catarina). Herman (2001b) listed P. sanctaecatharinae (junior
synonym of P. sulcatus) from Paraguay too; Newton and Peck
(2006) also report P. sulcatus from the Galápagos Islands (Isabela
and Santa Cruz).
Biological notes
Piestus sulcatus has been found in leaf litter, fruit falls or
under bark of decaying logs. Six specimens from Panamá were
observed in ‘7 day old tapir dung’. This species also has been
collected in flight intercept and Malaise traps, Berlese extraction
of leaf litter, and at light trap. Blackwelder (1943) noted some
similar habitats.
Remarks
Piestus sulcatus is similar to P. gounellei and P. mexicanus
differing from those species by incomplete V-shaped frontal
sulcus (gently complete in some specimens) (Fig. 7), undulate
microstriae and moderately sized punctures on head and
pronotum, developed stria 6 on elytra (Fig. 5), lateral lobes of
aedeagus exceeding the apex of median lobe by one-quarter of its
length (Fig. 212), and spermatheca elongate (Fig. 272).
Piestus gounellei has a complete V-shaped frontal sulcus
(Fig. 87), moderate to large punctures on head and pronotum,
undulate microstriae only on lateral areas, developed stria 6, short
lateral lobes exceeding a little the apex of median lobe (Fig. 215),
and globose apex of spermatheca (Fig. 273).
Piestus mexicanus has a complete V-shaped frontal sulcus
(Fig. 88), large punctures on head and pronotum, undulate
microstriae only on lateral areas, no stria 6, short lateral lobes
exceeding a little the apex of median lobe (Fig. 218), and
somewhat elongate spermatheca (Fig. 274).
Piestus gounellei Fauvel, 1902
(Figs 39, 87, 215–217, 273)
Piestus gounellei Fauvel, 1902: 24 (original description, type locality:
‘Brésil: Minas Geraes, Caraça’); Blackwelder, 1944: 100 (distribution);
Herman, 2001b: 1791 (distribution).
Piestus (Piestus) gounellei: Bernhauer & Schubert, 1910: 7 (distribution);
Scheerpeltz, 1933: 994 (mention); Scheerpeltz, 1952: 285 (characters,
distribution).
Piestus wasmanni Fauvel, 1902: 25 (original description, type locality:
‘Rio de Janeiro, Colonia Alpina’); Blackwelder, 1944: 101
(distribution); Herman, 2001b: 1795 (distribution). New synonym.
Piestus (Piestus) wasmanni: Bernhauer & Schubert, 1910: 7
(distribution); Scheerpeltz, 1952: 285 (characters, distribution).
Type material
Piestus gounellei Fauvel, 1902. Holotype deposited in IRSNB, male, with
labels: (1) ‘Caraça(Minas Geraes)/Brésil/E.Gounelle 1.2.1885’ [white label,
printed in black]; (2) ‘gounellei/Fvl.’ [handwritten]; (3) ‘R.I.Sc.N.B. 17.479
[printed in black]/Piestus [handwritten]/Coll. et det. A. Fauvel [printed in
black]’; (4) ‘Type’ [red label, printed in black]. Note: In the original
description Fauvel (1902) specified only one specimen studied.
Piestus wasmanni Fauvel, 1902. Holotype deposited in IRSNB,
female, with labels: (1) ‘Col. Alpina/Rio de Jan.’ [white label,
Invertebrate Systematics
535
handwritten]; (2) ‘Wasmanni/FvL.’ [white label, handwritten]; (3) ‘R.I.Sc.
N.B. 17.479/Piestus/Coll. et det. A. Fauvel’ [white label, first and third lines
printed in black and second line handwritten]; (4) ‘Type’ [red label, printed in
black]. Note: In the original description Fauvel (1902) specified only one
specimen studied.
Additional material
See Appendix 2.
Redescription
BL: 4.4–5.9 mm, BW: 1.2–1.6 mm. Body flattened
dorsoventrally; reddish brown to reddish dark brown (Fig. 39).
Dorsal integument of head with moderately sized punctures,
not contiguous (Fig. 87); pronotum with moderate to large
punctures, sometimes contiguous, there are some areas not
punctured and, undulate microstriae only on lateral areas;
striae of elytra with moderately sized punctures and interstriae
with inconspicuous fine punctures; metaventrite, only on lateral
regions, and abdominal tergites with moderately sized punctures
uniformly distributed.
Male. Head with front slightly deflected (Fig. 87); V-shaped
frontal sulcus complete; anterior angles curved and prominent,
slightly median fovea at base; eyes with two moderately sized
punctures with long setae on basal half of dorsal margin, basal
the longest. Antennae reaching apex of abdomen (Fig. 39);
scape with slightly enlarged area and long setae on basal half
of exposed dorsal face; scape shorter than antennomere 2 and 3
combined; antennomeres 4–11 oblong shape. Labrum with six
setae medially, equal in length; each lateral third, antero-internal
seta shorter than postero-external seta. Mandibles with dorsal
teeth weakly developed; internal border slightly asymmetrical,
each with one small and acute tooth at middle and, anterior to it
in right mandible with weak area that resembles a tooth, absent
in left mandible. Maxillary palpus with palpomere 4 longer than
2 and 3 combined. Mentum ~1.5 as wide as long. Pronotum
wider than long (PW/PL = 1.41) (Fig. 87); anterior angles
weakly projected; longitudinal median sulcus conspicuous;
longitudinal sulcus of posterior angles developed and reaching
the middle of pronotum; abrupt constriction at basal third and,
anterior to it, in each side with a short tooth and slight
emargination; apical two-thirds with curved sides. Prosternum
with conspicuous ovate set of fine punctures in median region
near apex. Femur of anterior legs with short, hard setae on
groove of ventral margin. Elytra as long as wide (EL/
BW = 1.04); stria 6 at basal half and stria 7 complete; striae
narrower than interstriae; epipleural carina developed.
Metaventrite with slightly median depression near apex.
Abdominal segments 3–6 subparallel sides, segments 3 and 4
slightly narrower than 5–7; sternite 3 with conspicuous
transverse carina; tergite 9 with short ventral struts; tergite 10
with two pairs of long setae on apex, apical the longest.
Median lobe of aedeagus with bulbous base in ventral view,
curved apex in lateral view, and projected apex in ventral
view (Figs 215, 217); lateral lobes exceeding a little the
apex of median lobe in lateral view; internal sclerites as in
Fig. 273.
Female. Similar to male, except for: antennae
exceeding apex of elytra, but not reaching apex of abdomen;
scape without enlarged area and with some long setae on
536
Invertebrate Systematics
basal half of exposed dorsal face; prosternum without ovate set of
fine punctures; ovipositor as in P. sulcatus; spermathecal
duct about same length of spermatheca, without sclerotised
basal region (Fig. 273); spermatheca globose at apex
(Fig. 273).
Distribution
In the current study, the species was examined from Brazil
(Espirito Santo, Rio de Janeiro, São Paulo, Paraná and Santa
Catarina), Paraguay (Caazapá and Itapúa) and Argentina
(Misiones).
Biological notes
Piestus gounellei has been found in litter or on or under bark of
decaying logs. This species also has been collected in flight
intercept traps.
Remarks
This species is similar to P. sulcatus and P. mexicanus, but differs
by characters cited above (see remarks under P. sulcatus).
Piestus mexicanus Laporte, 1835
(Figs 40, 88, 218, 219, 274)
Piestus mexicanus Laporte, 1835: 130 (original description, type locality:
‘Mexique’); Erichson, 1840: 836 (characters, distribution); Fauvel,
1864: 30 (characters, distribution); Sharp, 1887: 714 (characters,
distribution); Blackwelder, 1944: 100 (distribution, error: Laporte,
1834: 30); Herman, 2001b: 1792 (distribution); Navarrete-Heredia
et al., 2002: 208 (distribution); Newton et al., 2005: 37 (distribution).
Piestus (Piestus) mexicanus: Bernhauer & Schubert, 1910: 7 (distribution,
error: Laporte, 1834: 30); Scheerpeltz, 1952: 287 (distribution, error:
Laporte, 1834: 30).
Piestus alternans Sharp, 1887: 714 (original description, type locality:
‘Panama, David in Chiriqui’); Blackwelder, 1944: 100 (distribution);
Herman, 2001b: 1788 (distribution). New synonym.
Piestus (Piestus) alternans: Bernhauer & Schubert, 1910: 6 (distribution);
Scheerpeltz, 1952: 287 (distribution).
Type material
Piestus mexicanus Laporte, 1835. Syntype deposited in BMNH, female
[damaged apex of antennae and abdomen], with labels: (1) ‘Piestus mexica-/
nus Type ex./coll. Chevrolat./Mexico. Lesneut.?’ [white label, together with
the specimen, handwritten]; (2) ‘Type’ [circle white label with border red,
printed in black]; (3) ‘Mexico.’ [white label, printed in black]; (4) ‘Mexico’
[white label, handwritten]; (5) ‘B.C.A. Col. I. 2./Piestus/mexicanus,/Lap.’
[white label, printed in black]; (6) ‘27.?’ [light red label, handwritten]; (7)
‘Sharp Coll./1905-313.’ [white label, printed in black]; (8) ‘Piestus/
mexicanus Lap. Er./836 11.? type tolp?/Mexico Oaxaca? D. Lesneur.’
[green label, handwritten, note: ‘Colombia’ handwritten underside]. Note:
In the original description Laporte (1835) did not specify how many
specimens he observed. We received from BMNH only one specimen
labelled as type.
Piestus alternans Sharp, 1887. Holotype deposited in BMNH, male
[broken right antenna], with labels: (1) ‘Piestus alternans/ Type D.S./David
Panama/Champion’ [white label, together with the specimen, handwritten];
(2) ‘Holo-/type’ [circle white label with border red, printed in black]; (3)
‘David,/Panama./Champion.’ [white label, printed in black]; (4) ‘B.C.A. Col.
i. 2./Piestus/alternans, Sharp.’ [white label, printed in black]; (5) ‘Sharp Coll./
1905-313.’ [white label, printed in black]; (6) ‘Holotype/Piestus/alternans/
Sharp, 1876/det. R.G. Booth 2008’ [white label, the four first lines
E. Caron et al.
handwritten and the last one printed in black]. Note: In the original
description Sharp (1887) specified only one specimen (male) observed.
Additional material
See Appendix 2.
Redescription
BL: 3.8–5.4 mm, BW: 1.1–1.5 mm. Body flattened
dorsoventrally; light brown to reddish dark brown (Fig. 40).
Dorsal integument of head and pronotum with large punctures
(Fig. 88), sometimes contiguous, there are some areas not
punctured and, undulate microstriae only on lateral areas;
striae of elytra with large punctures and interstriae with
inconspicuous fine punctures; metaventrite, only on lateral
regions, and abdominal tergites with moderately sized
punctures uniformly distributed.
Male. Head with front slightly deflected (Fig. 88); V-shaped
frontal sulcus complete; anterior angles curved and prominent;
eyes with two moderately sized punctures with long setae on
basal half of dorsal margin, basal the longest. Antennae reaching
apex of abdomen (Fig. 40); scape with slightly enlarged area
and long setae on basal half of exposed dorsal face; scape
shorter than antennomere 2 and 3 combined; antennomeres
4–10 oblong shape. Labrum with six setae medially, equal in
length; each lateral third, antero-internal seta shorter than posteroexternal seta. Mandibles with dorsal teeth weakly developed;
internal borders slightly asymmetrical, each with one small and
acute tooth at middle and, anterior to it, in right mandible with
weak area that resembles a tooth, absent in left mandible.
Maxillary palpus with palpomere 4 longer than 2 and 3
combined. Mentum ~1.5 as wide as long. Pronotum wider
than long (PW/PL = 1.54) (Fig. 88); anterior angles weakly
projected; longitudinal median sulcus conspicuous;
longitudinal sulcus of posterior angles developed and reaching
middle of pronotum; abrupt constriction at basal third and,
anterior to it, in each side with short tooth and slight
emargination; apical two-thirds with curved sides. Prosternum
with conspicuous ovate set of fine punctures in median region
near apex. Femur of anterior legs with short, hard setae on groove
of ventral margin. Elytra as long as wide (EL/BW = 1.00); stria 6
absent and stria 7 complete; striae narrower than interstriae;
epipleural carina developed. Metaventrite with slight median
depression near apex. Abdominal segments 3–6 subparallel
sides, segments 3 and 4 slightly narrower than 5–7; sternite 3
with conspicuous transverse carina; tergite 9 with short ventral
struts; tergite 10 with two pairs of long setae on apex, apical the
longest. Median lobe of aedeagus with bulbous base in ventral
view and curved apex in lateral view (Fig. 218); lateral lobes
exceeding little apex of median in lateral view; internal sclerites
as in Fig. 219.
Female. Similar to male, except for: antennae reaching apex
of elytra; scape without enlarged area and with two long setae at
middle of dorsal face; prosternum without ovate set of fine
punctures; ovipositor as in P. sulcatus; spermathecal duct
about same length of spermatheca (or ~2 times, specimens
from Nicaragua, Costa Rica and Panama), without sclerotised
basal region (Fig. 274); spermatheca somewhat elongate
(Fig. 274).
Revision of Piestus with remarks on related genera
Distribution
In the current study, the species was examined from Mexico
(Tamaulipas, Durango, San Luís Potosí, Hidalgo, Veracruz,
Oaxaca and Chiapas), Belize (Cayo), Guatemala (Zacapa and
Escuintla), Nicaragua (Granada), Costa Rica (Guanacaste,
Alajuela, Heredia, San José, Cartago and Puntarenas), Panama
(Chiriquí and Panamá) and Colombia (Boyacá). Herman (2001b)
cites this species from some of the countries listed above,
plus Honduras, Brazil and the West Indies. Navarrete-Heredia
et al. (2002) consider Brazil and West Indies as doubtful
distributions.
Biological notes
Piestus mexicanus has been found in leaf litter near streams and
under bark of decaying logs. Some species have been observed in
‘fruit fall’, ‘flower fall’ or ‘mushrooms rotting vegetation’. The
species was also collected in flight intercept traps.
Remarks
Piestus mexicanus is similar to P. sulcatus and P. gounellei, but
differs by characters cited under remarks of P. sulcatus.
Piestus ecuadorensis, sp. nov.
(Figs 41, 89, 220, 221, 275)
Type material
Holotype deposited in AMNH, male, label: (1) ‘Ecuador: Pichincha Prov.:/W
of Alluriquin, Tinalandia,/2600-2800 ft, May 19-20/1993, Lee Herman
#2724-/2725; litter nr. stream [white label, printed in black]. 6 paratypes
deposited in following collections: in AMNH: 1 female and 1 sex
undetermined, the same label of holotype; 1 female, labels: (1) ‘Ecuador:
Pichincha Prov.:/W of Alluriquin, 3.3–5.3 km/SW road to Cooperativa/
Bolivar, near Tinalandia,’ [white label, printed in black]; (2) ‘3100-3500,
V-20-93; L. Herman/#2733-2734, litter nr stream’ [white label, printed in
black]; in DZUP: 1 sex undetermined, label: (1) ‘Ecuador: Pichincha: 17/ km
SE Sto. Domingo de/Colorados, Tinalandia/3000’X-16-21-88/litter, leg.
L. Herman’ [white label, printed in black]; in SEMC: 1 female, labels: (1)
‘Ecuador: Pichincha/Tinalandia, Santo Domingo/16 km E, 750m/0160 53’S,
7930 39’W/26–27 Mar 1999, R. Brooks, D. Brzoska/ECU1B99 048 ex: flight
intercept trap’ [white label, printed in black]; (2) ‘bar code/SM0158037/
KUNHM-ENT’ [white label, printed in black]; 1 male, the first same label, (2)
‘bar code/SM0158147/KUNHM-ENT’ [white label, printed in black].
Invertebrate Systematics
537
face; scape shorter than antennomere 2 and 3 combined;
antennomeres 4–11 oblong shape. Labrum with six setae
medially, equal in length; each lateral third, antero-internal
seta shorter than postero-external seta. Mandibles with dorsal
teeth weakly developed; internal border slightly asymmetrical,
each with one small and acute tooth at middle and, anterior to it,
in right mandible with weak area that resembles a tooth, absent
in left mandible. Maxillary palpus with palpomere 4 longer
than 2 and 3 combined. Mentum ~1.5 times as wide as long.
Pronotum wider than long (PW/PL = 1.49) (Fig. 89); anterior
angles weakly projected; longitudinal median sulcus
conspicuous, broad and deep; longitudinal sulcus of posterior
angles strongly developed and reaching middle of pronotum;
abrupt constriction at basal third and, anterior to it, in each side
with short tooth and slight emargination; apical two-thirds with
curved sides. Prosternum with conspicuous ovate set of fine
punctures in median region near apex. Femur of anterior legs
with short, hard setae on groove of ventral margin. Elytra as long
as wide (EL/BW = 1.00); stria 6 absent and stria 7 complete;
striae narrower than interstriae; epipleural carina developed.
Metaventrite with slightly median depression near apex.
Abdominal segments 3–6 subparallel sides, segments 3 and 4
slightly narrower than 5–7; sternite 3 with conspicuous transverse
carina; tergite 9 with short ventral struts; tergite 10 with one pair of
long setae on apex, sometimes with one inconspicuous seta near
long setae. Median lobe of aedeagus with bulbous base in ventral
view and curved apex in lateral view (Fig. 220); lateral lobes
exceed little apex of median lobe in lateral view; internal sclerites
as in Fig. 221.
Female. Similar to male, except for: antennae exceeding
apex of elytra, but not reaching apex of abdomen; scape without
enlarged area and with two long setae at middle of the dorsal
face; prosternum without ovate set of fine punctures; ovipositor
as in P. sulcatus; spermathecal duct about same length of
spermatheca, without sclerotised basal region (Fig. 275);
spermatheca somewhat elongate (Fig. 275).
Distribution
Ecuador (Pichincha).
Biological notes
Piestus ecuadorensis has been collected in flight intercept traps
and found in leaf litter near streams.
Description
BL: 4.0–4.6 mm, BW: 1.2–1.3 mm. Body flattened
dorsoventrally; reddish dark brown (Fig. 41). Dorsal
integument of dorsum of head, except anterior angles, with
large punctures contiguous (Fig. 89); almost all pronotum with
large punctures contiguous, there are few some areas not
punctured; striae of elytra with large punctures; metaventrite,
only on lateral regions, and abdominal tergites with moderately
sized punctures uniformly distributed.
Male. Head with front slightly deflected (Fig. 89); V-shaped
frontal sulcus complete; anterior angles curved and prominent;
eyes with two moderately sized punctures with long setae on
basal half of dorsal margin, basal the longest. Antennae
reaching apex of abdomen (Fig. 41); scape with slightly
enlarged area and long setae on basal half of exposed dorsal
Remarks
Piestus mexicanus is very similar to P. foveolatus because they
share contiguous large punctures on the head and a broad and deep
longitudinal sulcus on the pronotum (Figs 89, 90). However,
P. ecuadorensis differs from P. foveolatus by the absence of a
median longitudinal sulcus plus fovea at the base on vertex
(Fig. 89), short lateral lobes exceeding slightly the apex of
median lobe (Fig. 220), and short spermathecal duct of the
same length as spermatheca (Fig. 275).
Piestus foveolatus has a median longitudinal sulcus with a
fovea at the base of vertex (Fig. 90), long lateral lobes exceeding
by one-quarter of its length the apex of median lobe (Fig. 222),
and long spermathecal duct, ~3 times the length of spermatheca
(Fig. 276).
538
Invertebrate Systematics
Piestus ecuadorensis, as well as P. foveolatus, is easily
separated from P. gounellei and P. mexicanus by contiguous
large punctures on the head and a broad and deep longitudinal
sulcus on the pronotum (Figs 89, 90) (non-contiguous punctures
on head and not broad and slightly deep longitudinal sulcus on
pronotum in P. gounellei and P. mexicanus; Figs 87, 88).
Etymology
The name is derived from Ecuador, the country where the type
material was collected.
Piestus foveolatus, sp. nov.
(Figs 42, 90, 222, 223, 276)
Type material
Holotype deposited in FMNH, male, labels: (1) ‘Panamá: Bocas d. Toro,/
Fortuna-Chiriquí Grande/road, 8470 N, 82110 W,/800m, 14–16.vii.1987’
[white labels, printed in black]; (2) ‘D.M. Olson #566,/premontane rain
forest,/sifting litter/Field Museum N.H.’ [white label, printed in black]; (3)
‘David M. Olson,/Staphylinidae: species#82’ [white label, printed in black,
except ‘82’]. 40 paratypes: in FMNH, 2 sex undetermined, the same label
of holotype (FMNH), one of them with additional label: (4) ‘Piestus/
(Trachypiestus)/det. Newton 1993’ [white label, the two first lines
manuscript, third printed in black]; in AMNH, 1 female, 10 sex
undetermined, labels: (1) ‘Panamá: Chiriquí Prov./4 km NW Volcán,
1450m/8490 19’N, 82400 31’W/XII-24–01, L. Herman/litter near stream’
[white label, printed in black]; in SEMC, 1 sex undetermined, labels: (1)
‘Costa Rica: San Jose/Cerros de Escazu/2 km, S. San Antonio, 1650m/
9530 30’N, 8490 0’W/13 Jun 1997, R.Anderson/CR1A97 013E/ex: berlese
forest litter’ [white label, printed in black]; (2) ‘bar code/SM0119958/
KUNHM-ENT’ [white label, printed in black]; 1 sex undetermined, the
same first label, (2) ‘bar code/SM0119965/KUNHM-ENT’ [white label,
printed in black]; 1 male, the same first label, (2) ‘bar code/SM0119943/
KUNHM-ENT’ [white label, printed in black]; 1 sex undetermined, the
same first label, (2) ‘bar code/SM0119960/KUNHM-ENT’ [white label,
printed in black]; 1 sex undetermined, the same first label, (2) ‘bar code/
SM0119939/KUNHM-ENT’ [white label, printed in black]; 1 sex
undetermined, the same first label, (2) ‘bar code/SM0119937/KUNHMENT’ [white label, printed in black]; 1 sex undetermined, the same first
label, (2) ‘bar code/SM0139469/KUNHM-ENT’ [white label, printed in
black]; 1 sex undetermined, the same first label, (2) ‘bar code/SM0119842/
KUNHM-ENT’ [white label, printed in black]; 1 sex undetermined, the
same first label, (2) ‘bar code/SM0119845/KUNHM-ENT’ [white label,
printed in black]; 1 sex undetermined, the same first label, (2) ‘bar code/
SM0119833/KUNHM-ENT’ [white label, printed in black]; 1 sex
undetermined, the same first label, (2) ‘bar code/SM0119836/KUNHMENT’ [white label, printed in black]; 1 sex undetermined, the same first
label, (2) ‘bar code/SM0119837/KUNHM-ENT’ [white label, printed in
black]; 1 sex undetermined, the same first label, (2) ‘bar code/SM0139464/
KUNHM-ENT’ [white label, printed in black]; 1 sex undetermined, the
same first label, (2) ‘bar code/SM0139443/KUNHM-ENT’ [white label,
printed in black]; 1 sex undetermined, the same first label, (2) ‘bar code/
SM0139444/KUNHM-ENT’ [white label, printed in black]; 1 sex
undetermined, the same first label, (2) ‘bar code/SM0139446/KUNHMENT’ [white label, printed in black]; 1 sex undetermined, the same first
label, (2) ‘bar code/SM0139448/KUNHM-ENT’ [white label, printed in
black]; 1 sex undetermined, the same first label, (2) ‘bar code/SM0139467/
KUNHM-ENT’ [white label, printed in black]; 1 sex undetermined, the
same first label, (2) ‘bar code/SM0139458/KUNHM-ENT’ [white label,
printed in black]; 1 sex undetermined, the same first label, (2) ‘bar code/
SM0139453/KUNHM-ENT’ [white label, printed in black]; 1 sex
undetermined, the same first label, (2) ‘bar code/SM0139452/KUNHM-
E. Caron et al.
ENT’ [white label, printed in black]; 1 sex undetermined, the same first
label, (2) ‘bar code/SM0139451/KUNHM-ENT’ [white label, printed in
black]; 1 sex undetermined, the same first label, (2) ‘bar code/SM0139450/
KUNHM-ENT’ [white label, printed in black]; 1 sex undetermined, the
same first label, (2) ‘bar code/SM0139468/KUNHM-ENT’ [white label,
printed in black]; 1 sex undetermined, labels, (1) ‘Costa Rica: San Jose/
Cerros de Escazu/2 km, S. San Antonio, 1650m/9530 30’N, 8490 0’W/13
Jun 1997, R.Anderson/CR1A97 013H/ex: berlese forest litter’ [white label,
printed in black]; (2) ‘bar code/SM0136291/KUNHM-ENT’ [white label,
printed in black]; in DZUP, 1 sex undetermined, labels: (1) ‘Costa Rica:
San Jose/Cerros de Escazu/2 km, S. San Antonio, 1650m/9530 30’N,
8490 0’W/13 Jun 1997, R.Anderson/CR1A97 013E/ex: berlese forest
litter’ [white label, printed in black], (2) ‘bar code/SM0119926/
KUNHM-ENT’ [white label, printed in black]; 1 sex undetermined, the
same first label, (2) ‘bar code/SM0119931/KUNHM-ENT’ [white label,
printed in black].
Additional material
See Appendix 2.
Description
BL: 3.4–5.3 mm, BW: 1.1–1.4 mm. Body flattened
dorsoventrally convex; reddish dark brown (Fig. 42). Dorsal
integument of head, except anterior angles, with large
punctures contiguous (Fig. 90); almost all pronotum with large
punctures contiguous, there are few some areas not punctured;
striae of elytra with large punctures; metaventrite, only on lateral
regions, and abdominal tergites with moderately sized punctures
uniformly distributed.
Male. Head with front slightly deflected (Fig. 90); V-shaped
frontal sulcus complete; anterior angles curved and prominent,
and median longitudinal sulcus with fovea at base; eyes with
two moderately sized punctures with long setae on basal half of
dorsal margin, basal the longest. Antennae reaching apex of
abdomen (Fig. 42); scape with slightly enlarged area and long
setae on basal half of exposed dorsal face; scape shorter than
antennomere 2 and 3 combined; antennomeres 4–11 oblong
shape. Labrum with six setae medially, equal in length; each
lateral third, antero-internal seta shorter than postero-external
seta. Mandibles with dorsal teeth weakly developed; internal
border slightly asymmetrical, each with one small and acute
tooth at middle and, anterior to it, in right mandible with a
very weak area that resembles a tooth, absent in left mandible.
Maxillary palpus with palpomere 4 longer than 2 and 3 combined.
Mentum ~1.5 times as wide as long. Pronotum wider than long
(PW/PL = 1.50) (Fig. 90); anterior angles not projected;
longitudinal median sulcus conspicuous, broad and deep;
longitudinal sulcus of posterior angles strongly developed and
reaching middle of pronotum; abrupt constriction at basal third
and, anterior to it, in each side with short tooth and slight
emargination; apical two-thirds with curved sides. Prosternum
with conspicuous ovate set of fine punctures in median region
near apex. Femur of anterior legs with short, hard setae on groove
of ventral margin. Elytra as long as wide (EL/BW = 1.04); stria 6
absent and stria 7 complete; striae narrower than interstriae;
interstriae not carinate; epipleural carina developed.
Metaventrite with conspicuous median depression near apex.
Abdominal segments 3–6 subparallel sides, segments 3 and 4
slightly narrower than 5–7; sternite 3 with conspicuous
transverse carina; tergite 9 with short ventral struts; tergite 10
Revision of Piestus with remarks on related genera
with two pairs of long setae on apex, apical the longest. Median
lobe of aedeagus with bulbous base in ventral view and curved
apex in lateral view (Fig. 222); lateral lobes exceeding apex of
median lobe by one-quarter of its length in lateral view; internal
sclerites as in Fig. 223.
Female. Similar to male, except for: antennae exceeding
apex of elytra, but not reaching apex of abdomen; scape without
enlarged area and with two long setae at middle of dorsal face;
prosternum without ovate set of fine punctures; ovipositor as in
P. sulcatus; spermathecal duct long, ~3 times length of
spermatheca, without sclerotised basal region (Fig. 276);
spermatheca as in Fig. 276.
Distribution
Costa Rica (Alajuela, Heredia, San José, Cartago and
Puntarenas), Panama (Bocas del Toro and Chiriquí) and
Colombia (Valle del Cauca).
Biological notes
Piestus foveolatus has been found in leaf litter, sometimes near
streams (‘litter near stream’). This species was also collected in
flight intercept traps and by Berlese extraction of leaf litter.
Remarks
Piestus foveolatus is very similar to P. ecuadorensis, and also to
P. gounellei and P. mexicanus, but differs by characters cited
under remarks of P. ecuadorensis.
Etymology
The specific name refers to the conspicuous fovea on the median
base of head.
Piestus sulcipennis Scheerpeltz, 1952
(Figs 43, 91, 224, 225, 277)
Piestus sulcipennis Scheerpeltz, 1952: 299 (original description, type
locality: ‘St. Catharina-Brasilien’); Herman, 2001b: 1795
(distribution).
Type material
Piestus sulcipennis Scheerpeltz, 1952. Lectotype (here designated)
deposited in NMW, male [damaged, left antenna incomplete], with labels:
(1) ‘St.-Catharina/Brasilien’ [white label, printed in black]; (2) ‘ex coll./
Scheerpeltz’ [blue label, printed in black]; (3) ‘Typus/Piestus (Trachy-/
piestus) sulci-/pennis/O. Scheerpeltz [printed in black]’ [red label, first and
fifth lines printed in black, second, third and fourth lines handwritten]; (4)
‘sulcipennis/Scheerp.’ [green label, handwritten]. Paralectotype deposited in
NMW, sex undetermined, with the same labels of lectotype. Note: In the
original description Scheerpeltz (1952) specified a male and female observed.
However, only one specimen was dissected and its sex confirmed (male)
(lectotype). The sex of the other specimen (paralectotype) is still
undetermined.
Invertebrate Systematics
539
Dorsal integument of head, except anterior angles, and almost all
pronotum with large punctures contiguous, there are some few
areas unpunctured (Fig. 91); striae of elytra with large punctures;
abdominal tergites with moderately sized punctures uniformly
distributed.
Male. Head with front slightly deflected (Fig. 91); V-shaped
frontal sulcus complete; anterior angles curved and prominent;
eyes with two moderately sized punctures with long setae on
basal half of dorsal margin, basal the longest. Antennae
exceeding apex of elytra, but not reaching apex of abdomen;
scape with two long setae at middle of dorsal face; scape
subequal to antennomere 2 and 3 combined; antennomeres
4–11 oblong shape (Fig. 43). Labrum with six setae medially,
equal in length; each lateral third, antero-internal seta shorter than
postero-external seta. Mandibles with dorsal teeth weakly
developed; internal borders slightly asymmetrical, each with
one small and acute tooth at middle and, anterior to it, in right
mandible with weak area that resembles a tooth, absent in the
left mandible. Maxillary palpus with palpomere 4 longer than 2
and 3 combined. Mentum ~1.5 times as wide as long. Pronotum
wider than long (PW/PL = 1.47) (Fig. 91); anterior angles weakly
projected; the apical half with slightly somewhat longitudinal
median depression and on basal half a transverse depression;
longitudinal sulcus inconspicuous; longitudinal sulcus of
posterior angles developed and reaching middle or pronotum;
abrupt constriction at basal third and, anterior to it, in each side
with short tooth and slight emargination; apical two-thirds with
sinuous sides. Prosternum with conspicuous ovate set of fine
punctures in median region near apex. Femur of anterior legs
with short, hard setae on groove of ventral margin. Elytra as
long as wide (EL/BW = 0.94); stria 6 at basal half and stria 7
complete; striae narrower than interstriae; interstriae not carinate;
epipleural carina developed. Abdominal segments 3–6
subparallel sides, segments 3 and 4 slightly narrower than 5–7;
sternite 3 with conspicuous transverse carina; tergite 9 with long
ventral struts; tergite 10 with two pairs of long setae on apex,
apical the longest. Median lobe of aedeagus with bulbous base in
ventral view and curved apex in lateral view (Fig. 224); lateral
lobes exceed a little apex of median lobe in lateral view; internal
sclerites as in Fig. 225.
Female. Similar to male, except for: prosternum without
ovate set of fine punctures; ovipositor as in P. sulcatus;
spermathecal duct about same length of spermatheca, with
sclerotised basal region (Fig. 277); spermatheca with globose
apex and accessory gland on base (Fig. 277).
Distribution
Brazil (Santa Catarina).
Biological notes
Unknown.
Additional material
Remarks
See Appendix 2.
Piestus sulcipennis is similar to P. aper, differing from that
species by complete V-shaped frontal sulcus (Fig. 91), two
long setae on dorsal margin of the eyes, two long setae on
scape, slightly asymmetrical internal borders of mandibles
with one small acute tooth, stria 6 developed at the basal half
Redescription
BL: 3.8–4.0 mm, BW: 1.1–1.1 mm (types). Body flattened
dorsoventrally; reddish brown to reddish dark brown (Fig. 43).
540
Invertebrate Systematics
E. Caron et al.
Figs 92–105. Head and pronotum, dorsal view. 92, Piestus imperfectus, sp. nov.; 93, P. costatus; 94, P. chiriquensis; 95, P. nevermanni; 96, P. paradoxus;
97, P. boliviensis, sp. nov.; 98, P. aper; 99, P. angularis; 100, P. acuminatus, sp. nov.; 101, P. rugosus; 102, P. elegans, sp. nov.; 103, Eleusis interrupta, comb.
rest.; 104, Hypotelus laevis, comb. nov.; 105, Hypotelus andinus, comb. nov. Scale bar = 0.50 mm.
Revision of Piestus with remarks on related genera
of elytra, subparallel sides of abdomen, segments 3 and 4 slightly
narrower than 5–7, and accessory gland at the base of spermatheca
(Fig. 277).
Piestus aper has an incomplete V-shaped frontal sulcus
(Fig. 98), one short seta on dorsal margin of the eyes, one long
seta on scape, smooth internal borders of mandibles, stria 6 absent
on elytra, abdominal segment 5 wider than others, and accessory
gland at apical third of spermatheca (Fig. 283).
Piestus imperfectus, sp. nov.
(Figs 44, 92, 226, 227, 278)
Type material
Holotype deposited in SEMC, male, with labels: (1) ‘Ecuador: Pichincha/
Nanegalito, 7 km S, 1500m/000 23’N, 78400 36’W/28 Oct 1999,
R. Anderson/ECU1A99 212B/ex: riparian mont. evergreen litter’ [white
label, printed in black]; (2) ‘bar code/SM0359240/KUNHM-ENT’ [white
label, printed in black]. 36 paratypes deposited in following collections: in
SEMC: 1 female, labels: (1) ‘Ecuador: Esmeraldas/Bilsa, 0200 0’S,
79430 0’W/28 Apr-10 May 1996/ECU1H96 015, P. Hibbs/ex: flight
intercept trap’ [white label, printed in black]; (2) ‘bar code/SM0088817/
KUNHM-ENT’ [white label, printed in black]; 1 sex undetermined, labels:
(1) ‘Ecuador: Pichincha/Maquipucuna For. Res./50 km NW Quito, 1660m/
21 Dec. 1991, C. Carlton/R. Leschen #22’ [white label, printed in black]; (2)
‘Piestus/(Tachypiestus)/sp./det. J.S. Ashe 19’ [white label, the first three
lines handwritten, fourth line printed in black]; 3) ‘near P./(Tachypiestus)/
chiriquensis Shp./det. J.S. Ashe 1992’ [white label, first three lines
handwritten, fourth printed in black]; 2 sex undetermined, with the same
first label: (1) ‘Ecuador: Pichincha/Maquipucuna For. Res./50 km NW
Quito, 1660m/21 Dec. 1991, C. Carlton/R. Leschen #22’ [white label,
printed in black]; 1 sex undetermined, labels: (1) ‘Ecuador: Pichincha/45 km
NNW Quito/Macquipucuna Station/1600-1650m/3-18 Apr 1996/ECU1H96
012, P. Hibbs/ex: flight intercept trap’ [white label, printed in black]; (2)
‘bar code/SM0090869/KUNHM-ENT’ [white label, printed in black]; 1 sex
undetermined, the same first label, (2) ‘bar code/SM0090885/KUNHMENT’ [white label, printed in black]; 1 sex undetermined, labels: (1)
‘Ecuador: Pichincha/45 km NNW Quito/Macquipucuna Station/16001650m/18 Apr-5 May 1996/ECU1H96 013, P. Hibbs/ex: flight intercept
trap’ [white label, printed in black]; (2) ‘bar code/SM0090741/KUNHMENT’ [white label, printed in black]; 1 sex undetermined, labels: (1)
‘Ecuador: Pichincha/Mindo, 10.6 km W, Mindo Road/040 23’S,
78450 14’W, 1460m/26–29 Mar 1999, R. Brooks, D. Brzoska/ECU1B99
056 ex: flight intercept trap’ [white label, printed in black]; (2) ‘bar code/
SM0157957/KUNHM-ENT’ [white label, printed in black]; 1 female,
labels: (1) ‘Ecuador: Pichincha/Mindo, 10.6 km W, Mindo Road/
040 23’S, 78450 14’W, 1445m/26–29 Mar 1999, R. Brooks, D. Brzoska/
ECU1B99 064 ex: flight intercept trap’ [white label, printed in black]; (2)
‘bar code/SM0157373/KUNHM-ENT’ [white label, printed in black]; 1
female, labels: (1) ‘Ecuador: Pichincha/Maquipucuna Biological Station/
070 0’N, 78380 6’W 1200m/27 Oct 1999, R. Anderson/ECU1A99 208B/ex:
montane evergreen forest litter’ [white label, printed in black]; (2) ‘bar code/
SM0362386/KUNHM-ENT’ [white label, printed in black]; 1 sex
undetermined, labels: 1) ‘Ecuador: Pichincha/Maquipucuna Biological
Station/060 25’N, 78370 18’W 1480m/27 Oct 1999, R. Anderson/
ECU1A99 209A/ex: montane evergreen forest litter’ [white label, printed
in black]; (2) ‘bar code/SM0363280/KUNHM-ENT’; ‘Piestus/spp./det.
J.S. Ashe 2000’ [white label, printed in black]; 1 sex undetermined,
labels: (1) ‘Ecuador: Pichincha/Maquipucuna Biological Station/River
Trail, 1200m/070 34’N, 78370 57’W/27 Oct 1999, Z.H. Falin/ECU1F99
025/ex: under bark’ [white label, printed in black]; (2) ‘bar code/
SM0350256/KUNHM-ENT’ [white label, printed in black]; 1 sex
undetermined, labels: (1) ‘Ecuador: Pichincha/Maquipucuna Biological
Invertebrate Systematics
541
Station/070 0’N, 78380 6’W 1200m/27 Oct 1999, R. Anderson/ECU1A99
208C/ex: montane evergreen forest litter’ [white label, printed in black]; (2)
‘bar code/SM0363664/KUNHM-ENT’ [white label, printed in black]; 1 sex
undetermined, the same first label, (2) ‘bar code/SM0363666/KUNHMENT’ [white label, printed in black]; 1 sex undetermined, labels: (1)
‘Ecuador: Pichincha/Maquipucuna Biological Station 1275m/070 22’N,
79390 0’W/27–29 Oct 1999, Z.H. Falin/ECU1F99 048/ex: flight intercept
trap’ [white label, printed in black]; (2) ‘bar code/SM0355819/KUNHMENT’ [white label, printed in black]; 1 sex undetermined, the same first
label, (2) ‘bar code/SM0355818/KUNHM-ENT’ [white label, printed in
black]; 1 sex undetermined, the same first label, (2) ‘bar code/SM0355820/
KUNHM-ENT’ [white label, printed in black]; 1 sex undetermined, the
same first label, (2) ‘bar code/SM0355821/KUNHM-ENT’ [white label,
printed in black]; 1 sex undetermined, the same first label, (2) ‘bar code/
SM0355839/KUNHM-ENT’ [white label, printed in black]; 1 female,
labels: (1) ‘Ecuador: Pichincha/Bellavista Reserve, Ridge Trail/12 km S
Nanegalito, 2250m/000 54’S, 78400 56’W/28 Oct 1999, R. Anderson/
ECU1A99 211H/ex: cloud forest litter’ [white label, printed in black];
(2) ‘bar code/SM0357088/KUNHM-ENT’ [white label, printed in black]; 1
sex undetermined, labels: (1) ‘Ecuador: Napo, 2300m/Sierra Azul, Hacienda
Arargon/0400 0’S, 77550 0’W/17 Feb-26 Mar 1996/ECU1H96 009,
P. Hibbs/ex: flight intercept trap’ [white label, printed in black]; (2) ‘bar
code/SM0087583’ [white label, printed in black]; 1 sex undetermined, the
same first label, (2) ‘bar code/SM0087727’ [white label, printed in black]; 1
female, labels: (1) ‘Ecuador: Napo, Cosanga/4.2 km S on Baeza-Tena/Road
then 2.9 km W on pipeline/access road, 2150m/0370 19’S, 77500 1’W/6
Nov 1999, Z.H. Falin/ECU1F99 113 ex: under bark’ [white label, printed in
black]; (2) ‘bar code/SM0352655/KUNHM-ENT’ [white label, printed in
black]; in AMNH: 3 sex undetermined, label: (1) ‘Ecuador: Pichincha:/
Palmeras, old Sto. Dgo./rd, km 59, 43 km NE/Alluriquin, X-15-88/6400’,
light, L. Herman’ [white label, printed in black]; 1 female, 2 sex
undetermined, label: (1) ‘Ecuador: Pichincha:/18–20 km NE Alluriquin/
old Quito-Sto. Domingo rd/X-21-88 4,700–4,900’/litter, L. Herman’ [white
label, printed in black]; 1 female, 3 sex undetermined, labels: (1) ‘Ecuador:
Pichincha Prov./W of Alluriquin, 3.3–5.3 km/SW road to Cooperativa/
Bolivar, near Tinalandia,’ [white label, printed in black]; (2) ‘3100-3500,
V-20-93; L. Herman/#2733-2734, litter nr stream’ [white label, printed in
black]; in DZUP, 1 male, 1 sex undetermined, label: 1) ‘Ecuador: Pichincha/
Maquipucuna For. Res./50 km NW Quito, 1660m/21 Dec. 1991, C. Carlton/
R. Leschen #22’ [white label, printed in black]; in FMNH, 2 sex
undetermined, label: (1) ‘Ecuador: Pichincha/Maquipucuna For. Res./
50 km NW Quito, 1660m/21 Dec. 1991, C. Carlton/R. Leschen #22’
[white label, printed in black].
Description
BL: 4.5–5.8 mm, BW: 1.4–1.6 mm. Body somewhat convex;
reddish brown to reddish dark brown (Fig. 44). Dorsal
integument of head, except anterior angles, and all pronotum
with large punctures contiguous (Fig. 89); striae of elytra with
large punctures; metaventrite, only on lateral regions, and
abdominal tergites with moderately sized punctures uniformly
distributed.
Male. Head with front slightly deflected (Fig. 92); V-shaped
frontal sulcus incomplete, curved arms not joined medially;
anterior angles curved and prominent; eyes with a moderately
sized puncture with long seta on basal half of dorsal margin.
Antennae reaching apex of abdomen; scape with slightly
enlarged area and long setae on basal half of exposed dorsal
face; scape shorter than antennomere 2 and 3 combined;
antennomeres 4–11 oblong shape (Fig. 44). Labrum with six
setae medially, equal in length; each lateral third, antero-internal
seta shorter than postero-external seta. Mandibles with ventral
542
Invertebrate Systematics
teeth curved, not projected; dorsal teeth weakly developed;
internal borders symmetrical, each with two acute teeth, basal
anteriorly projected. Maxillary palpus with palpomere 4 longer
than 2 and 3 combined. Mentum subquadrate, as wide as long.
Pronotum wider than long (PW/PL = 1.45) (Fig. 92); anterior
angles weakly projected; on apical half with slightly somewhat
longitudinal median depression and basal half a transverse
depression; longitudinal median sulcus inconspicuous;
longitudinal sulcus of posterior angles developed and reaching
middle of pronotum; abrupt constriction at basal third and,
anterior to it, in each side with short tooth and slight
emargination; apical two-thirds with slightly sinuous sides.
Prosternum with conspicuous ovate set of fine punctures in
median region near apex. Femur of anterior legs with short,
hard setae on groove of ventral margin. Elytra somewhat
longer than wide (EL/BW = 1.07); stria 6 absent and stria 7
complete; striae wider than interstriae; interstriae not carinate;
epipleural carina developed. Metaventrite with conspicuous
median depression near apex. Abdominal segments 3–6
subparallel sides; sternite 3 with conspicuous transverse
carina; tergite 9 with short ventral struts; tergite 10 with pair
of long setae on apex. Median lobe of aedeagus with bulbous base
in ventral view and slightly curved apex in lateral view, tube
longer than base (Fig. 226); lateral lobes reaching apex of median
lobe in lateral view; internal sclerites as in Fig. 227.
Female. Similar to male, except for: antennae exceeding
apex of elytra, but not reaching apex of abdomen; scape without
enlarged area and with one long setae at middle of dorsal face;
prosternum without ovate set of fine punctures; ovipositor as in
P. sulcatus; spermathecal duct long, ~3 times length of
spermatheca, without sclerotised basal region (Fig. 278);
spermatheca L-shaped (Fig. 278).
Distribution
Ecuador (Esmeralda, Pichincha and Napo).
Biological notes
This species has been found in litter, under bark and also
collected in flight intercept traps.
Remarks
Piestus imperfectus is similar to P. chiriquensis, but differs
mainly by having an incomplete V-shaped frontal sulcus
(Fig. 92), curved apex of median lobe of aedeagus and tube
longer than base (Fig. 226), long spermathecal duct, ~3 times
the length of spermatheca and an L-shaped spermatheca
(Fig. 278).
Piestus chiriquensis has a complete V-shaped frontal
sulcus (Fig. 94), slightly curved apex of median lobe and short
tube with the same length as base (Fig. 230), short spermathecal
duct, about the same length as spermatheca, and sinuous shaped
spermatheca (Fig. 280).
Piestus imperfectus, as well as P. chiriquensis, may be easily
separated from P. nevermanni and P. costatus by slightly
enlarged area with long setae on scape (somewhat as a tuft)
and two acute teeth on symmetrical internal borders of mandibles
(only a tuft of long setae on scape of male and only one acute tooth
E. Caron et al.
on internal borders of mandibles in P. nevermanni and
P. costatus).
Etymology
The name refers to the incomplete V-shaped frontal sulcus on the
median of head.
Piestus costatus Sharp, 1887
(Figs 45, 93, 228, 229, 279)
Piestus costatus Sharp, 1887: 715 (original description, type locality:
‘Panama, Volcan de Chiriqui 2000 to 3000 feet’); Blackwelder, 1944:
100 (distribution); Herman, 2001b: 1790.
Piestus (Piestus) costatus: Bernhauer & Schubert, 1910: 7 (distribution).
Piestus (Trachypiestus) costatus: Scheerpeltz, 1952: 294 (characters,
distribution).
Type material
Holotype deposited in BMNH, female, with labels: (1) ‘Piestus costatus/
Type D.S./V. de Chiriqui/2-3000ft. Champion’ [white labels, together the
specimen, handwritten]; (2) ‘Holo-/Type’ [circle white label with border red,
printed in black]; (3) ‘B.C.A. Col. I. 2./Piestus/costatus,/Sharp.’ [white label,
printed in black]; (4) ‘Sharp Coll./1905-313.’ [white label, printed in black];
(5) ‘Holotype/Piestus/costatus/Sharp, 1887/det. R.G. Booth 2008’ [white
label, the four first lines handwritten, the last one printed in black]. Note: In the
original description Sharp (1887) specified only one specimen (female)
observed.
Additional material
See Appendix 2.
Redescription
BL: 3.8–5.2 mm, BW: 1.1–1.4 mm. Body somewhat convex;
reddish brown to reddish dark brown (Fig. 45). Dorsal
integument of head, except anterior angles, and all pronotum
with large punctures contiguous (Fig. 93); striae of elytra with
inconspicuous inclinate microstriae and large punctures;
metaventrite, only on lateral regions, and abdominal tergites
with moderately sized punctures uniformly distributed.
Male. Head with front slightly deflected (Fig. 93); V-shaped
frontal sulcus complete, sometimes inconspicuous at middle;
anterior angles curved and prominent; eyes with a moderately
sized puncture with long seta on basal half of dorsal margin.
Antennae almost reaching apex of abdomen (Fig. 45); scape with
a few long setae on basal half of exposed dorsal face (somewhat
as a tuft); scape shorter than antennomere 2 and 3 combined;
antennomeres 4–11 oblong shape. Labrum with six setae
medially, equal in length; each lateral third, antero-internal
seta shorter than postero-external seta. Mandibles with dorsal
teeth weakly developed; internal border slightly asymmetrical,
each with one small and acute tooth at middle and, anterior to it,
in right mandible with weak area that resembles a tooth, absent
in the left mandible. Maxillary palpus with palpomere 4 longer
than 2 and 3 combined. Mentum subquadrate, as long as wide.
Pronotum wider than long (PW/PL = 1.28) (Fig. 93); anterior
angles not projected; apical half with somewhat longitudinal
median depression and on basal half a transverse depression;
longitudinal median sulcus inconspicuous; longitudinal sulcus
of posterior angles developed and reaching middle of pronotum;
Revision of Piestus with remarks on related genera
abrupt constriction at basal third and, anterior to it, in each side
with short tooth and slight emargination; apical two-thirds with
slightly sinuous sides. Prosternum with conspicuous ovate set
of fine punctures in median region near apex. Femur of anterior
legs with short, hard setae on groove of ventral margin. Elytra
somewhat longer than wide (EL/BW = 1.05), sometimes
occulting tergite 3; stria 6 absent and stria 7 complete; striae
wider than interstriae; interstriae carinate; epipleural carina
developed. Metaventrite with conspicuous median depression
near apex. Abdominal segments 3–6 subparallel sides; sternite 3
with conspicuous transverse carina; tergite 9 with short ventral
struts; tergite 10 with pair of long setae on apex. Median lobe of
aedeagus with bulbous base in ventral view and slightly curved
apex in lateral view, tube longer than base (Fig. 228); lateral
lobes not reaching apex of median lobe in lateral view; internal
sclerites as in Fig. 229.
Female. Similar to male, except for: antennae reaching
apex of elytra; scape without enlarged area and with one long
setae at middle of dorsal face; prosternum without ovate set of
fine punctures; ovipositor as in P. sulcatus; spermathecal duct
long, ~5 times length of spermatheca, without sclerotised basal
region (Fig. 279); spermatheca L-shaped (Fig. 279).
Distribution
Costa Rica (San José, Cartago and Puntarenas) and Panama
(Bocas del Toro and Chiriquí).
Biological notes
Piestus costatus has been found in litter, near streams or
associated with decaying logs. Some specimens were collected
in flight intercept traps and by Berlese extraction of leaf litter.
Remarks
Piestus costatus is very similar to P. nevermanni but differs by
having a long spermathecal duct, ~5 times the length of
spermatheca, without sclerotised basal region, and
spermatheca L-shaped (Fig. 279) (short spermathecal duct,
about the same length as spermatheca, and spiral-shaped
spermatheca, in P. nevermanni; Fig. 281).
Piestus costatus is also similar to P. imperfectus and
P. chiriquensis, from which it differs by characters cited above
(see remarks under P. imperfectus).
Piestus chiriquensis Sharp, 1887
(Figs 46, 94, 230, 231, 280)
Piestus chiriquensis Sharp, 1887: 715 (original description, type locality:
‘Panama, Volcan de Chiriqui 2000 to 4000 feet’); Blackwelder, 1944:
100 (distribution); Herman, 2001b: 1790 (distribution).
Piestus (Piestus) chiriquensis: Bernhauer & Scheerpeltz, 1910: 7
(distribution).
Piestus (Trachypiestus) chiriquensis: Scheerpeltz, 1952: 294 (characters,
distribution).
Type material
Syntype deposited in FMNH, sex undetermined, labels: (1) ‘Piestus
chiriquen/sis D.S./V. de Chiriqui 2500/4000ftt. Champion’ [white label,
together with the specimen, handwritten]; (2) ‘V. de Chiriquí,/2-3000 ft/
Champion’ [white label, printed in black]; (3) ‘B.C.A. Col. I. 2./Piestus/
Invertebrate Systematics
543
chiriquensis/Sharp.’ [white label, printed in black]; (4) ‘Chicago Nat. Hist.
Mus./(ex. D. Sharp Colln./by exchange with/Brit. Mus. Nat. Hist.)’ [white
label, printed in black]. Note: There are 4 syntypes (not examined) deposited
in BMNH (Dr Roger Booth, BMNH, pers. comm.).
Additional material
See Appendix 2.
Redescription
BL: 4.2–5.8 mm, BW: 1.2–1.6 mm. Body somewhat convex;
light brown to reddish dark brown (Fig. 46). Dorsal
integument of head, except anterior angles, and all pronotum
with large punctures contiguous (Fig. 94); striae of elytra with
large punctures; metaventrite, only on lateral regions, and
abdominal tergites with moderately sized punctures uniformly
distributed.
Male. Head with front slightly deflected (Fig. 94); V-shaped
frontal sulcus complete; anterior angles curved and prominent;
eyes with a moderately sized puncture with long seta on basal
half of dorsal margin. Antennae reaching apex of abdomen;
scape with slightly enlarged area and long setae on basal half
of exposed dorsal face; scape shorter than antennomere 2 and 3
combined; antennomeres 4–11 oblong shape (Fig. 46). Labrum
with six setae medially, equal in length; each lateral third,
antero-internal seta shorter than postero-external seta.
Mandibles with dorsal teeth weakly developed; internal border
symmetrical, each with two acute teeth, basal anteriorly
projected. Maxillary palpus with palpomere 4 longer than 2
and 3 combined. Mentum subquadrate, as wide as long.
Pronotum wider than long (PW/PL = 1.45) (Fig. 94); anterior
angles weakly projected; on apical half with somewhat
longitudinal median depression and on basal half a transverse
depression; longitudinal median sulcus inconspicuous;
longitudinal sulcus of posterior angles developed and reaching
middle of pronotum; abrupt constriction at basal third and,
anterior to it, in each side with short tooth and slight
emargination; apical two-thirds with slightly sinuous sides.
Prosternum with conspicuous ovate set of fine punctures in
median region near apex. Femur of anterior legs with short,
hard setae on groove of ventral margin. Elytra somewhat
longer than wide (EL/BW = 1.04), occulting tergite 3; stria 6
absent and stria 7 complete; striae wider than interstriae;
interstriae carinate; epipleural carina developed. Metaventrite
with conspicuous median depression near apex. Abdominal
segments 3–6 with subparallel sides; sternite 3 with
conspicuous transverse carina; tergite 9 with short ventral
struts; tergite 10 with two pairs of long setae on apex, apical
the longest. Median lobe of aedeagus with bulbous base in ventral
view and slightly curved apex in lateral view, tube short with
same length of base (Fig. 230); lateral lobes reaching apex of
median lobe in lateral view; internal sclerites as in Fig. 231.
Female. Similar to male, except for: antennae exceeding
apex of elytra, but not reaching apex of abdomen; scape without
enlarged area and with one long seta at middle of dorsal face;
prosternum without ovate set of fine punctures; ovipositor as in
P. sulcatus; spermathecal duct short, about same length of
spermatheca, without sclerotised basal region (Fig. 280);
spermatheca with sinuous shape (Fig. 280).
544
Invertebrate Systematics
Distribution
In the current study the species was examined from Nicaragua
(Río San Juan), Costa Rica (Guanacaste, Alajuela, Heredia, San
José, Cartago, Limón and Puntarenas), Panama (Bocas del Toro,
Chiriquí, Panamá and Darién), Colombia (Cundinamarca and
Nariño), Venezuela (Lara), Ecuador (Esmeraldas, Sucumbios,
Pichincha, Napo, Cotopaxi and Tungurahua), Peru (Huánuco,
Cusco) and Bolivia (Cochabamba).
Remarks
Piestus chiriquensis is very similar to P. imperfectus and also to
P. costatus and P. nevermanni, from which it is easily separated
by characters cited above (see remarks under P. imperfectus).
Biological notes
Piestus chiriquensis has been found in litter, near streams or
associated with decaying logs. Some specimens were collected
in flight intercept and Malaise traps and by Berlese extraction
of leaf litter.
Piestus nevermanni Scheerpeltz, 1952
(Figs 47, 95, 232, 233, 281)
Piestus (Trachypiestus) nevermanni Scheerpeltz, 1952: 294, 302
(original description, type locality: ‘Costarica’).
Piestus nevermanni: Herman, 2001b: 1792 (distribution).
Type material
Holotype deposited in NMW, female [damaged specimen: without right
anterior and median legs], with labels: (1) ‘<’ [white label, printed in
black]; (2) ‘Costarica/leg. nevermann’ [white label, handwritten]; (3)
‘Dr M Bernhauer/8.11. donavit 1933’ [white label, first line printed in
black, second line handwritten, except the word ‘donavit’ printed in
black]; (4) ‘ex coll./Scheerpeltz’ [blue label, printed in black]; (5)
‘TYPUS/ Piestus (Trachy-/piestus) costa-/ricensis/O. Scheerpeltz’ [red
label, first line and fifth printed in black, the others handwritten]; (6)
‘nevermanni’ [light green label, handwritten]. Note: In the original
description Scheerpeltz (1952) specified one male specimen observed.
However, the observed type from NMW is female.
E. Caron et al.
third, antero-internal seta shorter than postero-external seta.
Mandibles with dorsal teeth weakly developed; internal border
slightly asymmetrical, each with one small and acute tooth at
middle and, anterior to it, in the right mandible with weak area
that resembles a tooth, absent in the left mandible. Maxillary
palpus with palpomere 4 longer than 2 and 3 combined. Mentum
subquadrate, as long as wide. Pronotum wider than long
(PW/PL = 1.48) (Fig. 95); anterior angles weakly projected; on
apical half with a somewhat longitudinal median depression
and on basal half a transverse depression; longitudinal median
sulcus inconspicuous; longitudinal sulcus of posterior angles
developed and reaching middle of pronotum; abrupt
constriction at basal third and, anterior to it, in each side with
a short tooth and slight emargination; apical two-thirds with
slightly sinuous sides. Prosternum with conspicuous ovate set
of fine punctures in median region near apex. Femur of anterior
legs with short, hard setae on groove of ventral margin. Elytra
somewhat longer than wide (EL/BW = 1.06); stria 6 absent and
stria 7 complete; striae wider than interstriae; interstriae carinate;
epipleural carina developed. Metaventrite with conspicuous
median depression near apex. Abdominal segments 3–6
subparallel sides; sternite 3 with conspicuous transverse
carina; tergite 9 with long ventral struts; tergite 10 with two
pairs of long setae on apex, apical the longest. Median lobe of
aedeagus with bulbous base in ventral view and curved apex in
lateral view, tube longer than base (Fig. 232); lateral lobes
reaching apex of median lobe in lateral view; internal sclerites
as in Fig. 233.
Female. Similar to male, except for: antennae exceeding
apex of elytra, but not reaching apex of abdomen; scape without
enlarged area and with one long seta at middle of dorsal face;
prosternum without ovate set of fine punctures; ovipositor as in
P. sulcatus; spermathecal duct short, about same length of
spermatheca, without sclerotised basal region (Fig. 281);
spermatheca with spiral shape (Fig. 281).
Distribution
Costa Rica and Panama (Chiriqui).
Additional material
Biological notes
See Appendix 2.
This species has been collected in flight intercept traps.
Redescription
BL: 6.0 mm, BW: 1.7 mm. Body somewhat convex; brown to
reddish dark brown (Fig. 47). Dorsal integument of head,
except anterior angles, and all pronotum with large punctures
contiguous (Fig. 95); striae of elytra with large punctures;
metaventrite, only on lateral regions, and abdominal tergites
with moderately sized punctures uniformly distributed.
Male. Head with front slightly deflected (Fig. 95); V-shaped
frontal sulcus complete; anterior angles curved and prominent;
eyes with a moderately sized puncture with long seta on basal
half of dorsal margin. Antennae reaching apex of abdomen;
scape with one tuft of long setae on basal half of exposed
dorsal face; scape shorter than the antennomere 2 and 3
combined; antennomeres 4–11 oblong shape (Fig. 47).
Labrum with six setae medially, equal in length; each lateral
Remarks
Piestus nevermanni is very similar to P. imperfectus and also to
P. costatus and P. chiriquensis, differing from these species by
characters cited above (see remarks under P. imperfectus).
Piestus paradoxus Bernhauer, 1917
(Figs 48, 49, 96, 127, 234, 235, 282)
Piestus paradoxus Bernhauer, 1917: 45 (original description, type
locality: ‘Caracas’); Blackwelder, 1944: 100 (distribution); Herman,
2001b: 1793 (distribution).
Piestus (Piestus) paradoxus: Scheerpeltz, 1952: 994 (distribution).
Piestus (Elytropiestus) paradoxus: Scheerpeltz, 1952: 294 (characters,
distribution).
Revision of Piestus with remarks on related genera
Type material
Holotype deposited in FMNH, female [damaged specimen: without left
antennomeres 3–11], with labels: (1) ‘Caracas/Bang. Haas.’ [white label,
handwritten]; (2) ‘paradoxus/Bernh./Typus unic.’ [light yellow label,
handwritten]; (3) ‘Chicago NHMus/M.Bernhauer/Collection’ [white label,
printed in black]. Note: In the original description Bernhauer (1917) specified
only one specimen observed.
Additional material
See Appendix 2.
Invertebrate Systematics
545
Female. Similar to male, except for: antennae almost
reaching apex of abdomen; scape without enlarged area and
with two long setae at middle of dorsal face; prosternum
without ovate set of fine punctures; each elytron with
interstriae 2 and 4 joined at apex and strongly posteriorly
extended, forming a spine; ovipositor as in P. sulcatus;
spermathecal duct short, ~1.5 times length of spermatheca,
without sclerotised basal region (Fig. 282); spermatheca with
somewhat globose apex (Fig. 282).
Distribution
Venezuela (Aragua).
Redescription
BL: 5.0–5.7 mm, BW: 1.5–1.6 mm. Body somewhat convex;
reddish brown to reddish dark brown (Figs 46, 47). Dorsal
integument of head, except anterior angles, and all pronotum
with undulate microstriae and large punctures contiguous
(Fig. 96); striae of elytra with large punctures; metaventrite,
lateral regions, and abdominal tergites with moderately sized
punctures uniformly distributed.
Male. Head with front slightly deflected (Fig. 96); V-shaped
frontal sulcus complete, sometimes inconspicuous at the middle;
anterior angles curved and prominent, and slightly median
longitudinal sulcus; eyes with a moderately sized puncture
with long seta on basal half of dorsal margin. Antennae
exceeding apex of abdomen (Fig. 48); scape with slightly
enlarged area and long setae on basal half of exposed dorsal
face; scape shorter than antennomere 2 and 3 combined;
antennomeres 4–11 oblong shape. Labrum with six setae
medially, equal in length; each lateral third, antero-internal
shorter than postero-external seta. Mandibles with dorsal teeth
weakly developed (Fig. 127); internal border symmetrical, each
with two acute teeth, basal anteriorly projected. Maxillary
palpus with palpomere 4 longer than 2 and 3 combined.
Mentum subquadrate, as wide as long. Pronotum wider than
long (PW/PL = 1.45) (Fig. 96); anterior angles not projected; on
apical half with somewhat longitudinal median depression and
on basal half a transverse depression; longitudinal sulcus
inconspicuous; longitudinal sulcus of posterior angles
developed and reaching middle of pronotum; abrupt
constriction at basal third and, anterior to it, in each side with
a short tooth and slight emargination; apical two-thirds with
slightly sinuous sides. Prosternum with conspicuous ovate set
of fine punctures in a median region near apex. Femur of anterior
legs with short, hard setae on groove of ventral margin. Elytra
together longer than wide (EL/BW = 1.17), occulting tergite 3 and
basal half of tergite 4; stria 6 at basal half and stria 7 complete;
striae with same width of interstriae; interstriae carinate;
interstriae 2 and 4 joined at apex and weakly posteriorly
extended; epipleural carina developed. Metaventrite with
conspicuous median fovea near apex. Abdominal segments
3–6 subparallel sides; sternite 3 with conspicuous transverse
carina; tergite 9 with short ventral struts; tergite 10 with one
pair of long setae on apex, sometimes with one of inconspicuous
seta near long setae. Median lobe of aedeagus with bulbous base
in ventral view and curved apex in lateral view (Fig. 234); lateral
lobes exceed a little apex of median lobe in lateral view; internal
sclerites as in Fig. 235.
Biological notes
This species has been noted only as ‘under bark’ and on ‘fungusy
log’.
Remarks
Piestus paradoxus is easily separated from the other species of
Piestus, except P. acuminatus, by the following character:
interstriae 2 and 4 of each elytron joined at the apex and
weakly posteriorly extended on males and strongly on females
(Figs 48, 49).
Piestus paradoxus may be confused with P. acuminatus
because they also have interstriae posteriorly extended on
elytra. However, P. paradoxus has very long antennae,
exceeding the apex of abdomen in males and almost reaching
the apex of abdomen in females (Figs 48, 49), only large punctures
on striae of elytra, conspicuous median fovea near the apex of
metaventrite, long tube of median lobe of aedeagus and lateral
lobes exceeding the apex of median lobe (Fig. 234), and
somewhat globose apex of spermatheca (Fig. 282).
Piestus acuminatus has shorter antennae, almost reaching
the apex of elytra in males and reaching the apex of elytra in
females (Fig. 53), microgranulate texture and large punctures on
striae of elytra, slightly median depression near the apex of
metaventrite, short tube of median lobe of aedeagus (Fig. 242)
and lateral lobes reaching the apex of median lobe, and very
elongate spermatheca (Fig. 285).
Piestus boliviensis, sp. nov.
(Figs 50, 97, 236, 237)
Type material
Holotype deposited in SEMC, male, labels: (1) ‘Bolivia: La Paz,
Coroico/Cerro Uchumachi, 2150m/1612.160 S, 6743.330 W/27-I-2001,
R.A. Anderson,/2nd growth cloud forest litter/BOL1A01-013’ [white
label, printed in black]; (2) ‘bar code/SM0454517/KUNHM-ENT’ [white
label, printed in black]; (3) ‘Piestus/sp./Det: J.S. Ashe, 2004’ [white label,
first two lines handwritten, third printed in black].
Description
BL: 5.3 mm, BW: 1.4 mm. Body flattened dorsoventrally;
reddish dark brown (Fig. 50). Dorsal integument of head,
except anterior angles, and all pronotum with large punctures
contiguous (Fig. 97); striae of elytra with large punctures;
546
Invertebrate Systematics
metaventrite and abdominal tergites with moderately sized
punctures uniformly distributed.
Male. Head with front slightly deflected (Fig. 97); V-shaped
frontal sulcus complete; anterior angles curved and prominent;
eyes with a moderately sized puncture with short seta on basal
half of dorsal margin. Scape antennal with long setae on basal
half of exposed dorsal face; scape subequal to antennomere 2
and 3 combined (Fig. 50); antennomeres 4–5 oblong shape
(material damaged, not possible to verify antennomeres 6–11).
Mandibles with dorsal teeth weakly developed; internal border
slightly asymmetrical, each with one small and acute tooth at
middle and, anterior to it, in right mandible with weak area
that resembles a tooth, absent in left mandible. Maxillary
palpus with palpomere 4 longer than 2 and 3 combined.
Pronotum wider than long (PW/PL = 1.42) (Fig. 97); anterior
angles weakly projected; on apical half with somewhat
longitudinal median depression and on basal half a transverse
depression; longitudinal median sulcus inconspicuous;
longitudinal sulcus of posterior angles developed and reaching
middle of pronotum; abrupt constriction at basal third and,
anterior to it, in each side with short tooth and slight
emargination; apical two-thirds with slightly sinuous sides.
Prosternum with conspicuous ovate set of fine punctures in
median region near apex. Femur of anterior legs with short,
hard setae on groove of ventral margin. Elytra together shorter
than wide (EL/BW = 0.91), apical margin strongly emarginate at
middle; stria 6 absent and stria 7 complete; striae with same width
of interstriae; interstriae carinate; epipleural carina developed.
Metaventrite with a conspicuous median depression near apex.
Abdominal segments 3–6 with subparallel sides; sternite 3 with
conspicuous transverse carina; tergite 9 with short ventral
struts; tergite 10 with two pairs of setae on apex (material
damaged, not possible to verify length of setae). Median lobe
of aedeagus with bulbous base in ventral view and slightly
curved apex in lateral view (Fig. 236); lateral lobes exceed
little the apex of median lobe in lateral view; internal sclerites
as in Fig. 237.
Female. Unknown.
E. Caron et al.
Piestus aper Sharp, 1876
(Figs 51, 98, 238, 239)
Piestus aper Sharp, 1876: 408 (original description, type locality:
‘St. Paulo’); Blackwelder, 1944: 100 (distribution); Herman, 2001a:
12 (mention); Herman, 2001b: 1788 (distribution).
Piestus (Piestus) aper: Bernhauer & Schubert, 1910: 6 (distribution).
Piestus (Trachypiestus) asper Scheerpeltz, 1952: 292 (emendation of
P. aper Sharp, 1876, characters, distribution).
Piestus asper: Herman, 2001a: 12 (note: unjustified emendation by
Scheerpeltz (1952)); Herman, 2001b: 1788 (mention).
Piestus (Trachypiestus) schadei Scheerpeltz, 1952: 295 (original
description, type locality: ‘Villa Rica (Paraguay)’). New synonym.
Piestus schadei: Herman, 2001b: 1794 (distribution).
Type material
Piestus aper Sharp, 1876. Lectotype deposited in BMNH, sex
undetermined, with labels: (1) ‘Piestus/aper./Amazons/Type D.S.’ [white
label, together with the specimen, handwritten]; (2) ‘Syn-/Type’ [circle white
label with border light blue, printed in black]; (3) ‘Type’ [circle white label
with border red, printed in black]; (4) ‘Amaz./Sr. Paula?’ [circle green label,
handwritten]; (5) S. America:/Brazil.’ [white label, printed in black]; (6)
‘Sharp Coll/1905-313.’ [white label, printed in black]; (7) ‘Syntype/Piestus
apes?/Sharp, 1876/det. R.G. Booth 2008’ [white label, the three first lines
handwritten, the last one printed in black]. Note: In the original description
Sharp (1876) specified two specimens of sex undetermined. We received
from BMNH only one specimen. Another specimen is deposited in BMNH
(Dr Roger Booth, BMNH, pers. comm.).
Piestus (Trachypiestus) schadei Scheerpeltz, 1952. Lectotype
deposited in NMW, sex undetermined, with labels: (1) ‘Villa Rica/
Paraguay’ [white label, printed in black]; (2) ‘Fr. Schade leg./8. XII. 1924’
[white label, first line printed in black, second line handwritten, except ‘1924’
printed in black]; (3) ‘ex coll./Scheerpeltz’ [blue label, printed in black]; (4)
‘TYPUS/Piestus (Trachy-/piestus) Schadei/O. Scheerpeltz’ [red label, the
first and fourth lines printed in black, the second and third handwritten]; (5)
‘Schadei/Scheerp.’ [light green label/handwritten]. Note: In the original
description Scheerpeltz (1952) specified a male and female observed.
However, we received from NMW only one specimen.
Additional material
See Appendix 2.
Distribution
Redescription
Bolivia (La Paz).
BL: 3.2–5.2 mm, BW: 1.0–1.4 mm. Body flattened
dorsoventrally; light brown to reddish dark brown (Fig. 51).
Dorsal integument of dorsum of head and all pronotum with
large punctures contiguous (Fig. 98); striae of elytra with large
punctures; metaventrite and abdominal tergites with moderately
sized punctures uniformly distributed.
Male. Head with front slightly deflected (Fig. 98); V-shaped
frontal sulcus incomplete, curved arms not joined medially;
anterior angles curved and prominent; eyes with a moderately
sized puncture with short seta on basal half of dorsal margin.
Antennae exceed little apex of elytra (Fig. 51); scape with
one long seta at middle of dorsal face; scape subequal to
antennomere 2 and 3 combined; antennomeres 4–11 oblong
shape. Labrum with six setae medially, equal in length; each
lateral third, antero-internal seta longer than postero-external seta.
Mandibles with dorsal teeth weakly developed; internal border
smooth. Maxillary palpus with palpomere 4 longer than to 2 and 3
combined. Mentum ~1.5 times as wide as long. Pronotum wider
Biological notes
The holotype was collected in ‘2nd growth cloud forest litter’.
Remarks
Piestus boliviensis is easily separated from the other species of
Piestus by the slightly asymmetrical internal borders of
mandibles, only one small and acute tooth in each one, and
elytra together shorter than wide with apical margin strongly
emarginate at the middle (Fig. 50).
Etymology
The species name is derived from the name of Bolivia, where
the holotype was collected.
Revision of Piestus with remarks on related genera
than long (PW/PL = 1.56) (Fig. 98); anterior angles strongly
projected; apical half with a somewhat longitudinal median
depression and on basal half a transverse depression;
longitudinal median sulcus inconspicuous; longitudinal sulcus
of posterior angles strongly developed and exceed middle of
pronotum; abrupt constriction at basal third and, anterior to it, in
each side with a short tooth and slight emargination; apical
two-thirds with sinuous sides. Femur of anterior legs with
short, hard setae on groove of ventral margin. Elytra together
as long as wide (EL/BW = 0.99); stria 6 absent and stria 7
complete; striae wider than interstriae; interstriae carinate;
basal half of interstriae 2 and 5 and apical half of interstria 4
prominent; epipleural carina developed. Abdominal segments
3–6 with subparallel sides; sternite 3 with conspicuous transverse
carina; tergite 9 with long ventral struts; tergite 10 with one
pair of long setae on apex, sometimes with one pair of
inconspicuous setae near long setae. Median lobe of aedeagus
with bulbous base in ventral view and curved apex in lateral view
(Fig. 238); lateral lobes reaching apex of median lobe in lateral
view; internal sclerites as in Fig. 239.
Female. Similar to male. Ovipositor as in P. sulcatus;
spermathecal duct ~2 times length of spermatheca, without
sclerotised basal region (Fig. 283); spermatheca with globose
apex and accessory gland at apical third (Fig. 283).
Invertebrate Systematics
Piestus (Piestus) crassicornis: Bernhauer & Schubert, 1910: 7
(distribution).
Piestus (Trachypiestus) crassicornis: Scheerpeltz, 1952: 292 (characters,
distribution).
Type material
Piestus angularis Fauvel, 1864. Holotype deposited in IRSNB, female,
with labels: (1) ‘Brésil/Sa Catharina’ [white label, handwritten]; (2) ‘Caracas’
[white label, handwritten]; (3) ‘Mexique’ [white label, handwritten]; (4)
‘crassicornis Shp./parait u? même’ [white label, handwritten]; (5)
‘angularis/FvL.’ [white label, handwritten]; (6) ‘R.I.Sc.N.B. 17.479/
Piestus/Coll. et det. A. Fauvel’ [white label, first and third lines printed in
black and second line handwritten]; (7) ‘Type’ [red label, printed in black].
Note: In the original description Fauvel (1864) specified only one specimen
observed.
Piestus crassicornis Sharp, 1887. Holotype deposited in BMNH, female,
with labels: (1) ‘Piestus crassi-/cornis Type D.S./V. de Chiriqui 2500-/4000ft.
Champion’ [white label, together with the specimen, handwritten]; (2) ‘Holo-/
Type’ [circle white label with border red, printed in black]; (3) ‘Sp. figured.’
[white label, printed in black]; (4) ‘V. de Chiriqui,/25-4000 ft./Champion.’
[white label, printed in black]; (5) ‘B.C.A. Col. I. 2./Piestus/crassicornis,/
Sharp.’ [white label, printed in black]; (6) ‘Sharp Coll./1905-313.’ [white
label, printed in black]; (7) ‘Holotype/Piestus/crassicornis/Sharp, 1887/det.
R.G. Booth 2008’ [white label, the four first lines handwritten, the last one
printed in black]. Note: In the original description Sharp (1887) specified only
one specimen observed.
Distribution
Additional material
In the current study the species was examined from Panama
(Bocas del Toro and Colón), Colombia (Amazonas), Suriname
(Marowijne), French Guiana (Saint-Laurent-du-Maroni and
Cayenne), Peru (Loreto, Cusco and Madre de Dios), Bolivia,
Brazil (Amazonas, Bahia and Mato Grosso) and Paraguay
(Guairá and Itapúa). Scheerpeltz (1952) also listed P. aper
from north Argentina.
See Appendix 2.
Biological notes
Piestus aper has been found in litter, and associated with fruit
falls, flower falls or decaying logs. Some specimens were
collected in flight intercept traps, or by Berlese and Winkler
extraction of leaf litter.
Remarks
This species is similar to P. sulcipennis but differs by characters
cited under remarks of P. sulcipennis.
Piestus angularis Fauvel, 1864
(Figs 52, 99, 240, 241, 284)
Piestus angularis Fauvel, 1864: 31 (original description, type locality:
‘Brésil (Santa-Catharina’); Blackwelder, 1944: 100 (distribution, error:
Fauvel, 1865: 35); Herman, 2001b: 1788 (distribution); NavarreteHeredia et al., 2002: 208 (distribution).
Piestus (Piestus) angularis: Bernhauer & Schubert, 1910: 6 (distribution,
error: Fauvel, 1865: 35).
Piestus (Trachypiestus) angularis: Scheerpeltz, 1952: 292 (characters,
distribution, error: Fauvel, 1865: 35).
Piestus crassicornis Sharp, 1887: 716 (original description, type locality:
‘Panama, Volcan de Chiriqui 4000 to 6000 feet’); Blackwelder, 1944:
100 (distribution); Herman, 2001b: 1790 (distribution). New synonym.
547
Redescription
BL: 4.6–6.8 mm, BW: 1.4–1.9 mm. Body somewhat convex;
reddish brown to reddish dark brown (Fig. 52). Dorsal
integument of head, except anterior angles, and all pronotum
with large punctures contiguous (Fig. 99); striae of elytra with
large punctures; metaventrite and abdominal tergites with
moderately sized punctures uniformly distributed.
Male. Head with front slightly deflected (Fig. 99); V-shaped
frontal sulcus complete; anterior angles curved and prominent,
and conspicuous median longitudinal sulcus with fovea at the
base; eyes with moderately sized puncture with short seta on
basal half of dorsal margin. Antennae reaching apex of abdomen
(Fig. 52); scape with one long seta at middle of dorsal face; scape
subequal to antennomere 2 and 3 combined; antennomeres 4–11
oblong shape. Labrum with six setae medially, equal in length;
each lateral third, antero-internal seta as long as postero-external
seta. Mandibles with dorsal teeth weakly developed; internal
border symmetrical, each with two acute teeth, basal anteriorly
projected. Maxillary palpus with palpomere 4 longer than to 2 and
3 combined. Mentum subquadrate, as wide as long. Pronotum
wider than long (PW/PL = 1.31) (Fig. 99); anterior angles not
projected; on apical half with somewhat longitudinal median
depression and on basal half a transverse depression; longitudinal
median sulcus inconspicuous; longitudinal sulcus of posterior
angles strongly developed and exceed middle of pronotum;
abrupt constriction at basal third and, anterior to it, in each
side with short tooth and a slight emargination; apical twothirds with sinuous sides. Prosternum with conspicuous ovate
set of fine punctures in median concavity near the apex. Femur of
anterior legs with short, hard setae on groove of ventral margin.
548
Invertebrate Systematics
Elytra together as long as wide (EL/BW = 0.98); stria 6 absent
and stria 7 complete; striae wider than interstriae; interstriae
carinate; basal half of interstriae 2 and 5 prominent; epipleural
carina developed. Metaventrite with conspicuous median fovea
near apex. Abdominal segments 3–6 with subparallel sides;
sternite 3 with conspicuous transverse carina; tergite 9 with
short ventral struts; tergite 10 with one pair of long setae on
apex, sometimes with one pair of inconspicuous setae near long
setae. Median lobe of the aedeagus with bulbous base in ventral
view and curved apex in lateral view (Fig. 240); lateral lobes
exceed little apex of median lobe in lateral view; internal sclerites
as in Fig. 241.
Female. Similar to male, except for: antennae exceeding
apex of elytra, but not reaching apex of abdomen; prosternum
without ovate set of fine punctures and without median
concavity near apex; ovipositor as in P. sulcatus; spermathecal
duct long, ~5 times length of spermatheca, without sclerotised
basal region (Fig. 284); spermatheca somewhat L-shaped
(Fig. 284).
Distribution
In the current study the species was examined from Mexico
(Veracruz), Belize (Cayo), Guatemala (Zacapa), Costa Rica
(Alajuela, Heredia, San José, Limón and Puntarenas), Panama
(Chiriquí), Suriname (Marowijne and Para), French Guiana
(Saint-Laurent-du-Maroni
and
Cayenne),
Ecuador
(Esmeraldas, Napo and Zamora-Chinchipe), Peru (Cusco),
Bolivia (Cochabamba), Brazil (Santa Catarina) and Paraguay
(Alto Paraná). Navarrete-Heredia et al. (2002) list P. angularis
also from Honduras, Nicaragua and Colombia.
Biological notes
Piestus angularis has been found in litter, near streams or
associated with decaying logs. Some specimens were collected
by flight intercept and Malaise traps.
Remarks
This species is easily distinguished from the other species of
Piestus by the conspicuous median longitudinal sulcus with a
fovea at the base of the vertex (Fig. 99), two acute teeth on
symmetrical internal borders of mandibles, and only one long seta
on scape (male and female).
Piestus acuminatus, sp. nov.
(Figs 53, 100, 135, 144, 242, 243, 285)
Type material
Holotype deposited in SEMC, male, labels: (1) ‘Ecuador: Napo/El Chaco,
4.8 km NW on road/to Oyacachi, 1750m/0180 22’S, 77500 38’W/7 Nov
1999, Z.H. Falin/ECU1F99 119 ex: pyrethrum/fogging mildewy downed
tree’ [white label, printed in black]; (2) ‘bar code/SM0351487/KUNHMENT’ [white label, printed in black]. 3 paratypes distributed: 2 in SEMC: 1
female, the same first label of holotype, (2) ‘bar code/SM0351486/KUNHMENT’ [white label, printed in black]; 1 male, labels: (1) ‘Ecuador: Napo/
Baeza-Tena Road, 15 km/W of Cosanga, Reserva Sierra/Azul, 2350m,
0400 55’S, 77560 9’W/5 Nov 1999, Z.H. Falin/ECU1F99 097 ex: rotting
cut lumber’ [white label, printed in black]; (2) ‘bar code/SM0355660/
E. Caron et al.
KUNHM-ENT’ [white label, printed in black]; 1 in FMNH, 1 male,
labels: (1) ‘Ecuador: Napo/Cosanga, 4.2 km S on Baeza-Tena/Road then
2.9 km W on pipeline/access road, 2150m/0370 19’S, 77500 1’W/7 Nov
1999, Z.H. Falin/ECU1F99 120 ex: on/under/bark downed logs’ [white
label, printed in black]; (2) ‘bar code/SM0152154/KUNHM-ENT’ [white
label, printed in black].
Description
BL: 5.0 mm, BW: 1.6 mm. Body convex; reddish brown to
reddish dark brown (Fig. 53). Dorsal integument of head and
all pronotum with large punctures contiguous; striae of elytra with
microgranulate texture and large punctures (Fig. 100);
metaventrite and abdominal tergites with moderately sized
punctures uniformly distributed.
Male. Head with front slightly deflected (Fig. 100); Vshaped frontal sulcus complete; anterior angles curved and
prominent, and with median fovea at base; eyes prominent
with moderately sized puncture with short seta on basal half of
dorsal margin. Antennae almost reaching apex of abdomen
(Fig. 53); scape with long seta on basal half of exposed dorsal
face; scape shorter than antennomere 2 and 3 combined;
antennomeres 4–11 oblong shape. Labrum with six setae
medially, equal in length; each lateral third, antero-internal
seta shorter than postero-external seta. Mandibles with dorsal
teeth weakly developed; internal border symmetrical, each with
two acute teeth, basal anteriorly projected. Maxillary palpus
with palpomere 4 longer than 2 and 3 combined. Mentum
subquadrate, as wide as long (Fig. 135). Pronotum wider than
long (PW/PL = 1.48) (Fig. 100); anterior angles weakly
projected; on apical half with somewhat longitudinal median
depression and on basal half a transverse depression; longitudinal
median sulcus conspicuous; longitudinal sulcus of posterior
angles developed and reaching middle of pronotum; abrupt
constriction at basal third and, anterior to it, in each side with
short tooth and slight emargination; apical two-thirds with
sinuous sides. Prosternum with conspicuous ovate set of fine
punctures in median region near apex. Femur of anterior legs with
short, hard setae on groove of ventral margin. Elytra somewhat
longer than wide (EL/BW = 1.05), occulting tergite 3; stria 6
absent and stria 7 complete; striae wider than interstriae;
interstriae carinate; interstriae 1 and 4 joined at apex and
weakly posteriorly extended (Fig. 53); epipleural carina
developed. Metaventrite with a conspicuous median
depression near apex. Abdominal segments 3–6 subparallel
sides; sternite 3 with conspicuous transverse carina (Fig. 144);
tergite 9 with short ventral struts; tergite 10 with two pairs of
long setae on apex, apical the longest. Median lobe of the
aedeagus with bulbous base in ventral view and curved apex
in lateral view (Fig. 242); lateral lobes reaching apex of median
lobe in lateral view; internal sclerites as in Fig. 243.
Female. Similar to male, except for: antennae reaching
the apex of elytra; scape with two long setae at the middle of
the dorsal face; prosternum without ovate set of fine punctures;
ovipositor as in P. sulcatus; spermathecal duct very short, about
the half of length of spermatheca, without sclerotised basal
region (Fig. 285); spermatheca elongate (Fig. 285).
Distribution
Ecuador (Napo).
Revision of Piestus with remarks on related genera
Biological notes
This species was collected under bark and cited from ‘rotting
cut lumber’.
Remarks
Piestus acuminatus is similar to P. rugosus and P. elegans
because they share microgranulate texture and large punctures
on striae of elytra. However, P. acuminatus is easily separated by
interstriae 1 and 4 of each elytron joined at the apex and weakly
posteriorly extended (male and female) (Fig. 53).
Piestus rugosus and P. elegans do not have interstriae
posteriorly projected.
Piestus acuminatus is distinguished from P. paradoxus by
characters cited under remarks for P. paradoxus.
Etymology
The name refers to the sharp point on the apex of each elytron.
Piestus rugosus Sharp, 1876
(Figs 54, 101, 244, 245, 286)
Piestus rugosus Sharp, 1876: 407 (original description, type locality:
‘Ega’); Blackwelder, 1944: 101 (distribution); Herman, 2001b: 1794
(distribution).
Piestus (Piestus) rugosus: Bernhauer & Schubert, 1910: 7 (distribution).
Piestus (Trachypiestus) rugosus: Scheerpeltz, 1952: 294 (characters,
distribution).
Type material
Holotype deposited in BMNH, female, with labels: (1) ‘Piestus rugosus/
Amazons/Type/D.S.’ [white label, together with the specimen, handwritten];
(2) ‘Holo-/type’ [circle white label with border red, printed in black]; (3)
‘Ega.’ [circle green label, handwritten]; (4) ‘S. America:/Brazil.’ [white label,
printed in black]; (5) ‘Sharp Coll./1905-313.’ [white label, printed in black];
(6) ‘Holotype/Piestus/rugosus/Sharp, 1876/det. R.G. Booth 2008’ [white
label, the four first lines handwritten and the last one printed in black].
Note: In the original description Sharp (1876) specified only one specimen
observed.
Invertebrate Systematics
549
internal seta shorter than postero-external seta. Mandibles with
dorsal teeth weakly developed; internal border symmetrical,
each with two acute teeth, basal anteriorly projected. Maxillary
palpus with palpomere 4 longer than 2 and 3 combined. Mentum
subquadrate, as wide as long. Pronotum wider than long (PW/
PL = 1.49) (Fig. 101); anterior angles not projected; on apical
half with somewhat longitudinal median depression and on
basal half a transverse depression; longitudinal median sulcus
inconspicuous; longitudinal sulcus of posterior angles developed
and reaching middle of pronotum; abrupt constriction at basal
third and, anterior to it, in each side with short tooth and slight
emargination; apical two-thirds with sinuous sides. Prosternum
with conspicuous ovate set of fine punctures in median region
near apex. Femur of anterior legs with short, hard setae on groove
of ventral margin. Elytra together as long as wide (EL/
BW = 0.97); stria 6 absent and stria 7 complete; striae wider
than interstriae; interstriae carinate; epipleural carina developed.
Metaventrite with a conspicuous depression near apex.
Abdominal segments 3–6 subparallel sides; sternite 3 with
conspicuous transverse carina; tergite 9 with long ventral
struts; tergite 10 with two pairs of long setae on apex, apical
the longest. Median lobe of aedeagus with bulbous base in ventral
view and curved apex in lateral view (Fig. 244); lateral lobes not
reaching apex of median lobe in lateral view; internal sclerites as
in Fig. 245.
Female. Similar to male, except for: antennae exceeding
apex of elytra, but not reaching apex of abdomen; scape with two
long setae at middle of dorsal face; prosternum without ovate
set of fine punctures; ovipositor as in P. sulcatus; spermathecal
duct short, about same length of spermatheca, without sclerotised
basal region (Fig. 286); spermatheca somewhat elongate
(Fig. 286).
Distribution
Guyana, Ecuador (Sucumbios),
(Cochabamba) and Brazil (Pará).
Peru
(Cusco),
Bolivia
Additional material
Biological notes
See Appendix 2.
This species has been noted occurring under bark and it was
also collected in flight intercept traps.
Redescription
BL: 4.0–6.0 mm, BW: 1.3–1.6 mm. Body somewhat convex;
reddish brown to reddish dark brown (Fig. 54). Dorsal
integument of head and all pronotum with large punctures
contiguous (Fig. 101); striae of elytra with microgranulate
texture and large punctures; metaventrite and abdominal
tergites with moderately sized punctures uniformly distributed.
Male. Head with front slightly deflected (Fig. 101); Vshaped frontal sulcus complete; anterior angles curved and
prominent, and median longitudinal sulcus with fovea at base;
eyes with moderately sized puncture with short seta on basal
half of dorsal margin. Antennae almost reaching apex of
abdomen; scape with few long setae on basal half of exposed
dorsal face (as a tuft); scape subequal to antennomere 2 and 3
combined; antennomeres 4–11 oblong shape (Fig. 101). Labrum
with six setae medially, equal in length; each lateral third, antero-
Remarks
Piestus rugosus is similar to P. acuminatus and P. elegans
because they share microgranulate texture and large punctures
on striae of elytra. However, P. rugosus differs from
P. acuminatus by characters cited under P. acuminatus, and it
differs from P. elegans by antennomeres 4–11 oblong shape
(Fig. 54), and short spermathecal duct about the same length as
spermatheca (Fig. 286). Piestus elegans has antennomeres 4–11
somewhat bead-like in females (Fig. 55), and long spermathecal
duct, ~2 times the length of spermatheca (Fig. 287).
Piestus elegans, sp. nov.
(Figs 55, 102, 287)
550
Invertebrate Systematics
E. Caron et al.
Type material
Etymology
Holotype deposited in SEMC, female, labels: (1) ‘Peru: Tambopata Prov./
Madre de Dios Dpto./15 km NE Puerto’ [white label, printed in black];
(2) ‘Maldonado Reserva/Cuzco Amazónico/12330 S, 69030 W/200m, Z2
E15’ [white label, printed in black]; (3) ‘26 June 1989, D. Silva/R.
A. Leschen #285/ex., flight intercept’ [white label, printed in black]; (4)
‘Piestus/Peru sp. 3/det J.S. Ashe 19’ [white label, first two lines handwritten,
third printed in black].
The specific name refers to the elegant, necklace-shaped
antennomeres 4–11.
Additional taxonomic notes
Subfamily PIESTINAE Erichson, 1839
Description
BL: 4.3 mm, BW: 1.2 mm. Body somewhat convex; reddish
brown (Fig. 55). Dorsal integument of head and all pronotum
with large punctures contiguous (Fig. 102); striae of elytra with
microgranulate texture and large punctures; metaventrite and
abdominal tergites with moderately sized punctures uniformly
distributed.
Female. Head with front slightly deflected (Fig. 102); Vshaped frontal sulcus complete; anterior angles curved and
prominent, and with slightly median fovea at base. Antennae
exceed base of elytra, but not reaching apex of elytra; scape with
one short seta at middle of dorsal face; scape subequal to
antennomere 2 and 3 combined; antennomeres 4–11 somewhat
bead-like (Fig. 55). Mandibles with dorsal teeth weakly
developed; internal border symmetrical, each with two acute
teeth, basal anteriorly projected. Maxillary palpus with
palpomere 4 longer than 2 and 3 combined. Pronotum wider
than long (PW/PL = 1.53) (Fig. 102); anterior angles weakly
projected; on apical half with slightly longitudinal median
depression and on basal half transverse depression;
longitudinal median sulcus inconspicuous; longitudinal sulcus
of posterior angles developed and reaching middle of pronotum;
abrupt constriction at basal third and, anterior to it, in each side
with a short tooth and slight emargination; apical two-thirds with
sinuous sides. Femur of anterior legs with short, hard setae on
groove of ventral margin. Elytra together as long as wide (EL/
BW = 1.00); stria 6 absent and stria 7 complete; striae wider than
interstriae; interstriae carinate; epipleural carina developed.
Metaventrite with conspicuous median depression near apex.
Abdominal segments 3–6 subparallel sides; sternite 3 with
conspicuous transverse carina; ovipositor as in P. sulcatus;
tergite 10 with two pairs of long setae on apex, apical the
longest. Spermathecal duct ~2 times length of spermatheca,
without sclerotised basal region (Fig. 287); spermatheca
somewhat elongate (Fig. 287).
Male. Unknown.
Distribution
Peru (Madre de Dios).
Biological notes
The holotype was collected in a flight intercept trap.
Remarks
Piestus elegans is similar to P. acuminatus and P. rugosus but
differs by characters listed under remarks for P. acuminatus and
P. rugosus.
Hypotelus Erichson, 1839
Hypotelus Erichson, 1839: 31. For review of literature of the genus, see
Herman (2001b).
Antropiestus Bernhauer, 1917: 45 (as subgenus of Piestus); Scheerpeltz,
1952: 295 (subgenus of Piestus); Herman, 2001b: 1788 (subgenus of
Piestus). Type-species. Piestus (Antropiestus) andinus Bernhauer,
1917 (fixed by original designation and monotypy). New synonym.
Eccoptopiestus Scheerpeltz, 1952: 295 (as subgenus of Piestus); Herman,
2001b: 1788 (subgenus of Piestus). Type species. Piestus laevis
Solsky, 1872 (fixed by original designation and monotypy). New
synonym.
Remarks
Eccoptopiestus and Antropiestus, previously considered as
subgenera of Piestus, are here placed as junior synonyms of
Hypotelus, since their type species, P. (E.) laevis Solsky, 1872
and P. (A.) andinus Bernhauer, 1917, are transferred to Hypotelus
based on some adult characters observed in type material (see
remarks under each species below).
Hypotelus laevis (Solsky, 1872), comb. nov.
(Figs 56, 104, 128, 136, 148, 246, 247)
Piestus laevis Solsky, 1872: 311 (original description: ‘Monte-Rico
(Pérou)’); Blackwelder, 1944: 100 (distribution, error: Solsky,
1871); Herman, 2001b: 1791 (distribution).
Piestus (Piestus) laevis: Bernhauer & Schubert, 1910: 7 (distribution,
error: Solsky, 1871).
Piestus (Eccoptopiestus) laevis: Scheerpeltz 1952: 295 (characters,
distribution, error: Solsky, 1871).
Type material
Neotype (here designated) deposited in IRSNB, sex undetermined, with
labels: (1) ‘Yuracaris/Bolivie’ [white label, handwritten]; (2) ‘laevis
Solsky/var?’ [white label, handwritten]; (3) ‘R.I.Sc.N.B. 17.479/
Hypotelus/Coll. et det. A. Fauvel’ [white label, first and third lines printed
in black, second line handwritten]. Note: In the original description Solsky
(1872) did not specify how many specimens he observed. However, type
material from Peru was not found in ZIN (Zoological Institute, Russian
Academy of Sciences, St. Petersburg, Russia) where Solsky’s collection is
deposited, and probably any types there are lost. However, we received
from IRSNB (Fauvel’s collection) one specimen, which is here designated
as neotype, and possibly it is an authentic specimen sent by Solsky to Fauvel.
There is another case where this happened, Aleochara sareptana Solsky
1874 (Dr Didier Drugmand, IRSNB, pers. comm.). In this way, we are
designating a neotype with the purpose of clarifying the taxonomic status
of the taxon (Article 75.3.1, ICZN 1999).
Additional material
See Appendix 2.
Revision of Piestus with remarks on related genera
Redescription
BL: 3.0–3.6 mm, BW: 0.8–0.9 mm. Body flattened
dorsoventrally; light brown to dark brown with one-quarter
apical of elytra darker (Fig. 56). Dorsal integument of head
and pronotum with fine punctures and undulate microstriae;
elytra with fine punctures disperse and only one longitudinal
striae finely punctate, closely to internal margin.
Male. Head with pair of broad and short pointed frontal
processes, as long as scape, and basal distance between processes
narrower than basal width of each one (Fig. 104); V-shaped
frontal sulcus incomplete; anterior angles curved and weakly
prominent; eyes with one moderately sized puncture with long
setae on basal half of dorsal margin. Antennae short, almost
reaching apex of elytra (Fig. 56); with one short seta near apex on
the dorsal face; antennomeres 5–11 entirely with microsetae and
some long setae dispersed; scape the longest, subequal to
antennomere 2 and 3 combined; antennomere 4 shortest and
5–11 somewhat oblong shape. Labrum with six setae medially,
external longest; each lateral third, three long setae, two apical
and one subapical, the shortest. Mandibles with bifurcate apex,
dorsal teeth shorter than ventral (Fig. 128); internal border with
one acute tooth at middle; prostheca well developed. Maxillary
palpus with palpomere 3 wider than long and 4 longer than 2 and
3 combined. Mentum 2 times wider than long and anterior
angles emarginate (Fig. 136); ligula slightly emarginate and
two pairs of conspicuous long setae on anterior margin, one
pair near median sclerite and other on apex of median sclerite.
Gular sutures joined. Pronotum wider than long (PW/PL = 1.33)
(Fig. 104); anterior angles not projected; longitudinal median
sulcus conspicuous; gradual constriction at basal third; apical
three-quarters with slightly curved sides. Prosternum with
anterior margin truncate. Elytra somewhat longer than wide
(EL/BW = 1.18), covering tergite 3; when together slightly
emarginate in middle; apical margin with short setae.
Abdominal segments 3–6 parallel sides; segments 3–6 with
two pair of paratergites, segment 7 with only one pair; sternite
7 with apical margin pointed; tergite 9 with short ventral struts;
tergite 10 with some long setae on apex and apical margin with
short fringes; sternite 9 with two pairs of long setae and truncate
apex. Median lobe of aedeagus with bulbous base in ventral
view and not curved apex in lateral view; lateral lobes almost
reaching apex of median lobe in lateral view (Fig. 246); internal
sclerites as in Fig. 247.
Female. Similar to male except for: pair of frontal process
less developed than on male; sternite 8 with short setae on apical
margin (Fig. 148); ovipositor as in P. andinus.
Distribution
Bolivia and Peru (Cusco).
Remarks
The species is excluded from the genus Piestus based on the
current phylogenetic study (see ‘Results’, ‘Cladistic analysis’)
and here transferred to the genus Hypotelus, with which it
shares the following characters: emarginate anterior angles of
mentum, pair of conspicuous setae at the apex of the median
sclerotised plate of the ligula (Fig. 137), joined gular sutures,
Invertebrate Systematics
551
only one complete longitudinal stria on each elytron
(Fig. 56), and short setae on apical margin of sternite 8 of
female (Fig. 148).
Hypotelus laevis differs from H. pusillus and H. andinus by a
pair of broad and short pointed frontal processes on the head
(Fig. 104) and well-developed dorsal teeth of mandibles, forming
a bifurcate apex (Fig. 128).
Hypotelus pusillus and H. andinus do not have a pair of
frontal processes on the head (Fig. 105), or dorsal teeth on the
mandibles (Fig. 129).
Biological notes
Hypotelus laevis has been collected on rotten palm and by
Winkler extraction of leaf litter.
Hypotelus andinus (Bernhauer, 1917), comb. nov.
(Figs 57, 105, 112, 129, 132, 137, 151, 155, 248–250)
Piestus (Antropiestus) andinus Bernhauer, 1917: 45 (original description,
type locality: ‘West-Kolumbien: Umgebung von Cali am Rio Cauca’);
Scheerpeltz, 1933: 993 (distribution); Scheerpeltz, 1952: 295
(characters, distribution).
Piestus andinus: Blackwelder, 1944: 100 (distribution); Herman, 2001b:
1788 (distribution); Newton et al., 2005: 37 (distribution).
Piestus (Antropiestus) strigipennis Bernhauer, 1921: 65 (original
description, type locality; ‘Bolivien: Yuracaris’); Scheerpeltz, 1933:
993 (distribution); Scheerpeltz, 1952: 295 (characters, distribution).
New synonym.
Piestus strigipennis: Blackwelder, 1944: 101 (distribution); Herman,
2001b: 1795 (distribution).
Type material
Piestus (Antropiestus) andinus Bernhauer, 1917. Lectotype deposited in
FMNH, male [damaged specimen: without right anterior leg] with labels: (1)
‘<’ [white label, handwritten, together with the specimen]; (2) ‘Columbia occ/
Cali. Fassl’ [white label/printed in black]; (3) ‘Antropiestus/andinus/Brnh.
Typus’ [light yellow label, handwritten]; (4) ‘Chicago NHMus/M.Bernhauer/
Collection’ [white label, printed in black]. 1 deposited in NMW with labels:
(1) ‘<’ [white label, printed in black]; (2) ‘Columbia occ/Cali. Fassl’ [white
label/printed in black]; (3) ‘Piestus (Subg./Antropiestus)’ [white label,
handwritten]; (4) ‘andinus Bernh./Cotyp.’ [white label, handwritten]; (5)
‘ex coll./Klima’ [blue label, printed in black]; (6) ‘ex coll./Scheerpeltz’ [light
blue label, printed in black]; (7) ‘COTYPUS/Piestus/(Antropiestus)/andinus/
Bernhauer’ [rose label, the first line printed in black, the others handwritten];
(8) ‘andinus/Bh.’ [green label, handwritten]. Paraletotype deposited in NMW,
male, with the same labels of lectotype. Note: In the original description
Bernhauer (1917) did not specify how many specimens he observed. We
received from FMNH and NMW two specimens with type labels, which here
we are considering as type material.
Piestus (Antropiestus) strigipennis Bernhauer, 1921. Syntype deposited in
FMNH, female, with labels: (1) ‘Yuracaris,/A. Fauvel’ (white label, printed in
black]; (2) ‘Bolivien/determ.’ [white label, printed in black]; (3) ‘Piestus
cavi?-/collis Fauvel?/Mus. Hamburg’ [light blue label, handwritten]; (4)
‘strigipennis/Bernh./Typus.’ [light yellow label, handwritten]; ‘Chicago
NHMus/M.Bernhauer/Collections’ [white label, printed in black]. Note: In
the original description Bernhauer (1921) did not specify how many
specimens he observed.
Additional material
See Appendix 2.
552
Invertebrate Systematics
E. Caron et al.
Figs 106–119. 106–108, Piestus sulcatus. 106, Antenna, male; 107, antenna, female; 108, labrum, left epipharynx and right
setae removed; 109, P. capricornis, labrum, left epipharynx and right setae removed; 110, P. fronticornis, sp. rev., labrum, left
epipharynx and right setae removed; 111, P. surrufus, sp. nov., labrum, left epipharynx and right setae removed; 112, Hypotelus andinus,
comb. nov., labrum, left epipharynx and right setae removed. 113–115, Piestus sulcatus, mandibles. 113, Dorsal view, left prostheca
removed; 114, right mandible, ventral view; 115, right mandible, lateral view; 116, P. lacordairei, apex of the mandibles, dorsal
view, setae and left prostheca removed. 117, 118, Piestus puncticollis, apex of the mandibles, setae and left prostheca removed. 117,
Dorsal view; 118, right mandible, lateral view; 119, P. capricornis, apex of the mandibles, setae and left prostheca removed. Scale bar
Figs 106, 107, 115, 117, 118 = 0.5 mm; Figs 108–111, 116, 119 = 0.25 mm; Fig. 112 = 0.15 mm.
Revision of Piestus with remarks on related genera
Invertebrate Systematics
Figs 120–137. 120, Piestus spinosus, apex of the mandibles, setae and left prostheca removed; 121, 122, P. fronticornis, sp.
rev., apex of the mandibles; 121, dorsal view, left prostheca removed; 122, right mandible, lateral view; 123, P. pennicornis, male,
apex of the mandibles, setae and left prostheca removed; 124, P. pennicornis, female, apex of the mandibles, setae and left
prostheca removed; 125, P. filicornis, apex of the mandibles, setae and left prostheca removed; 126, P. surrufus, sp. nov., apex of
the mandibles, setae and left prostheca removed; 127, P. paradoxus, apex of the mandibles, setae and left prostheca removed; 128,
Hypotelus laevis, comb. nov., mandibles, left prostheca removed; 129, Hypotelus andinus, comb. nov., apex of the mandibles,
setae and left prostheca removed; 130, P. fronticornis, sp. rev., maxilla; 131, P. sulcatus, maxillary palpus (palpomeres 2–4); 132,
H. andinus, maxillary palpus (palpomeres 2–4); 133, P. fronticornis, sp. rev., labium, left setae of the ligula and right setae of the
mentum removed; 134, P. sulcatus, mentum, right setae removed; 135, P. acuminatus, sp. nov., mentum, right setae removed;
136, H. laevis, mentum, right setae removed; 137, H. andinus, comb. nov., ligula, right palpus removed. Scale bar Figs 120,
123–130, 133–134 = 0.25 mm; Figs 121, 122 = 0.50 mm; Figs 131, 132, 137 = 0.15 mm; Fig. 136 = 0.10 mm.
553
554
Invertebrate Systematics
E. Caron et al.
Figs 138–148. 138–142, Piestus sulcatus. 138, Anterior leg, posterior view; 139, anterior leg, anterior view; 140,
median leg, posterior view; 141, posterior leg, posterior view; 142, pre-tarsus; 143, P. fronticornis, sp. rev., sternite
1–3; 144, P. acuminatus, sp. nov., sternite 1–3. 145–147, Piestus sulcatus. 145, Tergite 8, male, right setae removed;
146, sternite 8, male, right setae removed; 147, sternite 8, female, right setae removed; 148, Hypotelus laevis, comb.
nov., sternite 8, female, right setae removed. Scale bar Figs 138–141, 143, 144 = 0.50 mm; Fig. 142 = 0.10 mm; Figs
145–148 = 0.25 mm.
Redescription
BL: 6.1 mm, BW: 1.3 mm. Body somewhat flattened
dorsoventrally; entirely black with legs a little lighter
(Fig. 57). Dorsal integument of head and pronotum with fine
punctures and microgranulate sculptures (Fig. 105); elytra with
one longitudinal striae finely punctate and closely to internal
margin, conspicuous only on basal half; metaventrite and
abdomen with fine punctures and microgranulate sculptures.
Revision of Piestus with remarks on related genera
Invertebrate Systematics
555
Figs 149–155. 149, Piestus sulcatus, tergite 9 and 10, male, right setae removed; 150, P. bicornis, tergite 9
and 10, male, right setae removed; 151, Hypotelus andinus, comb. nov., tergite 10, female, right setae removed.
152, 153, Piestus sulcatus. 152, Sternite 9, male; 153, ovipositor, female; 154, P. fronticornis, sp. rev.,
ovipositor, female; 155, H. andinus, comb. nov., ovipositor, female. Scale bar Figs 149, 152–154 = 0.25 mm;
Fig. 150 = 0.50 mm; Figs 151, 155 = 0.15 mm.
Male. Head with front slightly deflected (Fig. 105); Vshaped frontal sulcus incomplete; anterior angles curved and
weakly prominent; eyes slightly prominent from above, and
line with five moderately sized punctures with long setae on
basal half of dorsal margin. Antennae short, almost reaching
apex of elytra (Fig. 57); with three long setae near the apex on
dorsal face; antennomeres 5–11 entirely with microsetae and
some long setae disperse; scape the longest, subequal to
antennomere 2 and 3 combined; antennomere 4 shortest and
5–11 somewhat quadrate. Labrum with six setae medially,
internal the shortest (Fig. 112); each lateral third, four long
setae, two apical and two subapical. Mandibles symmetrical,
one acute tooth on each internal border (Fig. 129); without
dorsal teeth. Maxillary palpus with palpomere 3 wider than
long and 4 longer than 2 and 3 combined (Fig. 132). Mentum
2 times wider than long and anterior angles emarginate; ligula
slightly emarginate and two pair of conspicuous long setae
on anterior margin, one pair near median sclerite and other on
apex of median sclerite (Fig. 137). Gular sutures joined.
Pronotum wider than long (PW/PL = 1.25) (Fig. 105); anterior
angles not projected; longitudinal median sulcus conspicuous;
conspicuous and deep large depression at middle; gradual
constriction at basal third; apical two-thirds with slightly
curved sides. Prosternum with anterior margin truncate.
Elytra somewhat longer than wide (EL/BW = 1.21), covering
tergite 3; when together slightly emarginate on the middle;
apical margin with short setae. Abdominal segments 3–6 with
parallel sides; segments 3–6 with two pair of paratergites,
segment 7 with only one pair; sternite 8 with apical margin
truncate; tergite 9 with short ventral struts; tergite 10 with
556
Invertebrate Systematics
Figs 156–171. Aedeagus, for each species with lateral view and apex in dorsal view respectively.
156–157, Piestus convexus, sp. nov.; 158–159, P. similis, sp. nov.; 160–161, P. lacordairei; 162–163,
P. heterocephalus; 164–165, P. puncticollis; 166–167, P. capricornis; 168–169, P. planatus; 170–171,
P. spinosus. Scale bar = 0.25 mm.
E. Caron et al.
Revision of Piestus with remarks on related genera
Invertebrate Systematics
Figs 172–189. Aedeagus. 172–175, Piestus longipennis. 172, Lateral view; 173, apex in dorsal view; 174, apex
in ventral view; 175, apex in lateral view (everted internal sac). 176–178, Piestus zischkai. 176, Lateral view; 177,
apex in dorsal view; 178, apex in ventral view. 179–181, Piestus bicornis. 179, Lateral view; 180, apex in dorsal
view; 181, ventral view. 182–183, Piestus fronticornis, sp. rev. 182, Lateral view; 183, apex in dorsal view. 184–
185, Piestus validus. 184, Lateral view; 185, apex in dorsal view. 186–187, Piestus rossii, sp. nov. 186, Lateral
view; 187, apex in dorsal view. 188–189, Piestus pennicornis. 188, Lateral view; 189, apex in dorsal view. Scale bar
Figs 172, 176, 179, 181, 182, 184 = 0.50 mm; Figs 173–175, 177, 178, 180, 183, 185, 186–189 = 0.25 mm.
557
558
Invertebrate Systematics
E. Caron et al.
Figs 190–205. Aedeagus, for each species with lateral view and apex in dorsal view respectively. 190–191,
Piestus pygmaeus; 192–193, P. extimus; 194–195, P. buquetii; 196–197, P. filicornis; 198–199, P. abdominalis, sp.
nov.; 200–201, P. niger; 202–203, P. minutus; 204–205, P. penicillatus. Scale bar = 0.25 mm.
some long setae on apex and apical margin with short
fringes, basal half not divided longitudinally; sternite 9 with
two pairs of long setae and truncate apex. Median lobe of
aedeagus with bulbous base in ventral view and somewhat
globose apex in lateral view, tube closed in dorsal view;
lateral lobes exceed little apex of median lobe and curved on
apex in lateral view (Figs 248, 249); internal sclerites as in
Fig. 250.
Female. Similar to male except for: elytra a little shorter,
covering only basal half of tergite 3; abdominal sternite 8 with
Revision of Piestus with remarks on related genera
Invertebrate Systematics
Figs 206–223. Aedeagus. 206–207, Piestus termitis, sp. nov. 206, Lateral view; 207, apex in dorsal view.
208–209, Piestus surrufus, sp. nov. 208, Lateral view; 209, apex in dorsal view. 210–211, Piestus formicinus.
210, Lateral view; 211, dorsal view. 212–214, Piestus sulcatus. 212, Lateral view; 213, apex in dorsal view;
214, ventral view. 215–216, Piestus gounellei. 215, Lateral view; 216, apex in dorsal view; 217, apex in
ventral view. 218–219, Piestus mexicanus. 218, Lateral view; 219, apex in dorsal view. 220–221, Piestus
ecuadorensis, sp. nov. 220, Lateral view; 221, apex in dorsal view. 222–223, Piestus foveolatus, sp. nov.
222, Lateral view; 223, apex in dorsal view. Scale bar = 0.25 mm.
559
560
Invertebrate Systematics
Figs 224–239. Aedeagus, for each species with lateral view and apex in dorsal view respectively. 224–225,
Piestus sulcipennis; 226–227, P. imperfectus, sp. nov.; 228–229, P. costatus; 230–231, P. chiriquensis;
232–233, P. nevermanni; 234–235, P. paradoxus; 236–237, P. boliviensis, sp. nov.; 238–239, P. aper. Scale
bar = 0.25 mm.
E. Caron et al.
Revision of Piestus with remarks on related genera
Invertebrate Systematics
561
Figs 240–250. 240–241, Piestus angularis, aedeagus. 240, Lateral view; 241, apex in dorsal view. 242–243, Piestus
acuminatus, sp. nov. 242, Lateral view; 243, apex in dorsal view. 244–245, Piestus rugosus. 244, Lateral view; 245,
apex in dorsal view. 246–247, Hypotelus laevis, comb. nov. 246, Lateral view; 247, dorsal view. 248–250, Hypotelus
andinus, comb. nov. 248, Lateral view; 249, dorsal view; 250, ventral view. Scale bar = 0.25 mm.
pointed apex and short setae on apical margin; tergite 10 divided
longitudinally on basal half (Fig. 151); ovipositor consisting of
pair of weakly pigmented hemisternites and pair of more apical
coxites, and with many long setae on apex (Fig. 155).
Spermatheca unknown.
Distribution
Colombia (Boyacá), Ecuador and Bolivia.
Biological notes
Hypotelus andinus has been collected in a Malaise trap.
Remarks
Hypotelus andinus is excluded from the genus Piestus based on
the current phylogenetic study (see ‘Results’, ‘Cladistic
analysis’) and here transferred to the genus Hypotelus, with
which it shares the same characters cited above (see remarks
under H. laevis).
The species H. andinus differs from H. pusillus by having a
pronotum with a conspicuous and deep large depression at the
middle (Fig. 105) and by tergite 10 of the female being
longitudinally divided on basal half (Fig. 151).
Hypotelus andinus differs from H. laevis by characters listed
above under remarks for H. laevis.
562
Invertebrate Systematics
E. Caron et al.
Figs 251–268. Spermatheca. 251, Piestus lacordairei; 252, P. heterocephalus; 253, P. capricornis; 254, P. planatus; 255,
P. spinosus; 256, P. longipennis; 257, P. bicornis; 258, P. fronticornis, sp. rev.; 259, P. validus; 260, P. pennicornis; 261,
P. pygmaeus; 262, P. extimus; 263, P. buquetii; 264, P. filicornis; 265, P. abdominalis, sp. nov.; 266, P. niger; 267, P. minutus; 268,
P. penicillatus. Scale bar = 0.25 mm; except Fig. 257 = 0.50 mm.
Revision of Piestus with remarks on related genera
Invertebrate Systematics
Figs 269–287. Spermatheca. 269, Piestus termitis, sp. nov.; 270, P. surrufus, sp. nov.; 271, P. formicinus, sp. nov.;
272, P. sulcatus; 273, P. gounellei; 274, P. mexicanus; 275, P. ecuadorensis, sp. nov.; 276, P. foveolatus, sp. nov.;
277, P. sulcipennis; 278, P. imperfectus, sp. nov.; 279, P. costatus; 280, P. chiriquensis; 281, P. nevermanni; 282, P.
paradoxus; 283, P. aper; 284, P. angularis; 285, P. acuminatus, sp. nov.; 286, P. rugosus; 287, P. elegans, sp. nov.
Scale bar = 0.25 mm.
563
564
Invertebrate Systematics
Subfamily OSORIINAE Erichson, 1839
Tribe ELEUSININI Sharp, 1887
Eleusis Laporte, 1835
Eleusis Laporte, 1835: 131. For review of literature of the genus and a
complete list of synonyms, see Herman (2001b).
Lissopiestus Scheerpeltz, 1952: 295 (as subgenus of Piestus); Herman,
2001b: 1788 (subgenus of Piestus). Type species. Isomalus interruptus
Erichson, 1840 (fixed by original designation and monotypy).
New synonym.
Remarks
Lissopiestus, previously considered as a subgenus of Piestus, is
here synonymised with Eleusis, as its type species P. (L.)
interruptus (Erichson, 1840) is transferred to Eleusis based on
morphological study of the holotype (see remarks under Eleusis
interrupta).
Eleusis interrupta (Erichson, 1840), comb. rest. (restored
combination)
(Figs 58, 103)
Isomalus interruptus Erichson, 1840: 839 (original description, type
locality: ‘Carthagenae in Columbia’); Fauvel, 1864: 35 (characters,
distribution).
Eleusis interrupta: Bernhauer & Schubert, 1910: 7 (distribution); Newton
et al. 2005: 29 (distribution).
Piestus (Piestus) interruptus: Scheerpeltz, 1933: 994 (distribution).
Piestus interruptus: Blackwelder, 1944: 100 (distribution); Herman,
2001b: 1791 (distribution).
Piestus (Lissopiestus) interruptus: Scheerpeltz, 1952: 295 (characters,
distribution).
Type material
Syntype deposited in ZMHB, female, with labels: (1) ‘6819’ [white
label, printed in black]; (2) ‘Type’ [red label, printed in black]; (3)
‘interruptus/E./Carthag. Gory’ [green label, handwritten]; (3) ‘Eleusis/
Cast./1835’ [white label, handwritten]; (4) ‘Zoll. Mus/Berlin’ [white label,
printed in black]; (5) ‘Holotypus/Isomalus/interruptus Erichson, 1840/
labelled by MNHUB 2008’ [red label, printed in black]. Note: In the
original description Erichson (1840) did not specify how many specimens
he observed.
Distribution
Brazil, Colombia, French Guiana, Ecuador, Peru, Bolivia and
Argentina (Herman 2001b; Newton et al. 2005).
Remarks
The specific name ‘interruptus’ is an adjective and is changed
again to the feminine spelling to agree with the gender of the
generic name (Article 34.2; ICZN 1999).
The species is transferred again to the genus Eleusis based on:
head with slight groove on dorsal edge of eye, pronotum strongly
narrowed at base and wider than long (Fig. 58), and abdomen
without paratergites, each abdominal segment with only a
membranous suture between tergum and sternum (Fig. 103).
The syntype of Eleusis interrupta is very similar to specimens
of E. humilis identified and deposited in FMNH. However, some
E. Caron et al.
specimens were dissected and some slight differences were
observed in aedeagi of those specimens, suggesting that a
careful morphological study is necessary before any
conclusion about the specific identity of Eleusis interrupta.
Eleusis interrupta was not used in the present cladistic
analysis because there is no topological correspondence in
character states among some observed characters.
Species Inquirendae (uncertain status)
Piestus longicornis (Lacordaire, 1833)
Zirophorus longicornis Lacordaire, 1833: 65 (original description, type
locality: ‘Cayenne’).
Piestus longicornis: Blackwelder, 1944: 100 (distribution); Herman,
2001b: 1792 (distribution).
Type material
Not found.
Distribution
French Guiana (Herman 2001b).
Biological notes
Unknown (Lacordaire 1833).
Remarks
Lacordaire (1833) described briefly P. longicornis based only on
some characters from the adult and from one larva. The brief adult
description (flat body, red elytra and long antennae) and the type
locality (Cayenne, French Guiana) of P. longicornis suggest this
species is similar to P. spinosus. However, the type material of
P. longicornis was not found and its taxonomic status is still
doubtful.
Piestus longicornis is listed as a valid species only in
Blackwelder (1944) and Herman (2001b), but has not been
otherwise used since its original description, and thus it should
be considered a nomen dubium.
Piestus fulvipes Erichson, 1840
Piestus fulvipes Erichson, 1840: 833 (original description, type locality:
‘Guadeloupe’); Fauvel, 1864: 23 (characters, distribution);
Blackwelder, 1943: 45 (characters, distribution, notes); Blackwelder,
1944: 100 (distribution); Herman, 2001b: 1791 (distribution).
Piestus (Piestus) fulvipes: Bernhauer & Schubert, 1910: 7 (distribution);
Scheerpeltz, 1952: 288 (characters, distribution).
Type material
Not examined. Syntype deposited in BMNH, sex undetermined ‘One
example in the British Museum from the Chevrolat collection. Collected
by F. de L’Herminier’ (Blackwelder, 1943: 46). Note: ‘One syntype labelled
as Type by D. Sharp from the Chevrolat collection’, with label: ‘Piestus
fulvipes Er. Type ex. coll. Chevrolat. Guadeloupe L’Herminier.’ and
deposited in BMNH (Dr Roger Booth, BMNH, pers. comm.). In the
original description Erichson (1840) did not specify how many specimens
he observed.
Distribution
West Indies (Herman 2001b).
Revision of Piestus with remarks on related genera
Biological notes
Unknown (Blackwelder 1943).
Remarks
All specimens previously identified in collections as P. fulvipes
were considered as P. penicillatus. Blackwelder (1943)
redescribed P. fulvipes and P. penicillatus, noting a little
difference on pronotum integument between the two species.
As the type material of P. fulvipes was not observed in the
current study, the species is placed as uncertain status, but we
suggest that it is a junior synonym of P. penicillatus.
Acknowledgements
We would like to thank Sonia A. Casari (MZSP), José Ricardo M. Mermudes
(Universidade Federal do Rio de Janeiro, UFRJ, Brazil), Albino
M. Sakakibara (UFPR), Lucia M. de Almeida (UFPR) and three
anonymous reviewers for valuable suggestions on the manuscript. We are
also grateful to Margaret K. Thayer (FMNH), Torsten Dikow (FMNH), Dave
J. Clarke (FMNH) and Angélico Asenjo (UFPR) for providing helpful
comments that improved this research. We thank M. K. Thayer for the use
of the Microptics imaging system provided by US National Science
Foundation collaborative grant, AToL: Assembling the Beetle Tree of Life
(EF-0531768, subcontract 144–439) and the TAXon line – ‘Rede Paranaense
de Coleções Biológicas (UFPR) for the photos. This research was supported
by MCT/CNPq Proc.no. 477520/2007-3, 473847/2009-4 and ‘Conselho
Nacional de Desenvolvimento Científico e Tecnológico’. Finally, we are
grateful to curators and institutions listed for loan of types and other
specimens used in this study. This paper is part of the Ph.D. project of the
first author, and contribution n 1864 of the Departamento de Zoologia,
Universidade Federal do Paraná, Brazil.
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Appendix 1. List of characters
Seventy parsimony-informative characters were selected, 62 binary and 10 multistate
Head:
0. Front: (0) without pair of frontal processes (Fig. 73); (1) with pair of frontal processes (Fig. 72).
1. Length of the frontal processes in male: (0) equal or longer than the antennal scape (Fig. 65); (1) shorter than the antennal scape (Fig. 64).
2. Basal distance between each frontal process: (0) 2 times or more the basal width of each one (Fig. 66); (1) the same or narrower than the basal width of each one
(Fig. 67).
3. Front, without frontal process: (0) slightly ventrally deflected (Fig. 86); (1) suddenly ventrally deflected (Fig. 82).
4. Front in frontal view: (0) without a slight concavity at middle; (1) with a slight concavity at middle.
5. V-shaped frontal sulcus: (0) incomplete (Fig. 86); (1) complete (Fig. 82).
6. Vertex: (0) without transverse carina on base (Fig. 86); (1) with transverse carina on base (Fig. 82).
7. Punctures: (0) fine and inconspicuous (Fig. 82); (1) fine and conspicuous (Fig. 86); (2) large (Fig. 89).
8. Moderate punctures with seta on basal half of the dorsal margin of the eyes: (0) one; (1) two; (2) three (in line); (3) five or more (in line).
9. Antennal length of male: (0) equal or slightly longer than female; (1) conspicuously longer than female (Figs 48, 49, 106, 107).
10. Antennal length of male: (0) short, reaching or exceeding a little the apex of elytra (Fig. 13); (1) long, almost reaching or reaching the apex of abdomen (Fig. 17).
11. Long setae on basal half of exposed dorsal face of the antennal scape: (0) absent; (1) present only in male (Figs 106, 107); (2) present in male and female.
12. Enlarged area on basal half of exposed dorsal face of the antennal scape: (0) absent; (1) present only in male (Figs 106, 107); (2) present in male and female.
13. Antennomere 4: (0) with dispersed microsetae, not densely covered; (1) densely covered with microsetae (Fig. 106).
14. Antenna (antennomere 4 to 11): (0) the same width from the base to the apex (Fig. 18); (1) gradually narrowing to the apex (Fig. 35).
15. Epipharynx as visible dorsally: (0) short, about the same or up to 2 times the median length of labrum (Fig. 108); (1) long, ~2.5 times or so the median length of
labrum (Fig. 110).
16. Epipharynx as visible dorsally: (0) with one lobe at each side (Fig. 108); (1) with two lobes at each side (Fig. 111).
17. Mandibles: (0) weakly projected anteriorly, shorter than or equal to the antennal scape (Fig. 26); (1) strongly projected anteriorly, longer than the antennal scape
(Fig. 25).
18. Dorsal tooth of the mandibles: (0) absent (Fig. 126); (1) present (Fig. 125).
19. Mandibles with dorsal tooth: (0) weakly developed, or developed but not forming bifurcate apex with first ventral tooth (Figs 113, 115, 123); (1) developed,
forming bifurcate apex with first ventral tooth (Figs 121, 122).
20. Mandibles with bifurcate apex: (0) dorsal tooth shorter than the ventral tooth (Fig. 121); (1) dorsal tooth longer than the ventral tooth (Fig. 120).
21. Teeth on internal borders of the mandibles (left–right): (0) 0–0 (Fig. 126); (1) 1–1 or 1–2, right mandible with an area that resembles a tooth (Figs 125, 129); (2)
1–2, right mandible with the apical tooth acute and not close to the other (Fig. 121); (3) 2–2, basal teeth projected anteriorly (Fig. 127); (4) 1–0.
22. Openings of glands at the external margin of mandibles: (0) absent; (1) present (Fig. 114).
23. Small basal tooth on external margin of the mandibles: (0) absent (Fig. 128); (1) present (Fig. 113).
24. Small basal tooth on external margin of the mandibles: (0) weakly acute (Fig. 121); (1) strongly acute (Fig. 113).
25. Maxillary palpus: (0) palpomere 4 equal to or a little longer than 2 and 3 combined (Fig. 131); (1) palpomere 4 shorter than 2 and 3 combined (Fig. 130).
26. Maxillary palpomere 3: (0) wider than long (Fig. 132); (1) as wide as long (Fig. 131).
27. Mentum: (0) rectangular, 2 times as wide as long (Fig. 136); (1) subquadrate, 1.5 times as wide as long or as wide as long (Fig. 135).
28. Anterior angles of the mentum: (0) not emarginate (Fig. 135); (1) emarginate (Fig. 136).
29. Pair of conspicuous setae at the apex of the median sclerotised plate of the ligula: (0) absent (Fig. 133); (1) present (Fig. 137).
30. Pair of conspicuous setae on apical margin of the ligula, near the median sclerotised plate: (0) absent (Fig. 133); (1) present (Fig. 137).
31. Gular sutures: (0) joined; (1) narrowly separated (Fig. 8); (2) broadly separated.
Thorax:
32. Anterior angles of the pronotum: (0) not or weakly anteriorly projected (Fig. 61); (1) anteriorly projected (Fig. 65).
33. Sides of the pronotum, anterior to constriction: (0) curved (Fig. 87); (1) subparallel to parallel (Fig. 83); (2) sinuous, with short teeth (Fig. 98).
34. Punctures on pronotum: (0) fine and inconspicuous, sometimes with a few scattered moderate punctures (Fig. 83); (1) fine and conspicuous, sometimes with a
few large punctures (Fig. 86); (2) only large punctures (Fig. 91).
35. Median longitudinal sulcus on pronotum: (0) inconspicuous (Fig. 92); (1) conspicuous (Fig. 82).
36. Basal constriction of the pronotum: (0) present on basal half or third (Figs 7, 85); (1) present on basal quarter (Fig. 82).
37. Slight emargination in each side, anterior to basal constriction of the pronotum (in lateral view): (0) absent; (1) present (Fig. 65).
38. Pronotum with lateral tooth anterior to the constriction: (0) absent (Fig. 61); (1) present (Fig. 7).
39. Anterior margin of the prosternum: (0) truncate; (1) anteriorly projected at the middle (Fig. 8).
40. Conspicuous ovate set of fine punctures in median region of prosternum of the male: (0) absent; (1) present (Fig. 8).
41. Mesoventrite process: (0) not carinate; (1) carinate (Fig. 9).
42. Moderate punctures on median region of metaventrite: (0) absent; (1) present.
43. Groove on ventral margin of the anterior femur: (0) absent; (1) present (Fig. 139).
44. Stout setae at the groove on ventral margin of the anterior femur: (0) short (Fig. 139); (1) long.
45. Small acute tooth on each basolateral angle of elytra: (0) absent (Fig. 56); (1) present (Fig. 52).
46. Complete longitudinal striae on each elytron: (0) one (Fig. 56); (1) three; (2) five (Fig. 16); (3) six (Fig. 5).
47. Stria 6 at the basal half on each elytron: (0) absent; (1) present (Fig. 5).
48. Epipleural carina of elytra: (0) absent; (1) present (Fig. 6).
49. Punctures on striae of elytra: (0) absent or inconspicuous (Fig. 16); (1) fine (Fig. 19); (2) large (Fig. 55).
50. Integument of striae of elytra: (0) not microgranulate; (1) microgranulate (Fig. 55).
(continued next page )
Revision of Piestus with remarks on related genera
Invertebrate Systematics
569
Appendix 1. (continued )
51. Integument of interstriae of elytra: (0) without undulate microstriae; (1) with undulate microstriae.
52. Interstriae of elytra: (0) not carinate (Fig. 86); (1) carinate (Fig. 55).
53. Apical margin of elytra: (0) truncate (Fig. 54); (1) projected (Fig. 53).
Abdomen:
54. Sternite 3: (0) without transverse carina (Fig. 143); (1) with transverse carina (Figs 10, 144).
55. Basolateral ridge on tergites 3–7: (0) absent; (1) present.
56. Paratergites on segments 3–6: (0) one pair; (1) two pairs.
57. Punctures on tergites: (0) uniformly distributed (Fig. 46); (1) only in anterolateral regions (Fig. 15).
58. Very small emargination on each posterolateral area on sternite 7, external openings of abdominal defensive gland complex (Caron et al. 2008): (0) absent; (1)
present (Fig. 11).
59. Lateral margin on sternite 7: (0) each without longitudinal set of long setae; (1) each with longitudinal set of long setae.
60. Apical margin of sternite 8 of female: (0) without short setae (Fig. 147); (1) with short setae (Fig. 148).
61. Apical margin of tergite 10: (0) without fringe (Fig. 149); (1) with fringe (Fig. 151).
62. Apex of tergite 10: (0) with 1 or 2 pairs of setae (Fig. 149); (1) with more than two pairs of setae (Fig. 151).
Genitalia:
63. Base of the median lobe of the aedeagus in ventral view: (0) not bulbous (Fig. 181); (1) bulbous (Fig. 214).
64. Apex of the median lobe of the aedeagus, in lateral view: (0) not curved (Fig. 188); (1) curved (Fig. 186).
65. Apex of the median lobe of the aedeagus, in ventral view: (0) not emarginated medially (Fig. 214); (1) emarginate medially (Fig. 181).
66. Region of median lobe for supporting the base of lateral lobe: (0) not extended anteriorly (Fig. 212); (1) extended anteriorly (Fig. 179).
67. Setae on apex of ovipositor of female: (0) many (Fig. 155); (1) few, one or two (Fig. 154).
68. External margin of ovipositor of female: (0) convexly curved (Fig. 153); (1) emarginate (Fig. 154).
69. Basal region of the spermathecal duct: (0) not sclerotised (Fig. 270); (1) sclerotised (Fig. 262).
570
Invertebrate Systematics
E. Caron et al.
Appendix 2.
Additional material examined
Piestus lacordairei Laporte, 1835
NICARAGUA: Río San Juan: 1 specimen, 60 km SE San Carlos Refugio Bartola, 1058.400 N 8420.300 W, 100 m, on bark downed logs, 25.V.2002, R. Brooks,
Z. Falin and S. Chatzimanolis coll. (SEMC). COSTA RICA: 1 specimen, Hamburgfarm, 30.I.[19]26, Nevermann coll. (FMNH); 1 specimen, the same locality
and collector, 4.II.1936 (FMNH); 1 specimen, no locality and collector, VII.[19]38 (FMNH); 3 specimens, Vara Blanca 2000 m, VIII.[19]38, no collector
(FMNH); 2 specimens, no locality, 1.V.1940, ilegible coll. (FMNH); 1 specimen, ilegible locality, 3.VIII.39, no collector (FMNH); 1 specimen, ilegible locality,
23–28.I.1941, no collector (FMNH); 1 specimen, ilegible locality, 25.I.1941, no collector (FMNH); Guanacaste: 5 specimens, Cação Biological Station
10550 38’N 85270 7’W, 1050 m, under bark, 10.VII.2000, J. Ashe, R. Brooks and Z. Falin coll. (SEMC); 2 specimens, 8 km NE Sta. Elena Santa Elena Forest
Res. 1020.701’N 8447.899’W, 1640 m, 11–17.VI.2001, S. and J. Peck coll. (SEMC); Alajuela: 1 specimen, Sarchí, 1000 m, 17.X.[19]34, Nevermann coll.
(FMNH); 1 specimen, E.B. San Ramon R.B. San Ramon 27 km N and 8 km W San Ramon, 10130 3000 N 84350 3000 W, 850–950 m, wet premontane forest,
20.VI-6.VII.1999, R. Anderson coll. (SEMC); 1 specimen, Est. Biologica Alberto M. Brenes, 1013.020 N 84350 5100 W, 900–1000 m, 29.VI.–6.VII.1999, Johnson
coll. (FMNH); Heredia: 4 specimens, La Selva Biol. Station nr Puerto Viejo de Sararapiqui, 18.II.1985, L. Herman coll. (AMNH); San José: 1 specimen, Naranjo,
VII.1939, Uloya coll. (AMNH); 4 specimens, Naranjo, VII.[19]39, no collector (1 AMNH, 3 FMNH); 2 specimens, St. Maria Dota, 2.12.1994, H. Forster coll.
(NMW); Cartago: 1 specimen, 13–16.II.1939, no collector (FMNH); Limón: 1 specimen, Lasmercedes Santa Clara, 200–300 m, 12.VI.[19]28, F. Nevermann
coll. (FMNH); 1 specimen, Hamburg Farm, loose bark in cracks of wood, 22.11.1935, F. Nevermann coll. (FMNH); Puntarenas: 3 specimens, La Reserva de
Monteverde, ~4 km E Monteverde, 5000–5500 ft, 4.III.1985, L. Herman coll. (AMNH); 1 specimen, Wilson Botanical Garden Las Cruces Biol. Stn, 1200 m, 27.
V.1993, J. S. and A. K. Ashe coll. (SEMC); 2 specimens, UVITA, 3–22.11.1994, H. Forster coll. (NMW); 1 specimen, Corcovado National Park Sirena Stn.
upper Ollas Trail, 140 m, 8290 700 N 83340 3900 W, flight intercept trap, 24–28.VI.2000, Z.H. Falin coll. (SEMC); 1 specimen, Corcovado National Park Sirena
Stn upper Rio Claro Trail, 8280 2900 N 83350 800 W, 100 m, flight intercept trap, 28.VI-1.VII.2000, Z. Falin coll. (SEMC); 1 specimen, Rincon de Osa 50 m,
841.141’N 8331.117’W, 23–26.VI.2001, S. and J. Peck coll. (SEMC); 1 specimen, Las Cruces Biol. Sta., 0847.140 N 8257.580 W, 1330 m, flight intercept
trap, 28–31.V.2004, J. S. Ashe, Z. Falin and I. Hinojosa coll. (SEMC). PANAMA: 4 specimens, Canal Zone Barro Colorado I., under bark, 14.I.1959, H. S. Dybas
coll. (FMNH); 6 specimens, the same locality and collector, fermented fibrous log and at light, 16.I.1959 (FMNH); 4 specimens, the same locality and collector,
Fairchild Trail, bark and under bark debris from fallen tree, 28.I.1959 (FMNH); 2 specimens, Canal Zone 4 mi. NW Gamboa, under bark, 23.II.1975, J. F. Lawrence
coll. (FMNH); 11 specimens, Canal Zone Barro Colorado Is., under bark fermenting, 16–22.II.1976, A. Newton coll. (3 DZUP, 8 FMNH); 2 specimens, the
same locality and collector, under bark fermenting logs, 25.II.1976 (FMNH); Bocas del Toro: 1 specimen, Almirante dam on Nigua Creek, berlese, 23.III.1959,
H. S. Dybas coll. (FMNH); 3 specimens, the same locality and collector, berlese, 25.III.1959 (FMNH); 1 specimen, the same locality and collector, berlese,
27.III.1959 (FMNH); Chiriquí: 1 specimen, La Fortuna 1200 m, Continental Divide Trail, 8460 0’N 82120 0’W, berlese forest litter, 9.VI.1995, R. Anderson coll.
(SEMC); 1 specimen, La Fortuna Cont. Divide Trail, 08460 N 82120 W, 1150 m, under bark, 9.VI.1995, J. Ashe and R. Brooks coll. (SEMC); 1 specimen, La
Fortuna Río Hornito Trail, 08420 N 82140 W, 1000 m, under bark, 11.VI.1995, the same collector (SEMC); Veraguas: 2 specimens, Cerro Santa Rita, chips
from cut end of log, 9.II.1959, H. S. Dybas coll. (FMNH); Colón: 1 specimen, Parque. Nac. Soberania Pipeline Rd km 61, 09070 N 79450 W, 40 m, flight intercept
trap, 7–21.VI.1995, J. Ashe, R. Brooks coll. (SEMC). COLOMBIA: Cundinamarca: 4 specimens, Finca San Pablo 3 km N Alban, 1800 m, 1–12.VIII.1967,
no collector (AMNH); Huila: 1 specimen, La Plata, no date and collector (NMW). FRENCH GUIANA: Saint-Laurent-du-Maroni: 5 specimens, Saül, under
bark, III.-IV.1999, A. Berkov coll. (AMNH); Cayenne: 3 specimens, no locality, date and collector (1 IRSNB, 2 ZMHB); 1 specimen, Roura 12.3 km SSE,
4380 55’N 52180 4’W, 260 m, under bark, 29.V.1997, J. Ashe and R. Brooks coll. (SEMC); 2 specimens, Roura 55.4 km SSE Kaw marshes, 4290 58’N 5230 0’W,
40 m, under fermenting bark, 11.VI.1997, the same collector (SEMC). ECUADOR: 1 specimen, no locality, date and collector (FMNH); Esmeraldas: 1
specimen, Cachabé low c., XI.[19]96, Rosenberg coll. (FMNH); 4 specimens, Cachabé low c., I.[19]97, Rosenberg coll. (2 FMNH, 2 IRSNB); 1 specimen,
Cachabé low c., XII.?, Rosenberg coll. (IRSNB); Pichincha: 1 specimen, 45 km NNW Quito Macquipucuna Station, 1600–1650 m, flight intercept trap, 18.IV–5.
V.1996, P. Hibbs coll. (SEMC); 1 specimen, Nanegalito 7 km S on Nono Road, 000 23’N 78400 36’W, 1540 m, pyrethrun fogging allen tree trunk, 30.X.1999, Z.
H. Falin coll. (SEMC); Napo: 1 specimen, Sierra Azul Hacienda Aragon 0400 0’S 77550 0’W, 2300 m, flight intercept trap, 17.II–26.III.1996, P. Hibbs coll.
(SEMC); Cotopaxi: 1 specimen, Naranjito nr San Francisco de la Pampas 1500ft., cane-sugar, 18.V.1993, J.J. Morrone, E. Tapia coll. (AMNH); 1 specimen, N of
San Francisco de las Pampas vic. Rio Esmeraldas, 4400–5000 ft., litter and debris, 14–15.V.[19]/93, L. Herman coll. (AMNH). PERU: Ucayali: 1 specimen,
Boqueron del Padre Abad, 470 m, 8.VIII.1946, no collector (AMNH); Cusco: 1 specimen, Consuelo Manu rd. km 165, rotten palm and leaf litter, 1.X.1982, L.E.
Watrous, G. Mazurek coll. (FMNH); 1 specimen, Marcapata, no date and collector (IRSNB). BRAZIL: Pará: 1 specimen, no locality, date and collector
(IRSNB); Rondônia: 2 specimens, Cacaulândia, II.1997, Vaz Mello coll. (AMBC). Undetermined country: 1 specimen, no locality, date and collector
(IRSNB).
Piestus capricornis Laporte, 1835
PERU: Loreto: 2 specimens, Napo River 80 mi NE of Iquitos, 7.XII.1980, T. King coll. (FMNH). BRAZIL: Amazonas: 1 specimen, Fonte Boa Estr.
Mamopina 023227S 660408W, 21–24.IX.2005, J.A. Rafael, F.F. Xavier Fo coll. (INPA); Pará: 3 specimens, Benevides, no date and collector (IRSNB);
Rondônia: 2 specimens, Porto Velho Campus UNIR, 08500 04’S 63560 35’W, 19.IV.2006, J.A. Rafael, F.F. Xavier Fo coll. (1 DZUP, 1 INPA); 5 specimens,
Vilhena, 12460 55’S 60220 18’W, 25.IV.2006, J.A. Rafael, F.F. Xavier Fo coll. (1 DZUP, 4 INPA). Undetermined country: 1 specimen, Cayenne, Guadalupe, no
date and collector (IRSNB); 1 specimen, no locality, date and collector (FMNH).
Piestus planatus (Sharp, 1887)
MEXICO: 1 specimen, no locality, date and collector (FMNH); 1 specimen, Sr. Durango, V.[19]05, Speyer coll. (ZMHB); Chiapas: 1 specimen, Palenque,
18.VII.1992, O. Hillert coll. (ZMHB); Veracruz: 1 specimen, Córdova, no date, J. Flohr coll. (ZMHB); Guerrero: 2 specimens, Atovac, no date, J. Flohr coll.
(ZMHB); Oaxaca: 2 specimens, no locality, date and collector (IRSNB); 2 specimens, 5.7 mi S Valle Nacional, 2000’, under bark hardwood, 11–18.VIII.1973, A.
Newton coll. (FMNH). GUATEMALA: 1 specimen, no locality, date and collector (IRSNB); Alta Verapaz: 1 specimen, Panzos, no date and collector (ZMHB).
NICARAGUA: Río San Juan: 11 specimens, 60 km SE San Carlos Refugio Bartola, 1058.400 N 8420.300 W, 100 m, on bark downned logs, 25.V.2002, R.
Brooks, Z. Falin and S. Chatzimanolis coll. (1 DZUP, 10 SEMC); 1 specimen, the same locality and collector, under bark, 27.V.2002 (SEMC). COSTA RICA: 1
specimen, no locality, date and collector (ZMHB); 1 specimen, Escocia, 7.IV.[19]24., Nevermann coll. (FMNH); Alajuela: 7 specimens, E.B. San Ramon
Revision of Piestus with remarks on related genera
Invertebrate Systematics
571
R.B. San Ramon 27 km N and 8 km W San Ramon, 10130 30’N 84350 30’W, 1120 m, rotting palm trunk, 29.VI–6.VII.1999, R. Anderson coll. (SEMC); Limón: 1
specimen, Hamburgfarm Reventazón, 14.X.[19]24, F. Nevermann coll. (FMNH); 1 specimen, the same locality and collector, no date (FMNH); 1 specimen, the
same locality and collector, 12.XI.1925 (FMNH); 1 specimen, the same locality and collector, 15.XI.1925 (AMNH); 1 specimen, the same locality and collector,
4.X.[19]28 (FMNH); 2 specimens, the same locality and collector, 6.VI.[19]30 (1 FMNH, 2 AMNH); 2 specimens, the same locality and collector, 3.XI.[19]33
(FMNH). PANAMA: Darién: 3 specimens, Cana ANCON Station, 0745.323’N 7741.069’W, 500 m, 3–9.VI.1996, S. Lingafelter coll. (1 DZUP, 2 SEMC); 4
specimens, Cana Biological Stn., Serranía de Pirre 7450 18’N 77410 6’W, 975 m, 4.VI.1996, A. Gillogly coll. (SEMC); 3 specimens, Cana ANCON Station,
07450 N 77410 W, 900–1100 m, 5.VI.1996, S. Lingafelter coll. (SEMC); 6 specimens, Cana Biological Station, 7450 18’N 77410 6’W, 530 m, 09.VI.1996, J.
Ashe, R. Brooks coll. (SEMC). ECUADOR: Esmeraldas: 1 specimen, Cachabé low c., no date, Rosenberg coll. (IRSNB); 1 specimen, Cachabé low c., XI.[19]
96, Rosenberg coll. (IRSNB). Undetermined country: 4 specimens, no locality, date and collector (ZMHB).
Piestus spinosus (Fabricius, 1801)
VENEZUELA: Bolívar: 1 specimen, El Pauchi bei Santa Elena, 01.III.1995, O. Hillert coll. (ZMHB). FRENCH GUIANA: Saint-Laurent-du-Maroni:
1 specimen, Juin, no date and collector (FMNH); 1 specimen, no locality, date and collector (AMNH); 1 specimen, Maripasoula Lawa River, under bark,
7.XII.1963, B. Malkin coll. (FMNH); 1 specimen, Saül under bark, III–IV.1999, A. Berkov coll. (AMNH); Cayenne: 1 specimen, Pariacabo, no date and
collector (IRSNB); 2 specimens, no locality, date and collector (ZMHB); 7 specimens, Roura 39.4 km SSE, 4320 43’N 5280 26’W, 270 m, under fermenting
bark, 10.VI.1997, J. Ashe, R. Brooks coll. (SEMC). ECUADOR: Napo: 1 specimen, no locality, date and collector (ZMHB); 2 specimens, 18 km S Tena,
28.IV.1978, C. W. and L. B. O’Brien and Marshall coll. (FMNH); Pastaza: 3 specimens, Cusuimi on Rio Cusuimi 150 km SE Puyo, 300 m, under bark,
19–23.VII.1971, B. Malkin coll. (FMNH). PERU: Loreto: 1 specimen, no locality, date and collector (ZMHB); Ucayali: 1 specimen, Boqueron del Padre
Abad, 470 m, 24.VIII.1946, F. Woytkowski coll. (AMNH); Huánuco: 1 specimen, TingoMaria, 13.VII.1968, C. W. and L. B. O’Brien coll. (AMNH); Lima:
1 specimen, Salinas, III.1939, no collector (NMW); Junín: 1 specimen, Rio Toro, no date and collector (ZMHB); Huánuco: 7 specimens, Panguana Sira,
12.IX.1988, Listabarth coll. (NMW); Cusco: 1 specimen, Consuelo Manu rd. km 165, rotten palm and leaf litter, 1.X.1982, L. E. Watrous and G. Mazurek
coll. (FMNH); 2 specimens, the same locality and collector, litter under crown of felled tree, 2.X.1982 (FMNH); 2 specimens, the same locality and collector,
rotten palm, 4.X.1982 (FMNH); 1 specimen, the same locality and collector, rotten palm, 5.X.1982 (FMNH); 2 specimens, the same locality and collector,
rotten palm, 6.X.1982 (FMNH); 1 specimen, the same locality and collector, flight intercept trap, 7.X.1982 (FMNH); 1 specimen, the same locality and
collector, beating palm branches, 8.X.1982 (FMNH); 2 specimens, the same locality and collector, under bark, 11.X.1982 (FMNH). BOLIVIA: 1 specimen,
Yuracaris, no date and collector (IRSNB); 2 specimens, Yuracaris, 6.XI.1900, no collector (1 FMNH, 1 NMW); Cochabamba: 2 specimens, El Chapa?,
X.1949, L. Peña coll. (FMNH); 1 specimen, Chapar, no date and collector (DZUP); Santa Cruz: 1 specimen, Loc. Parque Amboró, 17 39’S 63 43’W, bajo
corteza de Ficus sp., Rio Saguayo, no date and collector (FMNH); 1 specimen, Prov. Ichilo Loc. Parque Amboró, bajo corteza de Ficus sp., Rio Saguayo,
700 m, 26.XII.1988, no collector (FMNH). BRAZIL: 1 specimen, Marco de Legua, no date and collector (IRSNB); Amazonas: 1 specimen, Manaus, no
date and collector (NMW); 3 specimens, no locality, date and collector (1 NMW, 2 FMNH); 1 specimen, Ega, no date and collector (IRSNB); Pará: 1
specimen, no locality and date, E. Sefer coll. (MNRJ); 2 specimens, no locality, date and collector (1 MZSP, 1 FMNH); 10 specimens, Aldeia Aracu Igarapé
Gurupu-Umu Maranhão 50 km E of Caninde under bark, V.1963, B. Malkin coll. (FMNH); Rondônia: 4 specimens, Guaporé, 121605S 604230W, 327 m,
23.IV.2006, J.A. Rafael, F.F. Xavier Fo coll. (INPA); 2 specimens, Vilhena, 12460 55’S 60220 18’W, 25.IV.2006, J.A. Rafael, F.F. Xavier Focoll. (INPA);
Mato Grosso: 7 specimens, Vila Vera, 55300 12460 , X.1973, M. Alvarenga coll. (AMNH); 4 specimens, Sinop, 12310 55370 , X.1974, M. Alvarenga coll.
(AMNH). Undetermined country: 4 specimens, no locality, date and collector (1 NMW, 1 FMNH, 1 ZMHB, 1 ZMUC).
Piestus longipennis Fauvel, 1864
COLOMBIA: 1 specimen, no locality, date and collector (IRSNB); Valle del Cauca: 6 specimens, Cali, no date, Fass coll. (1 DZUP, 5 NMW); 1 specimen,
Carmen, 1400 m, 17.VI.1908, Fass coll. (FMNH); 1 specimen, Villa Elvira, 1800 m, 2.VII.1908, Fass coll. (FMNH); 1 specimen, Rio Fivaco, 2000 m, IX.1908,
Fass coll. (FMNH); 1 specimen, S. Antonio, 2000 m, XI.1908, Fass coll. (FMNH). ECUADOR: Carchi: 1 specimen, 46 km W Tufino west slope, 2600 m,
under dead wood bark, 19.XI.1987, C. Young and R. Davidson coll. (FMNH); Pichincha: 1 specimen, near Calicali, 1800 m, 19.XII.87, M. Huybensz coll.
(FMNH); Napo: 2 specimens, 52 km N. Tena, 6500ft., 24.V.1993, L. Herman coll. (AMNH); 1 specimen, Cosanga, 4.2 km S on Baeza-Tena Road then 2.9 km W
on pipeline access road, 2150 m, 0370 19’W 77500 1’W, on under bark downed logs, 7.XI.1999, Z. H. Falin coll. (SEMC); Cotopaxi: 1 specimen, Canton Sigchos
Las Pampas Otonga Natural Reserve, 25–28.VII.2005, W. Rossi coll. (DZUP); 3 specimens, Canton Sigchos Las Pampas Bosque Integral Otonga, 7–10.VII.06,
W. Rossi coll. (FMNH); 1 specimen, Bosque Integral Otonga, 1975 m, 79000 204’ 00250 166’, 02.06.[20]07, A.C. Proaño C. and A. Barragán coll. (DZUP);
Tungurahua: 1 specimen, Baños, no date and collector (ZMHB).
Piestus zisckai Scheerpeltz, 1951
ECUADOR: Tungurahua: 1 specimen, Santa Jnéz, no date and collector (ZMHB); Zamora-Chinchipe: 1 specimen, Rio Bombuscaro, 1100 m, 470 0’S
78370 48’W, malaise trap, 26.VI–4.VII.1996, P. Hibbs coll. (SEMC). PERU: 1 specimen, no locality, date and collector (ZMHB). BOLIVIA: 1 specimen,
Yuracaris, no date and collector (IRSNB).
Piestus bicornis (Olivier, 1811)
COSTA RICA: Cartago: 4 specimens, Turialba, 800 m, no date and collector (ZMHB); Limón: 2 specimens, Rio Barbilla, no date and collector (NMW);
1 specimen, Hamburgfarm Reventazon, 27.IX.[19]24, F. Nevermann coll. (FMNH); 1 specimen, Las Mercedes 200–300m Sta. Clara, 20.III.1922, the same
collector (FMNH). PANAMA: 7 specimens, Canal Zone, Madden Dam, under bark of hardwood ferment. stage, 12.VI.1976, A. Newton coll. (FMNH);
Panamá: 1 specimen, Ciricito Canal Zone, 16.I.[19]31, no collector (AMNH). COLOMBIA: 3 specimens, no locality, date and collector (1 ZMUC, 1
ZMHB, 1 IRSNB); Cundinamarca: 3 specimens, Bogota, no date and collector (ZMHB); Meta: 1 specimen, Villavicencio, 11.VII.[19]38, H. Dybas coll.
(FMNH); 5 specimens, Villavicencio Meta, 25.VII.[19]38, H. S. Dybas coll. (FMNH); Putumayo: 19 specimens, Santa Rosa headwaters of Rio San Miguel,
under bark of log, 16–20.X.1970, B. Malkin and P. Burchard coll. (FMNH). VENEZUELA: 1 specimen, Cafetal, no date and collector (FMNH); 3
specimens, Cafetal, V.1922, no collector (FMNH); Carabobo: 15 specimens, Las Trincheiras, VI.1922, L. R. Reynold coll. (FMNH). FRENCH GUIANA:
572
Invertebrate Systematics
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Saint-Laurent-du-Maroni: 1 specimen, 3.5mi N Saül, Les Eaux Claires, Treefall, 16.IV.[19]96 C. Chaboo coll. (AMNH); Cayenne: 2 specimens, no
locality, date and collector (NMW). ECUADOR: Sucumbios: 1 specimen, Sacha Lodge, 0.5S 76.5W, 270 m, malaise, 27.VIII–10.IX.1994, Hibbs coll.
(SEMC); Napo: 1 specimen, Tena, 14.II.1923, F.X. Williams coll. (FMNH); 1 specimen, Tena, 2100’, 3.XI.[19]88, L. Herman coll. (AMNH); Pastaza: 9
specimens, Cusuimi on Rio Cusuimi 150 km SE Puyo, 300 m, under bark, 19–23.VII.1971, B. Malkin coll. (FMNH); 8 specimens, Puyo nr. Santa Clara, 950
m, 18–21.VII.2008, W. Rossi and I. Tapia coll. (FMNH); Morona-Santiago: 1 specimen, Macas, no date and collector (FMNH). PERU: 5 specimens, no
locality, date and collector (1 ZMHB, 2 AMNH, 2 FMNH); Loreto: 1 specimen, 20 km from Ucayali on R.Calleria Colonia Calleria, 5.X–10.X.1961, B.
Malkin coll. (FMNH); 10 specimens, Yagua Indian Village headwaters of Loreto-Yacu, 2.V.1970, B. Malkin coll. (FMNH); San Martín: 7 specimens, Hera
(Jera) Moyobamba San Martin, 860 m, 27.VI.1947, F. Woytkowski coll. (AMNH); Huánuco: 1 specimen, no locality and collector, 15.V.[19]38 (NMW); 3
specimens, Panguana Sira, 12.IX.1988, Listabarth coll. (NMW); 2 specimens, Panguana Rio Pachitea, Rio Yuyapichis, 9370 S 74560 W, 260 m, 1–14.
IX.1968, Listabarth coll. (NMW); 3 specimens, Bella Durmiente near Tingo Maria, 14.VII.1968, C. W. and L. B. O’Brien coll. (AMNH); Junín: 2
specimens, no locality, date and collector (FMNH); 2 specimens, Chanchamajo, no date and collector (FMNH); 2 specimens, the same locality, 1000 m, no
date and collector (ZMHB); 2 specimens, Rio Toro, no date and collector (ZMHB); 4 specimens, San Ramón de Pangoa 40 km SE Satipo, 750 m, 24.
III.1972, R.T. and J.C. Schuh coll. (AMNH); Cusco: 1 specimen, Marcapata Perou Yungas Bolivie, 1000 m, no date and collector (IRSNB); 1 specimen,
Marcapata, no date and collector (IRSNB); 5 specimens, Callanga, no date and collector (2 NMW, 3 ZMHB); 2 specimens, Quincemil, 2400ft., 15.IV.1947,
J. C. Pallister coll. (AMNH); 3 specimens, Consuelo Manu rd. km 165, rotten palm, 3.X.1982, L. E. Watrous and G. Mazurek coll. (FMNH); 2 specimens, the
same locality and collector, rotten palm, 4.X.1982 (FMNH); 1 specimen, the same locality and collector, rotten palm, 5.X.1982 (FMNH); 1 specimen, the
same locality and collector, rotten palm, 5–15.X.1982 (FMNH); 1 specimen, the same locality and collector, rotten palm, 6.X.1982 (FMNH); 2 specimens,
the same locality and collector, beating palm branches, 8.X.1982 (FMNH); Madre de Dios: 2 specimens, Tambopata 15 km NE Puerto Maldonado Reserva
Cuzco Amazónico, 12330 S 69030 W, 200 m, under bark, 9.VII.1989, J. S. Ashe, R. A. Leschen (MUSM). BOLIVIA: 1 specimen, Yuracaris, no date and
collector (IRSNB); 1 specimen, VIII.1953, L. Peña coll. (FMNH); La Paz: 9 specimens, Yungas de la Paz, no date and collector (8 ZMHB, 1 FMNH); Beni:
1 specimen, Rurrenabaque, 23.X.1956, L. Pena coll. (FMNH); Cochabamba: 1 specimen, no locality, date and collector (DZUP); 11 specimens,
Cristalmayu, 28.VIII.1949, L. Peña coll. (NMW); 3 specimens, Chapare, 400 m, no date, R. Zischka coll. (1 NMW, 2 ZMHB); 7 specimens, 20 mi SW Villa
Tunari, 2.IV.1978, L. and C. W. O’Brien and G. B. Marshall coll. (FMNH); 2 specimens, 16.7 km W. Villan Tunari Avispas, 1710 13’S 65320 46’W, 500 m,
11.II.1999, R. Anderson coll. (SEMC); 5 specimens, 16.7 km W. Villa Tunari Parque Machias, 16580 20’S 65240 42’W, 300 m, 12.II.1999, R. Anderson
coll. (SEMC); Santa Cruz: 1 specimen, Prov. Ichilo Loc. Parque Amboró, 17390 S 63430 W, Rio Saguayo bajo corteza de Ficus sp., no date and collector
(AMNH); 1 specimen, Ichilo, 17390 S 63430 W, bajo corteza de Ficus sp., 31.VII.1988, no collector (FMNH); 3 specimens, Ichilo Cafetal, under bark of
stump, 6.VIII.1990, no collector (FMNH); 1 specimen, Potrerillo de Guenda, 2.IX.[19]98, no collector (FMNH). BRAZIL: 4 specimens, no locality, date
and collector (3 NMW, 1 IRSNB); 1 specimen, Pebas Ega, no date and collector (IRSNB); 1 specimen, Santa Rita, no date and collector (IRSNB); Acre: 1
specimen, Iquiri, VIII.[19]51, no collector (MZSP); Amazonas: 1 specimen, Manaus, Reserva Adolpho Ducke, 02550 5100 S 59580 5900 W, 13.III.2006, V.
Rodrigues coll. (INPA); Pará: 16 specimens, Caninde Rio Gurupi, under bark, 6.IV.1963, B. Malkin coll. (FMNH); Goiás: 1 specimen, Jatahy Rio Verde, no
date and collector (IRSNB); 1 specimen, Jatahy, no date and collector (FMNH); 1 specimen, Pires do Rio, 2.XI.1956, J. R. Pacheco coll. (DZUP); Minas
Gerais: 1 specimen, no date and collector (ZMHB); Espirito Santo: 1 specimen, no date and collector (IRSNB); 3 specimens, Corrego Itá, XI.1956, W.
Zikan coll. (MNRJ); Rio de Janeiro: 1 specimen, Neue Friburg., no date and collector (ZMHB); São Paulo: 1 specimen, no locality, date and collector
(MZSP); 2 specimens, Umgbg.v.Ribeirão Preto, III–VII.1899 (FMNH); Paraná: 1 specimen, Salto Mauá, V.[19]43, no collector (DZUP); 1 specimen,
Caviuna, XII.1945, A. Maller coll. (AMNH); 1 specimen, Rolandia, IV.1948, A. Maller coll. (AMNH); Santa Catarina: 2 specimens, Blumenau, no date
and collector (NMW); 1 specimen, N. Teutonia, no date, Plaumann coll. (FMNH); 1 specimen, no locality, date and collector (FMNH); 1 specimen, Corupá,
I.1938, A. Maller coll. (MNRJ); 1 specimen, no locality and collector, III.1940 (NMW); 25 specimens, Nova Teutonia, VII.41, Dirings coll. (3 DZUP, 22
MZSP); 1 specimen, Corupa, XI.1948, A. Maller coll. (AMNH). PARAGUAY: Guairá: 2 specimens, 300 m, 25.VII.1951, C. Pfannel coll. (FMNH).
Undetermined country: 10 specimens, no locality, date and collector (5 ZMHB, 2 NMW, 2 IRSNB, 1 FMNH).
Piestus fronticornis (Dalman, 1821)
NICARAGUA: 1 specimen, no locality, date and collector (IRSNB); Granada: 1 specimen, Res. Nat. Volcan Mombacho, 1150.050 N 8558.830 W, 1060
m, flight intercept trap, 1–5.VI.2002, R. Brooks, Z. Falin and S. Chatzimanolis coll. (SEMC); Río San Juan: 1 specimen, 60 km SE San Carlos Refugio
Bartola, 1058.400 N 8420.300 W, 100 m, on bark downed logs, 25.V.2002, R.Brooks, Z.Falin, and S.Chatzimanolis coll. (SEMC). GUATEMALA:
Guatemala: 2 specimens, no locality, 27.VI.1969, G. Ekis coll. (AMNH). COSTA RICA: 1 specimen, no locality, date and collector (ZMHB);
Guanacaste: 1 specimen, Puerto Humo, 17.VIII.[19]66, F. Fisk coll. (FMNH); Alajuela: 2 specimens, San Carlos, no date and collector (1 DZUP, 1
ZMHB); Heredia: 1 specimen, La Selva Biol. Statíon nr. Puerto Viejo de Sarapiqui, 18.II.1985, L. Herman coll. (AMNH); San José: 2 specimens, La Uruca,
1100 m, no date and collector (IRSNB); 29 specimens, no locality and date, E. Schmidt coll. (AMNH); 15 specimens, no locality, date and collector
(AMNH); 1 specimen, La Caja 8 Kil.W., 1931, Schmidt coll. (FMNH); 2 specimens, La Caja, 1934, no collector (1 AMNH, 1 FMNH); 2 specimens, no
locality and collector, 1936 (FMNH); 5 specimens, La Caja, II.1940, no collector (ZMHB); 3 specimens, no locality, VIII.[19]64, F. Fisk coll. (FMNH); 1
specimen, San Antonio de Escazu, malaise trap, 25.III–9.IV.1984, S.A. Cameron coll. (AMNH); 11 specimens, St. Maria Dota, 2.XII.1994, H. Forster coll.
(NMW); Limón: 1 specimen, 1000–1200 m, X.[19]28, F. Nevermann coll. (FMNH); 1 specimen, 4.X.[19]28, F. Nevermann coll. (FMNH); 1 specimen,
willted banana leaves, 25.XI.1928, F. Nevermann coll. (FMNH); 1 specimen, 1.V.1931, Nevermann coll. (FMNH); 1 specimen, 26.XI.[19]33, Nevermann
coll. (FMNH); 1 specimen, decaying banana blossoms leaves, 26.I.1935, F. Nevermann coll. (FMNH); Puntarenas: 21 specimens, UVITA, 3.XI.–22.
XI.1994, H. Forster coll. (NMW); 1 specimen, Corcovado National Park Sirena Stn. lower Ollas Trail, 5 m, fogging rotting sap flow, 24.VI.2000, Z. H. Falin
coll. (SEMC); 4 specimens, Corcovado National Park Sirena Stn. lower Ollas Trail 5m 8240 4800 N 83350 2200 W under bark, 25.VI.2000, the same collector
(SEMC). PANAMA: 1 specimen, no locality, date and collector (IRSNB); 1 specimen, Canal Zone Barro Colorado I., 13.I.1959, H.S. Dybas coll. (DZUP); 4
specimens, the same locality and collector, 14.I.1959 (1 DZUP, 3 FMNH); 4 specimens, the same locality and collector, fermented fibrous log at light, 16.
I.1959 (FMNH); 1 specimen, the same locality and collector, 21.I.1959 (FMNH); 2 specimens, Canal Zone, Madden Dam, under bark of hardwood ferment
stage, 12.VI.1976, A. Newton coll. (FMNH); 5 specimens, Canal Zone Barro Colorado Is., under bark fermenting, 16–22.II.1976, the same collector
(FMNH); 1 specimen, Canal Z. Barro Colo. Is., sift fermenting bark, 20.VIII.1978, Q. D. Wheeler (FMNH); 2 specimens, the same locality and collector, 26.
VIII.1978 (FMNH); Bocas del Toro: 1 specimen, Almirante trail to dam on Nigua Crk., decayed palm log, 29.III.1959, H. S. Dybas coll. (FMNH); Chiriquí:
2 specimens, V. de Chiriqui 25–4000ft., no date, Champion coll. (FMNH); 3 specimens, Bugaba, no date, the same collector (1 NMW, 1 AMNH, 1 ZMHB);
1 specimen, La Fortuna Cont. Divide Trail, 08460 N 82120 W, 1150 m, under bark, 9.VI.1995, J. Ashe and R. Brooks coll. (SEMC); Veraguas: 1 specimen,
Revision of Piestus with remarks on related genera
Invertebrate Systematics
573
no locality and collector, IX–X.1938 (FMNH); Colón: 1 specimen, no locality and collector, 28.I.1925 (FMNH); 1 specimen, Canal Zone 4mi. NW
Gamboa, under bark, 23.II.1975, J. F. Lawrence coll. (FMNH); 3 specimens, Black Tank Rd. NW Gatun Locks, under bark, 2.VI.1995, B. Ratcliffe and M.
Jameson coll. (SEMC); 1 specimen, 10–15 km N jct. Escobal and Piña Rds, ~30 m, under bark, 2.VI.1996, J. Ashe, R. Brooks coll. (SEMC); Panamá: 2
specimens, Ciricito Canal Zone, 13.III.[19]30, no collector (AMNH); 5 specimens, the same locality, 14.III.[19]30, no collector (AMNH); 2 specimens, the
same locality, 19.III.[19]30, no collector (AMNH); 1 specimen, the same locality, 28.II.[19]31, no collector (AMNH); Darién: 2 specimens, Cana Biological
Station, 530 m, 7450 N 77410 6’W, 09.VI.1996, J. Ashe and R. Brooks coll. (SEMC). COLOMBIA: 1 specimen, no locality, date and collector (ZMHB);
Cundinamarca: 1 specimen, Bogota, no date and collector (ZMHB); Valle del Cauca: 2 specimens, R. Dagua, IV.[18]97, W. Rosenberg coll. (IRSNB); 1
specimen, PNN Farallones de Cali Anchicaya, 3260 N 76480 W, 650 m, malaise, 19.XII.2000–02.I.2001, S. Sarria coll. (SEMC). FRENCH GUIANA:
Saint-Laurent-du-Maroni: 1 specimen, no locality, date and collector (FMNH); 1 specimen, Les Eaux Claires, 3. XI.1995, A. Berkov coll. (AMNH); 2
specimens, 3.5mi N Saül Les Eaux Claires, IV.[19]96, A. Berkov coll. (AMNH); 4 specimens, Saül 7 km N Las Eaux Claires 3390 4600 N 53130 1900 W, 220 m,
under bark, 1.VI.1997, J. Ashe and R. Brooks coll. (SEMC); 2 specimens, Saül, Lecythis zabucajo fallen branch, III–IV.1999, A. Berkov coll. (AMNH);
Cayenne: 3 specimens, no locality, date and collector (IRSNB). ECUADOR: Esmeraldas: 3 specimens, San Marco, 14.IX.1956 (FMNH); 2 specimens,
Zapallo Grande, 25–30.X.1987, M. Huybensz coll. (FMNH); Pastaza: 5 specimens, Puyo nr. Santa Clara, 950 m, 18–21.VII.2008, W. Rossi and I. Tapia
coll. (FMNH). BRAZIL: Amazonas: 1 specimen, Manaus, XI.1961, R. Arlé coll. (INPA); 1 specimen, Est. Br 17 km 34, 7.VIII.[19]69, E.V. Silva coll.
(INPA); 1 specimen, 35 km NE Manaus Res. Flor. Ducke, 25.VII–08.VIII.[19]95, Arndt and Gröger coll. (ZMHB); Amapá: 1 specimen, Serra do Navio,
IX.1957, K. Lenko coll. (MZSP); Pará: 1 specimen, no locality, date and collector (IRSNB); 2 specimens, Taperinha Santarem, 1–10.VII.[19]27, Zerny coll.
(NMW); 2 specimens, Tiriós Alto Parú d’Oeste, I–II.1963, Machado, Pereira coll. (MZSP); 6 specimens, Caninde Rio Gurupi, under bark, 6.IV.1963, B.
Malkin coll. (FMNH). Undetermined country: 1 specimen, no locality, date and collector (NMW).
Piestus validus Sharp, 1876
COLOMBIA: Meta: 1 specimen, Villavicencio, 25.VII.[19]88, H. S. Dybas coll. (FMNH). VENEZUELA: Carabobo: 2 specimens, San Esteban Ven. nr.
Puerto Cabello, 1–20.XII.1939, P. J. Anduze coll. (FMNH); Aragua: 3 specimens, Rancho Grande Biol. Stn., 10210 000 N 67410 000 W, 1300 m, flight intercept
trap, 12–14.V.1998, J. Ashe, R. Brooks and R. Hanley coll. (SEMC); 1 specimen, the same locality and collector, flight intercept trap, 14.V–2.VI.1998
(SEMC). ECUADOR: 1 specimen, Macas, no date and collector (FMNH); 2 specimens, no locality, date and collector (FMNH); 2 specimens, Libertad,
12.V.[19]63, L. Peña coll. (FMNH); 2 specimens, 8 km NE Puyo, 28.IV.1978, C. W. and L. B. O’Brien and Marshall coll. (FMNH); 1 specimen, Mera,
25.I.[19]23, F.X. Williams coll.; Napo: 4 specimens, Coca, V.1965, L. Pena coll. (FMNH); Pastaza: 1 specimen, Pomona on Rio Pastaza, 2000ft.,
9.VIII.1960, T. Sabine coll. (FMNH); 1 specimen, Cusuimi on Rio Cusuimi 150 km SE Puyo, 300 m, under bark, 19–23.VII.1971, B. Malkin coll. (FMNH);
Loja: 1 specimen, Pompeya, 13–25.V.1965, L. Pena coll. (FMNH). PERU: Junín: 1 specimen, Chanchamayo, no date and collector (FMNH); Huánuco:
1 specimen, Chinchao 25 km. Below Carpish, 2500 m, 6.IX.1946, F. Woykowski coll. (AMNH); 1 specimen, Panguana Sira, 12.IX.1988, Listabarth coll.
(NMW); 1 specimen, Panguana Río Pachitea Rio Yuyapichis, 9370 S 74560 W, 260 m, VI.1986, Listabarth coll. (NMW); Cusco: 1 specimen, Marcapata, no
date and collector (IRSNB); 9 specimens, Callanga, no date and collector (NMW); 3 specimens, Consuelo Manu rd. km 165, rotten palm, 3.X.1982,
L.E. Watrous and G. Mazurek coll. (1 DZUP, 2 FMNH); 7 specimens, the same locality and collector, rotten palm, 4.X.1982 (1 DZUP, 6 FMNH);
8 specimens, the same locality and collector, 5–15.X.1982 (1 DZUP, 7 FMNH); 5 specimens, the same locality and collector, rotten palm, 6.X.1982
(FMNH); 7 specimens, the same locality and collector, beating dead branches, 8.X.1982 (FMNH); 4 specimens, the same locality and collector, palm litter,
11.X.1982 (FMNH); 4 specimens, the same locality and collector, rotten palm, 12.X.1982 (FMNH); Madre de Dios: 2 specimens, Tambopata, 25.X.1982,
L. E. Watrous and G. Mazurek coll. (FMNH). BOLIVIA: 3 specimens, the same locality and collector (1 ZMHB, 2 FMNH); 2 specimens, Yuracaris, 6.
XI.1900, no collector (1 NMW, 1 FMNH); 5 specimens, Yuracaris, no date and collector (1 IRSNB, 4 NMW); 1 specimen, Yuracaris, S. Antonio, no date
and collector (IRSNB); La Paz: 1 specimen, Yungas de la Paz, no date and collector (ZMHB); 1 specimen, Yungas de La Paz 1000 m, no date, H. Rolle
coll. (ZMHB); 1 specimen, Yungas de la Paz, no date, V. Heyne coll. (ZMHB); Cochabamba: 4 specimens, Cochabamba 109 km E Yungas, CochabambaVila Tunari Rd., 1780 5000 S 65420 2900 W, 1480 m, flight intercept trap, 1–6.II.1999, F. Genier coll. (SEMC); 1 specimen, Cochabamba 109 km E Yungas,
Cochabamba-Vila Tunari Rd., 1780 5200 S 65420 5400 W, 1400 m, flight intercept trap, 8–12.II.1999, F. Genier (SEMC). BRAZIL: 4 specimens, Pebas, no date
and collector (1 FMNH, 3 IRSNB). Undetermined country: 5 specimens, no locality, date and collector (1 NMW, 4 ZMHB).
Piestus pennicornis Fauvel, 1864
NICARAGUA: Matagalpa: 1 specimen, 6 km N Matagalpa, Selva Negra Hotel, 1259.990 N 8554.530 W, 1350 m, under bark, 21.V.2002, R. Brooks, Z.
Falin and S. Chatzimanolis coll. (SEMC); Granada: 1 specimen, Res. Nat. Volcan Mombacho, 1150.050 N 8558.830 W, 800–1000 m, under bark and
upright tree, 2.VI.2002, R. Brooks, Z. Falin and S. Chatzimanolis coll. (SEMC). PANAMA: Colón: 1 specimen, Fort Sherman, 15.I.1980, Stockwell coll.
(FMNH); Veraguas: 9 specimens, Cerro Tule, 4 km W of Santa Fe, 850 m, 30.VII.1995, A. Gillogly coll. (SEMC). COLOMBIA: 2 specimens, Bogota, no
date and collector (1 ZMHB, 1 FMNH). VENEZUELA: 1 specimen, Caracas, no date and collector (IRSNB); 3 specimens, Las Trincheras, VI.1922, L. R.
Reynold coll. (FMNH). ECUADOR: Sucumbios: 2 specimens, Sacha Lodge, 0280 1400 S 76270 3500 W, 270 m, fruit fall, 23.III.1999, R. Brooks coll.
(SEMC); Napo: 1 specimen, Cosanga, 4.2 km S on Baeza-Tena Road then 2.9 km W on pipeline access road, 0370 1900 S 77500 100 W, 2150 m, under bark, 6.
XI.1999, Z. H. Falin coll. (SEMC); 1 specimen, the same locality and collector, 7.XI.1999 (SEMC). PERU: Cusco: 1 specimen, Marcapata, no date and
collector (IRSNB); 10 specimens, Consuelo, Manu rd km 165, rotten palm, 3.X.1982, L. E. Watrous and G. Mazurek coll. (1 DZUP, 9 FMNH); 1 specimen,
the same locality and collector, beating palm branches, 8.X.1982 (FMNH); 1 specimen, the same locality and collector, rotten palm, 11.X.1982 (DZUP).
BOLIVIA: 3 specimens, Yuracaris, no date and collector (IRSNB); Cochabamba: 1 specimen, 17 mi N Villa Tunari, 1.IV.1978, L. and C.W. O’Brien and
G.B. Marshall coll. (FMNH). BRAZIL: 3 specimens, Pebas, no date and collector (IRSNB); 1 specimen, no locality and date, A. Böttcher coll. (ZMHB); 2
specimens, no locality, date and collector (1 MNRJ, 1 FMNH); Rondônia: 1 specimen, Guaporé, 12160 0500 S 60420 3000 W, coleta manual em tronco, 23.
IV.2006, J.A. Rafael and F.F. Xavier Fo, coll. (INPA); Pará: 2 specimens, no locality, date and collector, ‘Ex-Typis’ (IRSNB); 1 specimen, no locality, date
and collector (FMNH); 4 specimens, Caninde, Rio Gurupi, under bark, 6.IV.1963, B. Malkin coll. (FMNH); Rio de Janeiro: 4 specimens, Nova Friburgo, no
date, L.W. Schaufuß coll. (ZMHB); 1 specimen, no locality and date, F. Sahlhberg coll. (FMNH); 3 specimens, no locality and collector, Fry coll. (2 FMNH,
1 ZMHB); 1 specimen, no locality, date and collector (FMNH); 2 specimens, Pétropolis, no date, Lfévrier coll. (IRSNB); 1 specimen, the same locality, no
date and collector (IRSNB); São Paulo: 6 specimens, no locality and date, M. Ráz coll. (5 FMNH, 1 NMW); 1 specimen, São Bernardo, IX.1932, J. Guerin
coll. (FMNH); Paraná: 1 specimen, Rolândia, XII.1947, A. Maller coll. (AMNH); 1 specimen, the same locality and collector, IX.1948 (AMNH); Santa
574
Invertebrate Systematics
E. Caron et al.
Catarina: 1 specimen, no locality and date, Klimsch coll. (FMNH); 2 specimens, Blumenau, no date and collector (NMW); 1 specimen, no locality, date
and collector (ZMHB); 1 specimen, Rio Vermelho, I.1946, A. Maller coll (AMNH); 1 specimen, Corupá, XI.1948, A. Maller coll. (AMNH). PARAGUAY:
Caazapá: 4 specimens, Hermosa, prop. Sosa family, San Rafael Reserve, 26190 1500 S 55440 5500 W, 90 m, fermenting tree wound, 4.XII.2000, Z.H. Falin
coll. (SMEC). Undetermined country: 1 specimen, no locality, date and collector (AMNH); 4 specimens, no locality and date, L.W. Schaufuß coll.
(ZMHB).
Piestus pygmaeus Laporte, 1835
MEXICO: 8 specimens, no locality, date and collector (3 ZMHB, 3 IRSNB, 1 FMNH, 1 NMW); 1 specimen, Tuxtepec, no date, Flohr coll. (ZMHB);
3 specimens, Almolonga, no date, Flohr coll. (ZMHB); 8 specimens, Jalapa, no date, W. Schaus coll. (AMNH); 5 specimens, no locality and date, J. Flohr
coll. (ZMHB); 4 specimens, Jalapa, no date, Hoege coll. (1 FMNH, 1 NMW, 2 ZMHB); 7 specimens, no locality and date, Truqui col. (FMNH); 4
specimens, no locality, 30.VI.1897, C. Hoge coll. (1 NMW, 3 AMNH); San Luís Potosí: 5 specimens, 12 mi S Cd. Mante, 21.VI.1975, L. E. Watrous coll.
(FMNH); Jalisco: 7 specimens, 1 Mi.S.W. La Resolana, 20.XI.1950, R. F. Smith coll. (AMNH); 1 specimen, Las Jarillas, 62 km S/P. Vallarta, under bark
and logs, 22.VII.1984, Chemsak and Doyen coll. (FMNH); Veracruz: 2 specimens, Las Vigas, no date, Hoege coll. (NMW); 2 specimens, Jalapa, no date
and collector (1 ZMHB, 1 AMNH); 1 specimen, Almolonga, no date and collector (ZMHB); 2 specimens, El Fortin, 20.VII.1936, C. H. Seevers coll.
(FMNH); 2 specimens, Tejeria, 4.VII.1944, H. Dybas coll. (FMNH); 1 specimen, Tierra Blanca, 28.VII.1944, H. Dybas coll. (FMNH); 1 specimen, 6 mi NE
Catemaco, rain for., 1500ft., under bark, 30.VII–8.VIII.1970, A. Newton coll. (FMNH); México: 2 specimens, Real de Arriba, Temescaltepec, 19.VI.[19]33,
H. E. Hinton and R. L. Usinger coll. (AMNH); Oaxaca: 4 specimens, Oaxaca, 4.5 mi S Valle Nacional, under bark hardwood, 13–16.VIII.1973, A. Newton
coll. (FMNH); 3 specimens, Oaxaca, 5.7 mi S Valle Nacional, under bark hardwood, 11–18.VIII.1973, A. Newton coll. (FMNH); 2 specimens, Tuxtepec, no
date and collector (ZMHB); Chiapas: 3 specimens, Cahabon, Vera Paz, no date, Champion coll. (1 AMNH, 1 NMW, 1 ZMHB); Nayarit: 1 specimen, El
Cora, Tepic, no date and collector (ZMHB). BELIZE: Orange Walk: 1 specimen, Rio Bravo Conservation Area, La Milpa Ruins, 17500 N 89020 W, FIT,
15–25.V.1997, C. Carlton coll. (SEMC); Belize: 8 specimens, Altun Ha, 11.VIII.1977, L. and C. W. O’Brien and G. B. Marshall coll. (FMNH); Toledo:
1 specimen, BARC San Pedro Columbia, 16160 4300 N 88570 4900 W, under bark at night, 25.IX.2004, P. W. Kovarik coll. (FMNH). GUATEMALA:
3 specimens, Zapote, no date, G.C. Champion coll. (1 ZMHB, 2 FMNH); Zacapa: 4 specimens, 24 km ENE Gualán, 280 m, under bark, 26.VI.1993, J. Ashe
and R. Brooks coll (SEMC); Escuintla: 1 specimen, Finca El Zapote, 2400ft., under bark, 6.VII.[19]48, R.D. Mitchell coll. (FMNH); 8 specimens, the same
locality and collector, 7.VII.[19]48 (FMNH); 1 specimen, the same locality and collector, 9.VII.[19]48 (FMNH); 5 specimens, the same locality and
collector, 10.VII.[19]48 (FMNH); 6 specimens, the same locality and collector, 11.VII.[19]48 (FMNH). HONDURAS: Santa Bárbara: 1 specimen, El
Mochilo, 13 km SE, 22.VII.1977, C. W. and L. O’Brien and Marshall coll. (FMNH); 1 specimen, Santa Barbara, La Fe, Finca La Roca, 5.3 km, S. Peña
Blanca, 14570 N 88020 W, 740 m, under bark, 21.VI.1994, Brooks and Ashe coll. (SEMC); Colón: 2 specimens, 3mi SW, Sonaguera, 23.VIII.1956, B. and
B. Valentine coll. (FMNH); Francisco Morazán: 1 specimen, Zamorano, 14N 87W, 820 m, rotting breadfruit, 6.VI.1994, J. Ashe and R. Brooks coll.
(SEMC). NICARAGUA: 1 specimen, no locality, date and collector (NMW); Granada: 20 specimens, Reserva Natural Volcan Mombacho, 1150.50 N
8600.740 W, 375 m, under fermenting guanacaste lumber, 1.VI.2002, R. Brooks, Z. Falin, and S. Chatzimanolis coll. (SEMC); 39 specimens, the same
locality and collector, 4.VI.2002 (SMEC); 3 specimens, Res. Nat. Volcan Mombacho, 1150.050 N 8558.830 W, 1200 m, under bark, 2.VI.2002, R. Brooks,
Z. Falin, and S. Chatzimanolis (SEMC); Río San Juan: 1 specimen, 60 km SE San Carlos, Refugio Bartola, 1058.400 N 8420.300 W, 100 m, on bark
downed logs, 25.V.2002, R.Brooks, Z. Falin and S. Chatzimanolis coll. (SEMC). COSTA RICA: 1 specimen, Costa Rica, no date, P. Biolley coll. (FMNH);
1 specimen, Rio Virilla, 26.XII.[19]31, H. Schmidt coll. (SEMC); 1 specimen, San Mateo, Macacona, no date and collector (IRSNB); Guanacaste: 1
specimen, S. Cañas, 9–14.II.1989, F.D. Parker coll. (SEMC); Heredia: 2 specimens, La Selva Biol. Station nr. Puerto Viejo de Sarapiqui, 18.II.1985,
L. Herman coll. (AMNH); Cartago: 1 specimen, Turialba, 800 m, no date, A. Heyne coll. (ZMHB); 2 specimens, Vulkan Turialba, 10 km N, no date and
collector (ZMHB); Limón: 1 specimen, Lasmercedes, Santa clara, 200–300 m, 17.X.[19]26, F. Nevermann coll. (FMNH); 1 specimen, Ramal Parisimina,
Santa Clara, 21.X.[19]26, the same collector (FMNH); 1 specimen, Hamburgfarm Reventazon Ebene Limon, 5.X.[19]28, the same collector (FMNH);
1 specimen, 31.X.[19]31, the same collector (FMNH); 1 specimen, Reventazon, Hamburg Farm, 27.IV.1934, the same collector (FMNH); 1 specimen, the
same locality and collector, on dry bark of Castilla, 21.VIII.[19]36 (FMNH); 6 specimens, the same locality and collector, in dry inner bark of Lecythis,
1.X.1936 (FMNH); 1 specimen, no localty and collector, VII.[19]38 (FMNH); Puntarenas: 8 specimens, UVITA, 3.XI–22.XI.1994, H. Forster coll.
(NMW); 2 specimens, Corcovado National Park, Sirena, logging recently splintered tree, 26.VI.2000, Z. H. Falin coll. (SMEC); 1 specimen, Corcovado
National Park, Sirena Stn., upper Rio Claro Trail, 8280 2900 N 83350 800 W, 100 m, flight intercept trap, 28.VI–1.VII.2000, Z. Falin coll. (SEMC); 14
specimens, Corcovado National Park, Sirena Stn., junc. Guanacaste-Sirena Trails, 8280 5500 N 83350 4600 W, 5 m, under bark, 30.VI.2000, Z. H. Falin coll.
(SEMC). PANAMA: 2 specimens, Almirante, IV.[19]43, no collector (FMNH); 12 specimens, Canal Zone, Barro Colorado I., under bark, 14.I.1959, H.S.
Dybas coll. (FMNH); 1 specimen, the same locality and collector, berlese, 25.I.1959 coll. (FMNH); 1 specimen, the same locality and collector, under bark,
27.I.1959 (FMNH); 4 specimens, the same locality and collector, berlese, 28.I.1959 (FMNH); 10 specimens, the same locality and collector, under bark, 4.
IV.1959 (FMNH); 1 specimen, the same locality, under bark, 3.II.1976, A. Newton coll. (FMNH); 3 specimens, the same locality and collector, under bark,
8–25.II.1976 (FMNH); 2 specimens, the same locality and collector, 10.II.1976 (FMNH); 5 specimens, the same locality and collector, 16–22.II.1976
(FMNH); 1 specimen, the same locality and collector, 25.II.1976 (FMNH); 3 specimens, no locality, date and collector (FMNH); 1 specimen, the same
locality, fermenting bark, 26.VIII.1978, Q. D. Wheeler coll. (FMNH); Bocas del Toro: 1 specimen, Almirante, dam on Nigua Creek, berlese, 23.III.1959, H.
S. Dybas coll. (FMNH); 4 specimens, the same locality and collector, 25.III.1959 (FMNH); 1 specimen, the same locality and collector, 27.III.1959
(FMNH); Chiriquí: 1 specimen, no locality, VII.1930, David coll. (FMNH); 3 specimens, Chiriqui, no date and collector (ZMHB); 2 specimens, Pto
Armuelles, no date and collector (FMNH); Veraguas: 2 specimens, Gabina, VI.[19]38, no collector (FMNH); Colón: 1 specimen, Parque Nac. Soberania,
Pipeline Rd km 6.1, 09070 N 79450 W, 40 m, flight intercept trap, 7–21.VI.1995, J. Ashe and R. Brooks coll. (SEMC); 2 specimens, 10–15 km N jct. Escobal
and Piña Rds., 30 m, under bark, 02.VI.1996, J. Ashe and R. Brooks coll. (SEMC); Panamá: 2 specimens, Las Cumbres, 15.II.1959, H. S Dybas coll.
(FMNH); 2 specimens, Fort San Lorenzo, 20 m, 09000 N 79380 W, under bark, 15.V.1995, J. and A. Ashe coll. (SEMC); 3 specimens, Old Plantation Trail
km 4.0, Parque Soberania, 175 m, 1–2.VII.1995, A. Gillogly coll. (SEMC); 1 specimen, Old Plantation Trail km 3.5, Parque Soberania, 150 m, 4.VII.1995,
A. Gillogly coll. (SEMC). GUADELOUPE: 3 specimens, no locality, date and collector (1 NMW, 1 IRSNB, 1 AMNH); 2 specimens, Trois Rivieres, no
date, Dufau coll. (ZMHB); 1 specimen, no locality and date, Goubeyre coll. (AMNH). SAINT VINCENT AND THE GRENADINES: Saint Vincent: 1
specimen, Leeward side, no date, H. H. Smith (IRSNB). GRENADA: 2 specimens, Balthazar, Windward side, no date, H. H. Smith (1 NMW, 1 IRSNB).
TRINIDAD AND TOBAGO: Trinidad: 1 specimen, no locality and date, Thailer coll. (FMNH); 1 specimen, no locality, 17.XI.[19]08, Peetz coll.
(FMNH); 7 specimens, Balandra Bay, 19.IV.[19]22, L. R. Reynold coll. (FMNH); 5 specimens, the same locality and date, F. Psota coll. (FMNH); 1
Revision of Piestus with remarks on related genera
Invertebrate Systematics
575
specimen, the same locality, V.[19]22, L. R. Reynold coll. (FMNH); 10 specimens, the same locality and date, no collector (FMNH); 5 specimens, the same
locality, 23.V.[19]22, L. R. Reynold coll. (FMNH); 7 specimens, the same locality and date, F. Psota coll. (FMNH); 3 specimens, the same locality, 24.V.
[19]22, L. R. Reynold coll. (FMNH); 1 specimen, the same locality 13.X[19]34, no collector (FMNH); 1 specimen, Arima Vallley, 800–1200ft., 10–22.
II.1964, J. G. Rozen and P. Wygodzinsky coll. (AMNH). COLOMBIA: 7 specimens, no locality, date and collector (ZMHB); 1 specimen, R. Dagua, IX–
XII.[18]94, W. Rosenberg coll. (IRSNB); Magdalena: 1 specimen, Sierra Nevada de Santa Marta, Puente de los Claves, 15 km. E. Pueblo Bello, 500 m,
under bark, 13.VI.1968, B. Malkin coll. (FMNH); Cesar: 14 specimens, Puento de Los Clavos, 15 km E of Pueblo Bello, Sierra Nevada de Santa Marta, 500
m, under bark, 13.VI.1968, the same collector (FMNH); Boyacá: 1 specimen, Muzo, no date and collector (FMNH); Meta: 1 specimen, Villavicencio, 12.
VII.1938, H. Dybas coll. (FMNH); Putumayo: 1 specimen, Santa Rosa, headwaters of Rio San Miguel, under bark, 16–10.X.1970, B. Malkin and P.
Burchard coll. (FMNH). VENEZUELA: 1 specimen, no locality, 1858, Moritz coll. (NMW); 2 specimens, St. Theresa, 13.V.[19]22, F. Psota coll. (FMNH);
2 specimens, Las Trincheras, VI.[19]22, F. Psota coll. (FMNH); Bolívar: 3 specimens, between Sta. Elena de Uairen and Icabaró, 30.VII.1987, M. A. Ivie
coll. (SEMC). GUYANA: Demerara-Mahaica: 2 specimens, Georgetown, 41 km, SW Soesdyke rd. to Linden, 6280 2400 N 58130 4800 W, 15 m, under bark,
10.VI.2001, R. Brooks and Z. Falin coll. (SEMC); Potaro-Siparuni: 1 specimen, Tumatumari, no date and collector (AMNH). FRENCH GUIANA: SaintLaurent-du-Maroni: 2 specimens, Maripassoula, Lawa River, 7.XII.1963, under bark, B. Malkin coll. (FMNH); 2 specimens, Les Eaux Claires, 3.XI.1995,
A. Berkov coll. (AMNH); 3 specimens, 3.5mi N Saül, Les Eaux Claires, IV.[19]96, A. Berkov coll. (AMNH); Cayenne: 1 specimen, Chimbo, VIII.[18]97,
Rosenberg coll. (IRSNB). ECUADOR: Sucumbios: 1 specimen, Sacha Lodge, 0280 1400 S 76270 3500 W, 270 m, fungus covered log, 23.III.1999, R. Brooks
coll. (SEMC); Pichincha: 1 specimen, 10 km SE Sto.Domingo de los Colorados, 2.V.1978, O’Brien and Marshall coll. (FMNH); Chimborazo: 3 specimens,
Chimbo, VIII.[18]97, Rosemberg coll. (FMNH); Pastaza: 1 specimen, Ashuara indian village on Rio Macuma nr. Rio Morona, beating dry foliage, 11–16.
VII.1971, B. Malkin coll. (FMNH); 5 specimens, Cusuimi on Rio Cusuimi, 150 km SE Puyo, 300 m, under bark, 19–23.VII.1971, B. Malkin coll. (FMNH).
PERU: 6 specimens, no locality and collector, 23.V.1936 (3 AMNH, 3 FMNH); Loreto: 1 specimen, Estiron, Rio Ampiacu, under corticals, 15–22.V.1966,
B. Malkin coll. (FMNH); 1 specimen, 1.5 km N Teniente Lopez, 235.660 S 7606.920 W, 210–240 m, flight intercept trap, 18.VII.1993, R. Leschen coll.
(SEMC); Cusco: 1 specimen, Callanga, no date and collector (IRSNB); Junín: 1 specimen, Jauja Prov., Saní Beni 8 km E Satipo, 840 m, 6–9.XI.1935, F.
Woytkwski coll. (SEMC); 1 specimen, San Ramón de Pangoa, 40 km SE Satipo, 750 m, 24.II.1972, R. T. and J. C. Schuh coll. (AMNH). BOLIVIA: 1
specimen, Yuracaris, no date and collector (IRSNB); Beni: 1 specimen, Chacobo Indian, Village on Rio Benicito, 66–12200 18–27.VII.1960, B. Malkin
coll. (FMNH); Cochabamba: 1 specimen, Regian Chaparé, 400 m, 20.XII.1949, R. Zischka coll. (NMW); 1 specimen, 20 mi SW Villa Tunari, 2.IV.1978, L.
and C. W. O’Brien and G. B. Marshall coll. (FMNH); Santa Cruz: 11 specimens, Santa Cruz de la Sierra, 24–25.VIII.1960, B. Malkin coll. (FMNH); 24
specimens, the same locality and collector, 23–25.IX.1960 (FMNH); 1 specimen, 14 mi SW Porta Chuelo, uv light trap, 24–III.[19]78, G .B. Marshall coll.
(FMNH); 1 specimen, 10 mi W. Puerto Chuelo, uv light trap, 26.III.1978, Marshall coll. (FMNH); 16 specimens, Ichilo, Cafetal by Rio Quebrada
Palometilla, 5.VIII.1990, P. Parrillo and P. Bettella coll. (FMNH); 2 specimens, the same locality and collector, 6.VIII.1990 (FMNH); 1 specimen, 3.7 km
SSE Buena Vista, Hotel Flora Y Fauna, 400–1729.950 S 6333.150 W, 440 m, under bark, 5.XI.2002, R. Leschen (SEMC); 1 specimen, the same locality and
collector, at light, 6.XI.2002 (SEMC). BRAZIL: 2 specimens, Pebas, no date and collector (IRSNB); 13 specimens, no locality, date and collector (1 MNRJ,
5 ZMHB, 2 IRSNB, 5 FMNH); 1 specimen, Petropolis, II.1850, no collector (FMNH); 1 specimen, the same locality, no date, H. Schulz coll. (NMW); 1
specimen, Guarujá, II.[19]18, A. Bierig coll. (FMNH); Pará: 1 specimen, no locality, date and collector (IRSNB); 6 specimens, no locality and date, Baker
coll. (5 FMNH, 1 NMW); 1 specimen, Caninde, Rio Gurupi, Gurupi Uma Maranhao, 50 km E Caninde, under bark, 6.IV.1963, B. Malkin coll. (FMNH); 1
specimen, Tiriós, Alto Pará d’Oeste, I–II.1963, Machado and Pereira coll. (MZSP); 5 specimens, Caninde, Rio Gurupi, under bark, 6.IV.1963, B. Malkin coll.
(FMNH); 1 specimen, the same locality and collector, 7–11.IV.1963 (FMNH); 1 specimen, Aldeia Aracu, Igarape, GurupuUmu, Maranhao 50 km E of
Caninde, under bark, V.1963, B. Malkin coll. (FMNH); Ceará: 3 specimens, Serra Ibiapaba, Ubajara, VIII.1953, Ulisses coll. (AMBC); Pernambuco: 1
specimen, Pery-Pery, no date and collector (IRSNB); Bahia: 8 specimens, no locality, date and collector (1 IRSNB, 5 FMNH, 2 NMW); Mato Grosso: 7
specimens, Barra do Tapirape, under bark, 8.XI.1964, B. Malkin coll. (FMNH); Goiás: 1 specimen, S. Isabel do Morro, I. Bananal, VI.1961, M. Alvarenga
coll. (MNRJ); Minas Gerais: 4 specimens, Viçosa, IX.1943, Wygodzinsky coll. (MNRJ); Espirito Santo: 1 specimen, Rio Bonito Conde, 21.VI.[19]28,
Leopoldina coll. (ZMHB); Rio de Janeiro: 1 specimen, Tijuca, I.1857, J. Gray coll. (IRSNB); 1 specimen, Petrópolis, no date and collector (IRSNB); 12
specimens, no locality and date, Fry coll. (2 ZMHB, 10 FMNH); 4 specimens, no locality, 21.II.1932, D. Mendes coll. (MNRJ); 11 specimens, Trapicheiro,
30.IX.1961, H.Schubart coll. (MNRJ); 2 specimens, the same locality and collector, 10.XI.1961 (INPA); 1 specimen, Pq. da Cidade, VI.1995, A. Bello coll.
(AMBC); São Paulo: 2 specimens, no locality and date, Mráz coll. (FMNH); 2 specimens, Santos, no date, J. Metz coll. (1 FMNH, 1 ZMHB); 1 specimen,
the same locality, no date, Brauns coll. (NMW); 3 specimens, 27.IX.[19]21, A. Bierig coll. (FMNH); Santa Catarina: 12 specimens, no locality, date and
collector (1 FMNH, 1 MZSP, 2 DZUP, 3 NMW, 5 ZMHB); 2 specimens, no locality and date, Lüderwaldt coll. (1 FMNH, 1 NMW); 1 specimen, Blumenau,
no date and collector (NMW); 6 specimens, the same locality, Hetschko coll. (NMW); 1 specimen, no locality and date, Fry coll. (FMNH); 17 specimens, N.
Teutonia, no date, Plaumann coll. (FMNH); 5 specimens, the same locality and collector, II.1950 (FMNH); 3 specimens, Nova Teutonia, 27110 B 52230 L.,
no date, F. Plaumann coll. (2 FMNH, MNRJ); 2 specimens, the same locality and collector, VIII.1935 (MNRJ); 2 the same locality, no date and collector
(FMNH); 2 specimens, blumenau, no date, Hetschko coll. (NMW); 2 specimens, Nova Teutonia, VII.1935, B. Pohl coll. (MZSP); 1 specimen, Corupa,
XI.1948, Humbolt coll. (AMNH). PARAGUAY: 7 specimens, no locality, date and collector (1 IRSNB, 6 ZMHB); 1 specimen, no locality, 1885, Drake
coll. (FMNH); 3 specimens, no locality and date, Fiebrig coll. (ZMHB); 26 specimens, San Luis, no date, Reimoser coll. (NMW); Asunción: 11 specimens,
Asunción, no date, B. Podtiaguin coll. (AMNH); Paraguarí: 1 specimen, Sapucai, 25400 S 56550 W, 190 m, 18.VI.1994, U. Drechsel coll. (SEMC);
Guairá: 3 specimens, Villa Rica, Sta. Barbara, I.1926, Schade coll. (1 NMW, 2 DZUP); 1 specimen, Calle Florida, 11.IV.1992, U. Dreschel coll. (SEMC); 1
specimen, the same locality and collector, 17.IX.1994 (SEMC); Caazapá: 1 specimen, Hermosa, prop. Sosa family San Rafael Reserve, 26190 1500 S
55440 5500 W, 90 m, under bark, 5.XII.2000, Z. H. Falin coll. (SEMC); 2 specimens, the same locality, data e coletor, fermenting tree wound (SEMC);
Itapúa: 6 specimens, Hohenau, Alto Parana, no date, H. Jacob coll. (NMW); 1 specimen, the same locality and collector, 23.V.[19]32 (NMW); 1 specimen,
the same locality and collector V.1932 (FMNH). ARGENTINA: 1 specimen, no locality, date and collector (IRSNB); 1 specimen, Rio Parana, Territorio
das Missiones, no date and collector (FMNH); Salta: 3 specimens, Tabacal, VI.1993, F. Schade coll. (AMNH); Tucumán: 2 specimens, no locality, 17.
IX.1928, H. E. Box coll. (1 FMNH, 1 AMNH); 2 specimens, no locality, 23.X.1929, H. E. Box coll. (AMNH); Misiones: 3 specimens, no locality and date,
M. Richter coll. (FMNH); 1 specimen, Iguazu, luz, 20.II.1967, R. Foerster coll. (ZMHB); 1 specimen, 20 km SE Puerto Iguazu, 220 m, forest uv light, 6.
I.1991, S. and J. Peck coll. (FMNH). Undetermined country: 14 specimens, no locality, date and collector (2 NMW, 3 IRSNB, 1 FMNH, 8 ZMHB).
Piestus extimus Sharp, 1887
UNITED STATES OF AMERICA: Arizona: 2 specimens, Santa Rita Mts., 5 to 8000ft., no date, F. H. Snow coll. (SEMC); 2 specimens, St. Catalina Mts.,
Molino Basin, dead sotol, 14.III.1970, K. Stephan coll. (FMNH); 1 specimen, Santa Rita Mts., Madera Cyn., dead sotol, 15.III.1970, K. Stephan coll.
576
Invertebrate Systematics
E. Caron et al.
(FMNH); 3 specimens, Dragon Mts. Wood Cyn., 29.IV.1972, K. Stephan coll. (FMNH); 1 specimen, Proctor Ranch, Madera Cyn., Pima Co., rotting barrel
cactus, 17.III.1973, D. S. Chandler coll. (FMNH); 4 specimens, the same locality, dead sotol, IV.1973, R. Lenczy coll. (1 DZUP, 3 AMNH); 2 specimens,
Pima, IV.1973, R. Leczy coll. (FMNH); 4 specimens, PimaCo., Madera Cn., dead sotol, IV.1973, Lenczy coll. (AMNH); 1 specimen, Santa Cruz co., vic.
Pena Blanca Lk, 14.XI.1998, Morris and Warmer coll. (FMNH). MEXICO: 11 specimens, no locality, date and collector (2 FMNH, 4 IRSNB, 2 ZMHB,
3 NMW); 1 specimen, no locality, 30.VI.1897, C. Höge coll. (NMW); Durango: 3 specimens, 2 mi NW Nombre de Dios, 12.VII.1975, L. E. Watrous coll.
(FMNH); Nayarit: 1 specimen, Tepic, 2–7.VIII.1947, B. Malkin coll. (AMNH); 1 specimen, San Blas, 4–5.VIII.1947, the same collector (AMNH);
1 specimen, Tepic, no date and collector (FMNH); Jalisco: 16 specimens, 1 Mi.S.W.La Resolana, 20.XI.1950, R. F. Smith coll. (1 DZUP, 15 AMNH);
1 specimen, Las Jarillas, 62 km S, P. Vallarta, under bark, 22.VII.1984, Chemsak and Doyen coll. (FMNH); 1 specimen, Chamela Biol. Stn., flight intercept
trap, 14.VII.1989, R. W. Brooks coll. (SEMC); 2 specimens, Atoyac, Isla Chica, 1300 m, 27.X.1997, J. L. Navarrete coll. (CZUG); 2 specimens, Unión de
Tula, camino a La MARTINIQUE, 1500 m, 9.I.1998, H. Fierros and J. L. Navarrete-Heredia coll. (CZUC); 2 specimens, Casimiro Castillo, Amoyo
Tacubaya, 600 m, 11.VII.2005, H. E. Fierros-López (CZUG); México: 2 specimens, Tejupilco, Temescaltepec, 15.VI.[19]33, H. E. Hinton, R. I. Usinger
coll. (AMNH); 1 specimen, Real de Arriba, Temescaltepec, 19.VI.[19]33, H.E. Hinton, R. I. Usinger coll. (AMNH); Guanajuato: 1 specimen, Guanajuato,
no date and collector (IRSNB); Oaxaca: 2 specimens, Ocotlan, no date and collector (ZMHB); 3 specimens, Zaachila, 4500ft., 21.VII.[19]37, A. R. Mead
coll. (FMNH); 8 specimens, Cuicatlan, 27.VII.[19]37, M. Embary coll. (FMNH); 2 specimens, 2.1 mi NW Totolapam, 3500ft., on rotting cacti columnar
Opuntia, 6.IX.1973, A. Newton coll. (FMNH); 3 specimens, 14.0 mi E Chacalpa, Hwy 175, montane tropical, (‘Correction: should be 14.0 mi S Suchixtepec,
16050 N, 96280 W), 1700 m, under bark fermenting sap, 30.IV.1977, J. S. Ashe coll. (FMNH). Undetermined country: 2 specimens, no locality, date and
collector (NMW).
Piestus buquetii Fauvel, 1864
PERU: Cusco: 7 specimens, Torentoy Canyon base of Machu Picchu, 2000 m, under bark, 20.VI.1964, B. Malkin coll. (1 DZUP, 6 FMNH). BOLIVIA:
3 specimens, Yuracaris, no date and collector (IRSNB); 1 specimen, no locality and date, N. Hobngren coll. (NHRS); 1 specimen, Rio Beni, La Paz Reyes,
1891, no collector (FMNH); La Paz: 1 specimen, 9.4 km E Chulumani, Apa Apa Ecoll. Reserve, 1620.990 S 6730.300 W, 2100–2400 m, fungus covered log,
19.I.2001, J. S. Ashe and R. S. Hanley coll. (SEMC); 1 specimen, La Paz 6 km W Yanacachi, 1625.950 S 6747.070 W, 2150–2350 m, fungus covered log,
24.I.2001, J. S. Ashe and R. S. Hanley coll. (SEMC); 1 specimen, Coroico, 1611.900 S 6743.440 W, 1850 m, under bark, 27.I.2001, J. S. Ashe and
R. S. Hanley (SEMC); Beni: 1 specimen, Yuracaris, no date and collector (FMNH); 1 specimen, Yuracaris, 6.II.1900, no collector (FMNH).
Piestus filicornis Fauvel, 1902
BOLIVIA: Cochabamba: 1 specimen, Cochabamba, 105 km E Yungas, nr. Rio Carmen Mayu, Villa Tunari Rd., 1780 5100 S 65430 5000 W, 1750 m, flight
intercept trap, 1–6.II.1999, R. Hanley coll. (FMNH); 1 specimen, the same locality, 1–6.II.1999, F. Genier coll. (SEMC); 1 specimen, Cochabamba, 109 km
E Yungas, Villa Tunari Rd., 1780 5000 S 65420 2900 W, 1480 m, flight intercept trap, 1–6.II.1999, R. Hanley coll. (SEMC); 1 specimen, Cochabamba, 105 km E
Yungas, nr. Río Carmen Mayu, Villa Tunari Rd., 1780 5100 S 65430 5000 W, 1750 m, flight intercept trap, 8–12.II.1999, R. Hanley coll. (SEMC); 1 specimen,
Cochabamba, 109 km E, Yungas, Villa Tunari Rd., 1780 5200 S 65420 5400 W, 1400 m, flight intercept trap, 8–12.II.1999, F. Genier coll. (DZUP); 1 specimen,
the same locality and collector, 6.II–8.VI.1999 (SEMC).
Piestus niger Fauvel, 1864
MEXICO: 1 specimen, no locality and date, Reitter coll. (FMNH); 1 specimen, ilegible locality, no date, J. Flohr coll. (ZMHB); 1 specimen, no locality and
date, Koltze coll. (FMNH); Veracruz: 7 specimens, Jalapa, no date, Flohr coll. (2 DZUP, 4 ZMHB, 1 NMW); 1 specimen, Fortin, 26.X.[1]961, Pereira and
Halfter coll. (MZSP); Oaxaca: 1 specimen, 1590m Pueblo Nuevo, FIT, 1–12.VIII.1986, H. and A. Howden coll. (FMNH). GUATEMALA: 1 specimen, no
locality, date and collector (NMW); Guatemala: 2 specimens, Guatemala City, no date, Champion coll. (FMNH). COSTA RICA: Guanacaste: 3
specimens, Cacao Biological Station, 10550 3800 N 85270 700 W, 1050 m, under bark, 10.VII.2000, J. Ashe, R. Brooks and Z. Falin coll. (SEMC); 4 specimens,
the same locality and collector, 11.VII.2000 (SEMC). PANAMA: 1 specimen, no locality, date and collector (ZMHB).
Piestus minutus Erichson, 1840
MEXICO: Nayarit: 1 specimen, El Cora, Tepic, no date and collector (ZMHB); BELIZE: Orange Walk: 2 specimens, Rio Bravo Conserv. Area, La
Milpa transect, vic. La Milpa Archeol. Site, under Pseudobombax bark, 7–14.IX.1995, P. Kovarik coll. (FMNH); 1 specimen, Rio Bravo Conservation Area,
La Milpa Ruins, 17500 N 89020 W, FITs, 15–25.V.1997, C. Carlton coll. (SEMC); 1 specimen, RBCA La Milpa Field Stat., FIT, 15–25.V.1997, the same
collector (SEMC). GUATEMALA: 2 specimens, San Juan, Vera Paz, no date, Champion coll. (FMNH); Alta Verapaz: 1 specimen, Lanquin, 1000ft., under
bark, 4.VI.1948, R. D. Mitchell coll. (FMNH); Izabal: 2 specimens, Quiriqua Ruins, 8.II.1996, D. Brzoska coll. (SEMC); Escuintla: 1 specimen, Finca El
Zapote, 2400ft., under bark, 8.VII.[19]48, R. D. Mitchell coll. (FMNH); 2 specimens, the same locality and collector, 9.VII.[19]48 (FMNH); 1 specimen, the
same locality and collector, 10.VII.[19]48 (FMNH); 1 specimen, the same locality and collector, 13.VII.[19]48 (FMNH); 1 the same locality and collector,
2000ft., 20.VII.1948 (FMNH). EL SALVADOR: Usulatán: 1 specimen, 3mi. E. El Trinufo, 1200’, 9.VI.1974, O’Briens and Marshall coll. (FMNH).
HONDURAS: Atlántida: 2 specimens, Lancetilla Bot. Grd., Tela, 15460 N, 87270 W, 10 m, under bark, 23.VI.1994, J. Ashe and R. Brooks coll. (SEMC).
NICARAGUA: Granada: 2 specimens, Res. Nat. Volcan Mombacho, 910m 1150.050 N 8558.830 W, flight intercept trap, 1–5.VI.2002, R. Brooks, Z. Falin
and S. Chatzimanolis coll. (SEMC); Río San Juan: 1 specimen, 60 km SE San Carlos, Refugio Bartola, 1058.400 N 8420.300 W, 100 m, Heliconia fruits,
27.V.2002, R. Brooks, Z. Falin and S. Chatzimanolis coll. (SEMC); 1 specimen, the same locality and collector, 28.V.2002 (SEMC). COSTA RICA:
1 specimen, no locality and date, B. Haas coll. (FMNH); Guanacaste: 1 specimen, Lomas Barbudal Biological Reserve, 10300 2200 N 85220 1500 W, 17m, on
and under bark, 15.VII.2000, J. Ashe, R. Brooks and Z. Falin coll. (SEMC); Heredia: 3 specimens, La Selva Biol. Station nr. Puerto Viejo de Sarapiqui,
18.II.1985, L. Herman coll. (AMNH); 1 specimen, the same locality, 25.III.1988, H. A. Hespenheide coll. (SEMC); 8 specimens, 3 km S. Puerto Viejo, OTSLa Selva, 100 m, X.1992, P. Hanson coll. (SEMC); 1 specimen, La Selva Biol. Sta., 3 km S Pto. Viejo, 10260 N 84010 W, malaise trap, 3–8.VIII.1992, G.
Wright coll. (SEMC); 3 specimens, the same locality, IX.1993, P. Hanson coll. (SEMC); San José: 1 specimen, no locality and collector, 1936 (FMNH);
1 specimen, La Caja, 1942, no collector (FMNH); Cartago: 2 specimens, Turialba, 800 m, no date and collector (ZMHB); Puntarenas: 1 specimen,
Revision of Piestus with remarks on related genera
Invertebrate Systematics
577
Surrubres, 300’, no date, A. Heyne, (FMNH); 32 specimens, UVITA, 3.XI.–22.XI.1994, H. Forster coll. (NMW); 5 specimens, Corcovado National Park,
Sirena Stn. junc. Guanacaste-Sirena Trails, 8280 5500 N 83350 4600 W, 5 m, under bark, 30.VI.2000, Z. H. Falin coll. (SEMC). PANAMA: 20 specimens, Canal
Zone, Barro Colorado Island, bark and debris from fallen tree, 14.I.1959, H. S. Dybas coll. (FMNH); 29 specimens, the same locality and collector,
fermenting fibrous log at light, 16.I.1959 (FMNH); 21 specimens, the same locality, Fairchild Trail, bark and under bark debris from fallen tree, berlese, 28.
I.1959, H. S. Dybas coll. (FMNH); 21 specimens, the same locality and collector, Zetek Trail, fibrous debris in tree trunk, 4.IV.1959 (FMNH); 1 specimen,
the same locality, Albrook Forest Site, 100ft., 20.I.1968, R. Hutton coll. (AMNH); 3 specimens, the same locality, 4mi. NW Gamboa, under bark, 23.II.1975,
J.F. Lawrence coll. (FMNH); 2 specimens, the same locality, under bark rotting log, 3.II.1976, A. Newton coll. (FMNH); 1 specimen, the same locality and
collector, under bark rotting logs, 10.II.1976 (FMNH); 1 specimen, the same locality and collector, under bark, 8–25.II.1976 (FMNH); 3 specimens, the same
locality, 28.VII.1978, Q. D. Wheeler coll. (FMNH); 4 specimens, the same locality, under bark fermenting, 16–22.II.1976, A. Newton coll. (FMNH); 3
specimens, the same locality and collector, under bark rotting logs, 25.II.1976 (FMNH); 7 specimens, Canal Zone, Madden Dam, under bark, 12.VI.1976, A.
Newton coll. (FMNH); 1 specimen, Ft. Gulick, at lights, 18.XI.1978, H. J. Haslan coll. (FMNH); 1 specimen, Canal Zone, Gamboa-Pipeline Rd KM7, flight
intercept trap, 20.V–11.VI.1996, S. Lingafelter coll. (SEMC); Chiriquí: 2 specimens, Rio Armuelles, 1930, no collector (FMNH); 1 specimen, 10.5 km NE
Caldera, 08420 N 82190 W, 340 m, tree fall litter, 24.V.1995, J. and A. Ashe coll. (SEMC); Veraguas: 6 specimens, Gabina, IX.[19]38, no collector
(FMNH); 5 specimens, Cerro Santa Rita, chips from cut end of log, 9.II.1959, H. S. Dybas coll. (FMNH); Colón: 1 specimen, no locality, 28.I.1925, G.
Bateson coll. (FMNH); 1 specimen, Parque Nac. Soberania, Pipeline Rd. Km 0.8, 09070 N 79450 W, 40 m, collected in Heliconia mariae, 2.VI.1995, Ashe
and Brooks coll. (SEMC); 3 specimens, Parque Nac. Soberania, Pipeline Rd. Km 6.1, 09070 N 79450 W, 40 m, flight intercept trap, 31.V–2.VI.1995, the
same collector (SEMC); 1 specimen, 10–15 km N jct. Escobal and Piña Rds, ~30 m, under bark, 02.VI.1996, the same collector (SEMC); 2 specimens, Santa
Rita Ridge, 400m, 17 km E Transisthmus Hwy, 16.VI.1996, A. Gillogly coll. (SEMC); Panamá: 1 specimen, Círicito, Canal Zone, 13.III.[19]30, coll.
(AMNH); 24 specimens, Canal Zone, Las Cumbres, compost heap, 15.II.1959, H. S. Dybas coll. (FMNH); 2 specimens, Altos de Majé, sifting under tree
bark, 6–15.X.1975, D. S. Chandler coll. (FMNH); 1 specimen, Cerro Azul, ~2000’, wet debris small forest stream, 21.II.1976, A. Newton coll. (FMNH); 2
specimens, Cerro Trinidad 400 m, 6.I.1980, Stockwell coll. (FMNH); 2 specimens, Pipeline Rd., flight intercept trap, 16–21.VI.1993, S. Lingfelter coll.
(SEMC); 9 specimens, 09050 N 79400 W, Old Plantation Rd. 6.9 km, S. Gamboa, 80 m, fungusy log, 3.VI.1995, J. Ashe and R. Brooks coll. (SEMC); 7
specimens, Old Plantation Trail Km 4.0, Parque Soberania, 175 m, 1–2.VII.1995, A. Gillogly coll. (SEMC); 1 specimen, Barro Colorado Island, FIT, 911N
7951W, 1–5.VII.2000, S. Chatzimanolis coll. (SEMC); 2 specimens, Old Plantation Trail km 3.5, Parque Soberania, 150 m, 4.VII.1995, A. Gillogly coll.
(SEMC); Darién: 2 specimens, Rio Tuquesa, 8270 N 77440 W, 20 m, 18.II.1981, Stockwell coll. (FMNH); 1 specimen, Cana, 7480 N 77390 W, 450 m, 24.
IX.[19]82 Stockwell coll. (FMNH). DOMINICA: 1 specimen, Salibiaj, under rotten banana leaves on ground, 28.I.1968, B. Malkin coll. (FMNH).
GRENADA: 4 specimens, Balthazar, Windward side, no date, H. H. Smith coll. (3 NMW, 1 IRSNB). TRINIDAD AND TOBAGO: Trinidad: 1 specimen,
no locality, date ans collector (FMNH); 1 specimen, no locality, 1904, F. Birch coll. (FMNH); 76 specimens, Balandra Bay, IV.1922, F. Psota coll. (FMNH);
49 specimens, the same locality and collector, 23.V.[19]22 (FMNH); 29 specimens, the same locality and date, L. R. Reynold coll. (FMNH); 22 specimens,
the same locality and collector, 24.V.[19]22 (FMNH); 1 specimen, W.I. Trinidad, 6 Mi. NE Arima, L’Oranga Valley, 15.VI.1973, no collector (AMNH); 1
specimen, W.I. Trinidad, Asa Wright N.C., under bark, 8.VI.1991, D. Brzoska coll. (SEMC). COLOMBIA: 1 specimen, the same locality, data e coletor
(NMW); 1 specimen, Villavicencio, 11.VII.[19]38, H. S. Dybas coll. (FMNH); 3 specimens, Puento de Los Clavos, 15 km E of Pueblo Bello, Sierra Nevada
de Santa Marta, 500 m, under bark of log, 13.VI.1968, B. Malkin coll. (FMNH); Magdalena: 1 specimen, PNN Tayrona Cañaveral, 11200 N 7420 W, 30 m,
Malaise, 30.VIII.–19.IX.[20]00, R. Henriquez coll. (SEMC); 2 specimens, the same locality and collector, 50 m, Malaise, 29.IX–17.X.2000 (SEMC); 1
specimen, the same locality and collector, 2–22.I.2001, R. Henriquez coll. (SEMC); Amazonas: 1 specimen, PNN Amacayacu Matamata, 3410 S 70150 W,
150 m, Malaise, 04–28.VIII.2000, A. Parente coll. (SEMC); 1 specimen, the same locality and collector, 17.XII.2000–02.I.2001 (SEMC). VENEZUELA: 1
specimen, no locality, date and collector (IRSNB); 1 specimen, Carupapo, no date and collector (IRSNB); 2 specimens, no locality, 1858, D’Moritz (1
FMNH, 1 NMW); 1 specimen, Tropical Research Sta., Rancho Grande nr. Maracay, 13.IV.1942, no collector (AMNH); Aragua: 1 specimen, Parque Nac.
Henri Pittier 4 km N El Limon, 700 m, on large dead tree, 19.VI.1987, M. A. Ivie coll. (SEMC). GUYANA: 1 specimen, Regio 8, Iwokrama Forest, Turtle
Mt. base camp, 4430 500 N 58430 500 W, 50 m, under bark, 1.VI.2001, R. Brooks and Z. Falin (SEMC); 14 specimens, the same locality and collector, 4.
VI.2001 (SEMC); 2 specimens, Region 8 Iwokrama Forest, Kabocalli Field Stn, 4170 400 N 58300 3500 W, 60 m, under bark, 5.VI.2001, R. Brooks and Z.Falin
(SEMC). FRENCH GUIANA: 1 specimen, Riv. Lumier, no date and collector (IRSNB); Saint-Laurent-du-Maroni: 1 specimen, Les Eaux Claires, 3.
XI.1995, A. Berkov coll. (AMNH); 5 specimens, 3.5mi N Saül, Les Eaux Claires, IV.[19]96, the same collector (AMNH); 11 specimens, Saül, under bark,
III.–IV.1999, the same collector (AMNH); Cayenne: 1 specimen, Camopi, no date and collector (IRSNB); 3 specimens, Roura, 55.4 km SSE, Kaw marshes,
4290 5800 N 5230 000 W, 40 m, under fermenting bark, 11.VI.1997, J. Ashe and R. Brooks coll. (SEMC). ECUADOR: Sucumbios: 6 specimens, Sacha Lodge,
0.5S 76.5W, 270 m, malaise, 12–22.II.1994, Hibbs coll. (SEMC); 2 specimens, the same locality and collector, 4–14.III.1994 (SEMC); 3 specimens, the
same locality and collector, 14–24.III.1994 (SEMC); 2 specimens, the same locality and collector, 3–13.IV.1994 (SEMC); 3 specimens, the same locality
and collector, 24.V–3.VI.1994 (SEMC); 2 specimens, the same locality and collector, 13–23.VI.1994 (SEMC); 1 specimen, the same locality and collector,
13–25.VII.1994 (SEMC); 1 specimen, the same locality and collector, 25.VII–3.VIII.1994 (SEMC); 1 specimen, the same locality and collector,
10–21.X.1994 (SEMC); 3 specimens, Sacha Lodge, 0280 1400 S 76270 3500 W, 270 m, under bark, 21–24.III.1999, R. Brooks coll. (SEMC);
2 specimens, the same locality and collector, 22.III.1999 (SEMC); 1 specimen, the same locality and collector, 24.III.1999 (SEMC); Pichincha:
1 specimen, Rio Palenque Science Center, 0360 000 S 79210 000 W, 200 m, malaise trap, 25.III–6.VII.1996, P. Hibbs coll. (SEMC); Napo: 2 specimens, 18 km
S. Tena, 28.IV.1978, C. W. and L. B. O’Brien and Marshall coll. (FMNH); 3 specimens, Jatun Sacha Biol. Station, 21 km E. Puerto Napo, 400 m, virgin rain
forest, F.I.T, 15.VII.1994, Levy and Génier coll. (SEMC); 1 specimen, the same locality and collector, 18.VII.1994 (SEMC); 1 specimen, Yuturi Lodge, Rio
Napo, 0320 5400 S 7620 1800 W, 270 m, flight intercept trap, 20–21.III.1999, R. Brooks and D. Brzoska coll. (SEMC); Pastaza: 1 specimen, Pomona, on Rio
Pastaza, 2000ft., 9.VIII.1960, T. Sabine coll. (FMNH); 5 specimens, Cusuimi on Rio Cusuimi, 150 km SE Puyo, 300 m, under bark, 19–23.VII.1971, B.
Malkin coll. (FMNH). PERU: Cusco: 1 specimen, Callanga, no date and collector (IRSNB); Madre de Dios: 1 specimen, Tambopata, 15 km NE Pto.
Maldonado, 200 m, at light, 10.VI.1989. J. S. Ashe and R. A. Leschen coll. (SEMC); 1 specimen, the same locality, Maldonado, Reserva Cuzco Amazónica,
12330 S 69030 W, 200 m, 20.VI.1989, R.A. Leschen coll. (SEMC); 1 specimen, the same locality, fight intercept, 24.VI.1989, J. S. Ashe and R. A. Leschen
coll. (SEMC); 1 specimen, the same locality and collector, Liana fruit fall, 25.VI.1989 (SEMC); 1 specimen, the same locality, berlese in fruit fall, 25.
VI.1989, R. A. Leschen coll. (SEMC); 1 specimen, the same locality and collector, 1.VII.1989 (SEMC); 1 specimen, the same locality and collector, 9.
VII.1989 (SEMC); 1 specimen, the same locality and collector, Swamp trail, flight intercept trap, 16.VII.1989 (SEMC); 1 specimen, the same locality and
collector, 17.VII.1989 (SEMC); 1 specimen, the same locality and collector, 19.VII.1989 (SEMC); 1 specimen, Manu, Parque Nac. Manu, Zona Res., Rio
Manu, Cocha Juarez, trail nr. Manu, Lodge, flight intercept trap, 18–24.IX.1991, A. Hartman (FMNH); 1 specimen, Cocha Cashu Bio. Stn., Manu National
578
Invertebrate Systematics
E. Caron et al.
Park, 11530 4500 S 71240 2400 W, 350 m, flight intercept trap, 17–19.X.2000, R. Brooks coll. (SEMC); 1 specimen, the same locality, Cocha Salvador,
Reserved Zone, Manu National Park, 1200 1300 S 71310 3600 W, 310 m, flight intercept trap, 20–21.X.2000, R. Brooks coll. (SEMC); 3 specimens, Pantiacolla
Lodge, Monk Saki Trap, Alto Madre de Dios River, 12390 2200 S 71130 5500 W, 400 m, 25.X.2000, R. Brooks coll. (SEMC). BOLIVIA: 1 specimen,
Yuracaris, no date and collector (IRSNB); Beni: 3 specimens, Chacobo Indian, Village on Rio Benícito, 66–12200 , under bark of fallen log, 28–31.
VII.1960, B. Malkin coll. (FMNH); Cochabamba: 2 specimens, Cochabamba, 67.5 km NE, Est. Biol. Valle del Sajita, Univ. de San Simon,, 1760 3300 S
64470 5200 W, 300 m, flight intercept trap, 7–9.II.1999, F. Genier coll. (SEMC); 1 specimen, Cochabamba, 117 km E., Yungas Lagunitas, 1000 m, 1760 2200 S
65400 5700 W, flight intercept trap, 8–12.II.1999, F. Génier coll. (SEMC); 1 specimen, Cochabamba, 109 km E., Yungas, Cochabamba-Villa Tunari Rd., 1480
m, 1780 5000 S 65420 2900 W, flight intercept trap, 8–12.II.1999, R. Hanley coll. (SEMC); Santa Cruz: 1 specimen, Sta. Cruz de la Sierra, 23–25.IX.1960, B.
Malkin coll. (FMNH); 2 specimens, Ichilo, Cafetal, forest, under bark of log, 6.VIII.1990, P. Parrilo coll. (FMNH). BRAZIL: 1 specimen, no locality, date
and collector (NMW); 3 specimens, Pebas, no date and collector (IRSNB); Amazonas: 2 specimens, Manaus, no date and collector (NMW); 1 specimen, no
locality, date and collector (ZMHB); 1 specimen, Benjamin Constant, 18–28.IX.1962, K. Lenko coll. (MZSP); 1 specimen, Igarape Belem, nr. Rio Solimoes,
70 km E of Leticia, under bark, 18–28.V.1970, B. Malkin coll. (FMNH); Pará: 3 specimens, Caninde, Rio Gurupi, under bark, 6.IV.1963, B. Malkin coll.
(FMNH); 3 specimens, Aldeia Aracu, Igarape, Gurupu-Umu, Maranhao, 50 km E of Caninde, under bark, V.1963, B. Malkin coll. (FMNH); Rondônia: 1
specimen, Porto Velho, Campus UNIR, 08500 0400 S 68560 3500 W, 19.IV.2006, J. A. Rafael and F. F. Xavier Fo coll. (INPA); 4 specimens, Ouro Preto do
Oeste, Reserva INPA, Ceplac, 10430 0000 S 62140 4500 W, 22.IV.2006, J. A.Rafael and F. F. Xavier Fo coll. (INPA); Goiás: 1 specimen, Jatahy, no date and
collector (IRSNB); São Paulo: 3 specimens, no locality and date, B. Pohl coll. (AMNH); Santa Catarina: 16 specimens, Nova Teutonia, no date, Plaumann
coll. (10 FMNH, 3 AMNH, 3 NMW); 2 specimens, no locality and date, Klimsch coll. (1 FMNH, 1 NMW); 4 specimens, Nova Teutonia, VII.1935, B. Pohl
coll. (DZUP); 2 specimens, the same locality, 27110 B 52230 L, VIII.1935, F. Plaumann coll. (MNRJ); 12 specimens, the same locality, II.1950, F. Plaumann
coll. (FMNH); Rio Grande do Sul: 6 specimens, no locality, date and collector (5 FMNH, 1 AMNH). PARAGUAY: Guairá: 4 specimens, Villa Rica, no
date and collector (DZUP); 10 specimens, the same locality, VII.1922, A. Schade coll. (NMW). ARGENTINA: Tucumán: 1 specimen, no locality, 17.
IX.1928, H.E. Box coll. (AMNH); 7 specimens, no locality and the same collector, 23.X.1929 (AMNH). Undetermined country: 8 specimens, no locality,
date and collector (5 ZMHB, 2 NMW, 1 IRSNB); 1 specimen, Guatemala, Nicaragua, Grenada, Mexico, no date and collector (IRSNB).
Piestus penicillatus (Dalman, 1821)
MEXICO: 1 specimen, Sierra de Durango, no date and collector (IRSNB). CUBA: 1 specimen, no locality and date, Gundlach coll. (ZMHB); 1 specimen,
no locality and collector, B. Haas coll. (AMNH); 1 specimen, Havana, no date and collector (IRSNB); 12 specimens, the same locality, no date, Baker coll.
(2 FMNH, 4 ZMHB, 4 SEMC, 2 IRSNB); 1 specimen, Ant. de los Baños, 9.I.[19]28 (ZMHB); 1 specimen, N. Sierra Maestra, X.[19]28, Rambousek coll.
(FMNH); Pina del Río: 18 specimens, Aspiro, I.1933, A. Bierig coll. (FMNH); 24 specimens, the same locality and collector, 3.VI.1934 (2 DZUP,
22 FMNH); 1 specimen, the same locality and collector, XII.[19]34 (FMNH); 5 specimens, Sierra Rangel, XII.[19]34, the same collector (FMNH);
1 specimen, the same locality, X.1938, no collector (FMNH); La Habana: 8 specimens, Las Palmas, IX.[19]28, no collector (FMNH); 1 specimen, Baños,
9.I.[19]28, A. Bierig coll. (ZMHB); 11 specimens, Sierra Bonilla, 27.I.[19]29, the same collector (FMNH); 1 specimen, the same locality, 5.II.1933, A.
Bierig coll. (FMNH); 9 specimens, the same locality, Cueva del Indio, 20.X.1932, A. Bierig coll. (FMNH); 5 specimens, Sítio Perdido, Jaruco, 4.IV.1932, A.
Bierig coll. (FMNH); 3 specimens, Somorostro, Sra. de Tapaste, 14.VIII.1932, A. Bierig coll. (FMNH); Matanzas: 6 specimens, Catalina, VIII.1937, A.
Bierig coll. (FMNH); Santiago de Cuba: 1 specimen, no locality, date and collector (ZMUC); Guantánamo: 1 specimen, 22.III.1914, C.T. Ramsden coll.
(AMNH). DOMINICAN REPUBLIC: 3 specimens, St. Domingo, no date and collector (ZMHB, 2 IRSNB). PUERTO RICO: 3 specimens, no locality
and date, Moritz coll. (ZMHB); 3 specimens, no locality and collector, S.G. Krug coll. (ZMHB). GUADELOUPE: 2 specimens, no locality and date, B.
Haas coll. (1 AMNH, 1 FMNH); 7 specimens, Camp Jacob, no date and collector (IRSNB); 1 specimen, no locality and collector, 16.9.1903 (FMNH); 4
specimens, no locality, date and collector (1 FMNH, 2 NMW, 1 ZMHB). Undetermined country: 3 specimens, no locality, date and collector (NMW); 1
specimen, B. Haas coll. (FMNH); 1 specimen, Cuba, Jamaica, 5.X.[19]28, Rambousek coll. (ZMHB).
Piestus formicinus, sp. nov.
GUYANA: Upper Demerara-Berbice: 80 specimens, Region 8 Iwokrama Forest, 1 km W Kurupukari, Iwokrama Field Stn., 4400 1900 N, 58410 400 W, 60m,
ex: Acromyrmex hystrix refuse pile, 21.V.2001, R. Brooks and Z. Falin coll. (SEMC); 1 specimen, the same locality and collector, 2.VI.2001 (9 FMNH, 9
DZUP, 9 AMNH, 4 MZSP, 49 SEMC); 1 specimen, Region 8 Iwokrama Forest, Turtle Mt. base camp, 50m, 4430 500 N, 58430 500 W, ex: Acromyrmex hystrix
refuse pile, 31.V.2001, the same collector (SEMC); 2 specimens, the same locality and collector, 1.VI.2001 (SEMC); 5 specimens, Region 8 Iwokrama
Forest, Kabocalli Field Stn., 60m, 4170 400 N, 58300 3500 W, 4.VI.2001, the same collector (SEMC); 2 specimens, the same locality and collector, 5.VI.2001
(SEMC).
Piestus sulcatus Gravenhorst, 1806
NICARAGUA: Río San Juan: 1 specimen, 8 km SE El Castillo, Refugio Bartola, 1058.60 N 8420.40 W, 30 m, rainforest litter, 24–31.V.2002, R. Anderson
coll. (SEMC); 2 specimens, the same locality, 100m, flight intercept trap, 28–30.V.2002, R. Brooks, Z. Falin and S. Chatzimanolis (SEMC). COSTA RICA:
4 specimens, Hamburgfarm, 1.II.[19]24, Neverm coll. (FMNH); 1 specimen, the same locality and collector, 27.XI.1936 (FMNH); 14 specimens, Farm
Castilla, VI.[19]38, no collector (FMNH); 3 specimens, Finca Castilla, VI.[19]38, no collector (2 FMNH, 1 AMNH); Guanacaste: 1 specimen, Pitilla
Biological Station, 10580 000 N 85250 000 W, 600 m, berlese leaf litter, 2.V.1995, R. Anderson coll. (SEMC); Alajuela: 1 specimen, Barroeta, 4–8.III.1942, no
collector (FMNH); 1 specimen, Alajuela, Peñas Blancas, 800 m, flight intercept, 19.V.1989, J. Ashe, R. Brooks and R. Leschen (SEMC); Heredia:
1 specimen, F. La Selva, 3 km S. Pto. Viejo, 10260 N 84010 W, 20.III.1980, H. A. Hespenheide coll. (SEMC); 3 specimens, La Selva Biol. Station nr., Puerto
Viejo de Sarapiqui, 18.II.1985, L. Herman coll. (AMNH); 2 specimens, La Selva, 3.2 km SE Puerto Viejo, 100 m, flight intercept trap, 19.III.1992, W. Bell
coll. (SEMC); 1 specimen, the same locality and collector, 24.III.1992 (SEMC); 2 specimens, the same locality and collector, 27.III.1992 (SEMC);
2 specimens, 3 km S. Puerto Viejo, OTS-La Selva, 100 m, IX.1992, P. Hanson coll. (SEMC); 17 specimens, the same locality and collector, X.1992,
P. Hanson coll. (1 CZUG, 16 SEMC); 1 specimen, La Selva, 80 m, flight intercept trap, 19.IV.1993, J. S. and A. K. Ashe coll. (SEMC); 2 specimens, La
Selva Biol. Res. Sta., 3 km S. Puerto Viejo, 10250 000 N 8400 000 W, 80 m, flight intercept trap, 2–15.VI.1996, R. Hanley coll. (SEMC); 1 specimen, the same
locality and collector, 4–10.VI.1996 (SEMC); 1 specimen, the same locality and collector, 4–15.VI.1996 (SEMC); Cartago: 1 specimen, Turrialba, 15.
Revision of Piestus with remarks on related genera
Invertebrate Systematics
579
VII.1973, 600 m, V. O. Becker coll. (DZUP); Limón: 1 specimen, Finca Castilla, VI.[19]38, C. Rica coll. (FMNH); Puntarenas: 3 specimens, Osa Penn.,
Fundación Neotrop., 10 km W. Rincon, 8450 3000 N 83250 000 W, 20 m, berlese forest litter, 21.VI.1997, R. Anderson coll. (SEMC); 7 specimens, Rincon, 7
km W, Osa Penn., Est. F.N. Aguas Buenas, 50 m, flight intercept trap, 21–25.VI.1997, S. and J. Peck coll. (SEMC); 1 specimen, Osa Penn., Fundación
Neotrop., 10 km W. Rincon, 8420 3000 N 83310 3000 W, 20 m, berlese forest litter, 22.VI.1997, R. Anderson coll. (SEMC); 2 specimens, Corcovado National
Park, Sirena Stn., upper Rio Claro Trail, 8280 2900 N 83350 800 W, 100 m, flight intercept trap, 28.VI–1.VII.2000, Z. H. Falin coll. (SEMC); 7 specimens,
Rincon de Osa, 841.1410 N 8331.1170 W, 50 m, FIT, 23–26.VI.2001, S. and J. Peck coll. (SEMC); 6 specimens, Rincon de Osa, 841.1410 N 8331.1170 W,
150 m, FIT, 23–26.VI.2001, S. and J. Peck (SEMC); 5 specimens, Corcovado National Park, Sirena Stn., Corcovado Trail, 8290 700 N 83340 3900 W, 150 m,
flight intercept trap, 28.VI–1.VII.2000, Z.H. Falin coll. (SEMC); 1 specimen, Corcovado National Park, Sirena Stn., lower Rio Claro Trail, 8240 4800 N
83350 2200 W, 40 m, UV light, 30.VI.2000, Z. H. Falin coll. (SEMC); 1 specimen, the same locality and collector, FIT, 23–26.VI.2001, S. and J. Peck coll.
(SEMC); 1 specimen, Osa Penninsula, Fnd. Neotrop., 10 km W. Rincon, 08420 3000 N 83310 3000 W, 200 m, lowland for litter, 24.VI.2001, R. Anderson coll.
(SEMC). PANAMA: 1 specimen, no locality and date, 7 day old tapir dung, H. Dybas coll. (FMNH); 1 specimen, Almirante, IV.[19]43, no collector
(FMNH); 4 specimens, Canal Zone, Barro Colorado I., fermented fibrous log at light, 16.I.1959, H. Dybas coll. (FMNH); 6 specimens, Barro Colorado I., 1.
II.1959, 7 day old tapir dung, H. Dybas coll. (FMNH); 1 specimen, the same locality and collector, Zetek Trail, 4.IV.1959, fibrous debris in tree trunk, H. S.
Dybas coll. (FMNH); 1 specimen, the same locality, 6.II.1976, wet leaves and debris forest stream, A. Newton coll. (FMNH); 3 specimens, San Blas,
Nusagandi Reserve, 09210 N 78590 W, 350 m, rotting palm flrs., 17.V.1995, J. and A. Ashe coll. (SEMC); 3 specimens, Bocas d. Toro, Fortuna, Chiriquí
Grande road, 8470 N 82110 W, 800 m, premontane rain forest sifting litter, 14–16.VII.1987, D.M. Oison coll. (FMNH); Veraguas: 1 specimen, Gabina, IX.
[19]38, no collector (FMNH); Colón: 1 specimen, Parque Nac. Soberania, Pipeline Rd., 09070 N 79450 W, leaf litter, 29.V.1995, J. Jolly and C. Chaboo coll.
(SEMC); 1 specimen, the same locality, 40 m, flight intercept trap, 4–7.VI.1995, J. Ashe and R. Brooks coll. (SEMC); 1 specimen, the same locality and
collector, 7–21.VI.1995 (SEMC); 2 specimens, 15 km N jct. Escobal and Piña Rds., ~30 m, flight intercept trap, 02–11.VI.1996, J. Ashe and R. Brooks coll.
(SEMC); Panamá: 1 specimen, Las Cumbres, 24.VIII.1975, no collector (FMNH); 4 specimens, the same locality, 24.X.1975, D. S. Chandler coll. (FMNH);
1 specimen, Old Gamboa Rd., flight intercept trap, 13.VII.1993, D. Windsor coll. (SEMC); 2 specimens, the same locality and collector, 09040 N 79400 W,
14–19.XI.1994 (SEMC); 2 specimens, road to Cerro Campana, 8410 0100 N 79550 2100 W, 760 m, litter on forest floor and near trail, 1.I.[20]02, L. Herman coll.
(AMNH); Darién: 2 specimens, Cana Biological Station, 7450 1800 N 77410 600 W, 550 m, flight intercept trap, 03–07.VI.1996, J. Ashe and R. Brooks coll.
(SEMC); 1 specimen, the same locality and collector, 06.VI.1996 (SEMC); 1 specimen, the same locality and collector, 07.VI.1996 (SEMC).
GUADELOUPE: 6 specimens, Capesterre, III-25–61, Neufchateau coll. (AMNH). DOMINICA: Saint David: 14 specimens, Salibia, under rotten
banana leaves, 28–29.I.1968, B. Malkin coll. (FMNH). MARTINIQUE: 1 specimen, Fort de France, 9–12.VI.1960, P. and C. Vaurie coll. (AMNH). SAINT
VINCENT AND THE GRENADINES: Saint Vincent: 2 specimens, Leeward side, no date, H. H. Smith coll. (1 NMW, 1 IRSNB). TRINIDAD AND
TOBAGO: Trinidad: 1 specimen, Balandra Bay, IV.1922, no collector (FMNH); 12 specimens, the same locality and date, L. R. Reynold coll. (2 AMNH,
10 FMNH); 5 specimens, the same locality and date, F. Psota coll. (FMNH); 1 specimen, Balandra Bay, 23.V.[19]22, F. Psota coll. (FMNH); 4 specimens,
the same locality and date, L.R. Reynold coll. (FMNH); 2 specimens, Balandra Bay, 24.IV.[19]22, the same collector (FMNH); 1 specimen, Arima Valley,
800–1.200ft., 10–22.II.1964, Rozen and Wygodzinsky coll. (AMNH); 1 specimen, Asa Wright N.C., 15.I.1981, G.E. Bohart coll. (SEMC); 2 specimens,
Simla Res. Sta., 2–15.VI.1981, H. Clemons coll. (SEMC). COLOMBIA: Chocó: 2 specimens, Quebrada Docordo, between Cucurrupi and Noanama, Rio
San Juan, beating dry foliage, 1–5.I.1969, B. Malkin coll. (FMNH). VENEZUELA: Aragua: 8 specimens, 18.9 km S Choroni, 2900ft., litter near stream, 2.
III.1992, L. Herman coll. (AMNH); 24 specimens, Rancho Grande-Maracay rd, first stream SE of Rancho Grande, 3600ft., litter near stream, 1.IV.1992, the
same collector (AMNH); 16 specimens, 20 km NW Maracay, Rancho Grande, sendero a la Toma de Agua, 3800ft., litter near stream, 22.III[19]92, the same
collector (AMNH); 17 specimens, the same locality and collector, La Toma, 1150m, litter nr stream, 15.IV.1994 (AMNH); 1 specimen, Rancho Grande Biol.
Stn., 10210 N 67410 W, 1140m, flight intecept trap, 25–28.II.1995, R.W. Brooks coll. (SEMC); 4 specimens, the same locality and collector, 1–8.III.1995
(SEMC); 1 specimen, the same locality and collector, 1115 m, along stream, 9.III.1995 (SEMC). GUYANA: Upper Demerara-Berbice: 1 specimen,
Region 8, Iwokrama Forest, 1 km W Kurupukari, Iwokrama Field Stn., 4400 1900 N 58410 400 W, 60 m, flight intercept trap, 26–29.V.2001, R. Brooks and Z.
Falin coll. (SEMC). SURINAMEE: Marowijne: 2 specimens, Nassau Mountain, 4470 5300 N 54310 1600 W, 500 m, flight intercept trap, 1–2.VI.1999, Z.H.
Falin and B. Dedjin coll. (SEMC); 1 specimen, the same locality, 4480 5600 N 54360 2000 W, 480 m, flight intercept trap, 3–4.VI.1999, Z.H. Falin and B. Dedijn
coll. (SEMC); 2 specimens, Palumeu, 3200 5600 N 55260 1800 W, 160 m, flight intercept trap, 5–9.VII.1999, Z. H. Falin and D. Konoe coll. (SEMC); 1
specimen, the same locality, flight intercept trap, 7–8.VII.1999, Z. H. Falin coll. (SEMC). FRENCH GUIANA: 2 specimens, no locality, date and collector
(IRSNB); 1 specimen, Cunani, no date and collector (IRSNB); 3 specimens, Boeuf Mort, ~3380 N 53130 W, 160 m, 10.X.1995, L. Herman coll. (AMNH); 1
specimen, Wanaboo, near Nason, Marowjine River, 4430 3500 N 54260 3600 W, 40 m, flight intercept trap, 31.V–5.VI.1999, Z. H. Falin, B. Dedjin coll.
(SEMC); Saint-Laurent-du-Maroni: 8 specimens, nr. Eaux Claires, 3.5 mi N Saul, 338–400 N 5313–140 W, 155–260 m, leaflitter near stream, 5–13.X.[19]
95, L. H. Herman coll. (AMNH); 2 specimens, the same locality, data e coletor, fig. fruit fall (AMNH); 7 specimens, the same locality, data e coletor, leaf
litter and Eschweilera flower fall (AMNH); 3 specimens, Saül, 7 km N, 1 km NW, Les Eaux Claires, along Rue de Belizon trail, 3390 4600 N 53130 1900 W, 280
m, flight intercept trap, 4–8.VI.1997, J. Ashe and R. Brooks coll. (SEMC); 4 specimens, Saül, Lecythis zabucajo fallen branch, III–IV.1999, A. Berkov coll.
(AMNH); 3 specimens, Saül, sifting, IV.1999, A. Berkov coll. (AMNH); 1 specimen, Saül, 7 km N, Les Eaux Claires, 3390 4600 N 53130 1900 W, 220 m, flight
intercept trap, 30.V–4.VI.1997, J. Ashe and R. Brooks coll. (SEMC); 1 specimen, Saül, Mt. Galbao summit, 3370 1800 N 53160 4200 W, 740 m, rotting fruit, 6.
VI.1997, J. Ashe and R. Brooks coll. (SEMC); Cayenne: 1 specimen, no locality, date and collector (IRSNB); 2 specimens, Roura, 18.4 km SSE, 4360 3800 N
52130 2500 W, 240 m, flight intercept trap, 22–24.V.1997, J. Ashe and R. Brooks coll. (SEMC); 1 specimen, Roura, 8.4 km SSE, 4400 4100 N 52130 2500 W, 200
m, flight intercept trap, 22–24.V.1997, J. Ashe and R. Brooks coll. (SEMC); 4 specimens, Roura, 18.4 km SSE, 4360 3800 N 52130 2500 W, 240 m, flight
intercept trap, 25–29.V.1997, J. Ashe and R. Brooks coll. (SEMC); 1 specimen, Cayenne, 33.5 km S and 8.4 km NW of Hwy N2 on Hwy D5, 4480 1800 N
52280 4100 W, 30 m, flight intercept trap, 26–28.V.1997, J. Ashe and R. Brooks coll. (SEMC); 1 specimen, the same locality and collector, 29.V–9.VI.1997
(SEMC); 1 specimen, Matoury, 41.5 km SSW on Hwy N2, 4370 2200 N 52220 3500 W, 50 m, flight intercept trap, 29.V–9.VI.1997, the same collector (SEMC);
1 specimen, Roura, 18.4 km SSE, 4360 3800 N 52130 2500 W, 240 m, flight intercept trap, 29.V–10.VI.1997, J. Ashe and R. Brooks coll. (SEMC); 1 specimen,
Roura, 13.0 km SSE, 4380 3800 N, 52170 5600 W, 240 m, rotting fruit, 10.VI.1997, J. Ashe and R. Brooks coll. (SEMC); 1 specimen, Roura, 27.4 km SSE,
4440 2000 N 52130 2500 W, 280 m, flight intercept trap, 10.VI.1997, J. Ashe and R. Brooks coll. (SEMC). ECUADOR: Esmeraldas: 1 specimen, Bilsa,
0200 000 S 79430 000 W, flight intercept trap, 28.IV–10.V.1996, P. Hibbs coll. (SEMC); 2 specimens, the same locality and collector 10.V–5.VI.1996 (SEMC);
5 specimens, the same locality and collector, 5.VI–7.VII.1996 (SEMC); 1 specimen, the same locality and collector, 7–19.VII.1996 (SEMC); 2 specimens,
Bilsa Biological Station, 0200 2400 N 79420 3600 W, 500 m, flight intercept trap, 7–19.VII.1996, P. Hibbs coll. (SEMC); Sucumbios: 2 specimens, Sacha
Lodge, 0.5S 76.5W, 270 m, malaise, 22.II–4.III.1994, Hibbs coll. (SEMC); 1 specimen, the same locality and collector, 14–24.III.1994 (SEMC); 1
580
Invertebrate Systematics
E. Caron et al.
specimen, the same locality and collector, 23.IV–4.V.1994 (SEMC); 2 specimens, the same locality and collector, 24.V–3.VI.1994 (SEMC); 1 specimen, the
same locality and collector, 23.IV–3.VII.1994 (SEMC); 1 specimen, the same locality and collector, 3–13.VII.1994 (SEMC); 3 specimens, the same locality
and collector, 13–25.VII.1994 (SEMC); 6 specimens, the same locality and collector, 23.VII–3.VII.1994 (SEMC); 2 specimens, the same locality and
collector, 25.VII–3.VIII.1994 (SEMC); 2 specimens, the same locality and collector, 3–16.VIII.1994 (SEMC); 1 specimen, the same locality and collector,
16–29.VIII.1994 (SEMC); 2 specimens, the same locality and collector, 27.VIII–10.IX.1994 (SEMC); 1 specimen, the same locality and collector, 20–30.
IX.1994 (SEMC); 20 specimens, the same locality and collector, 21–24.III.1999 (SEMC); 2 specimens, Sacha Lodge, 0280 1400 S 76270 3500 W, 270 m,
Bignoniaceae, large fruits, 22.III.1999, R. Brooks coll.; Pichincha: 1 specimen, 16 km E Sto.Domingo, Tinalandia, 680 m, malaise rainforest, 4.V–25.
VII.1985, S. and J. Peck coll. (FMNH); 2 specimens, 17 km SE Sto. Domingo de Colorado, Tinalandia, 3000’, 16–21.X.[19]88, L. Herman coll. (AMNH); 1
specimen, W of Alluriquim, Tinalandia, 2600–2800ft., mercury vapor light 19–20.V.1993, L. Herman coll. (AMNH); 1 specimen, 45 km NNW Quito,
Macquipucuna Station, 1600–1650 m, flight intercept trap, 3–18.IV.1996, P. Hibbs coll. (SEMC); 3 specimens, Tinalandia, Santo Domingo, 16 km E,
0160 5300 S 7930 3900 W, 750 m, flight intercept trap, 26–27.III.1999, R. Brooks and D. Brzoska coll. (SEMC); Napo: 1 specimen, Jatun Sacha Biol. Station, 21
km E. Puerto Napo, 400 m, 8–11.VII.1994, F. Génier coll. (SEMC); 3 specimens, the same locality and collector, 9.VII.1994 (SEMC); 5 specimens, the same
locality and collector, 21.VII.1994 (SEMC); 2 specimens, the same locality and collector, 13.VII.1994 (SEMC); 3 specimens, the same locality, 18.VII.1994,
Levy and Génier coll. (SEMC); 1 specimen, km 7.3 Sarayacu Loreto Rd., 1200 m, cloud forest, leces trap, 20.VII.1994, F. Génier coll. (SEMC); 1 specimen,
Onkone Gare Camp, 00390 1000 S 76260 0000 W, 220 m, under leaf litter, 4–12.X.1995, T.L. Erwin, G.E. Ball and D. Shpeley coll. (SEMC); 2 specimens,
Sierra Azul, Hacienda Aragon, 0400 000 S 77550 000 W, 2300 m, flight intercept trap, 17.II–26.III.1996, P. Hibbs coll. (SEMC); 1 specimen, Yuturi Lodge, Rio
Napo, 0320 5400 S 7620 1800 W, 270 m, flight intercept trap, 20–21.III.1999, R. Brooks and D. Brzoska coll.; Cotopaxi: 1 specimen, Guasaganda, 500 m, 1.
IV.1988, M. Huybensz coll. (FMNH); Pastaza: 3 specimens, Puyo, nr. Santa Clara, 950 m, 18–21.VII.2008, W. Rossi and I. Tapia coll. (FMNH). PERU: 1
specimen, Tambopata Prov., 15 km NE Pto. Maldonado, 200 m, flight intercept, 3.VI.1989, J. Ashe and R. Leschen coll. (SEMC); 1 specimen, the same
locality and collector, at light, 10.VI.1989 (SEMC); 1 specimen, the same locality and collector, flight intercept, 22.VI.1989 (SEMC); 2 specimens, the same
locality and collector, 18.VII.1989 (SEMC); Loreto: 1 specimen, Campamento San Jacinto, 218.750 S 7551.770 W, 175–215 m, berlese, 2.VII.1993, R.
Leschen coll. (SEMC); 3 specimens, the same locality and collector, flight intercept trap, 3.VII.1993 (SEMC); 1 specimen, the same locality and collector, 7.
VII.1993 (SEMC); 4 specimens, 1.5 km N Teniente Lopez, 235.660 S 7606.920 W, 210–240 m, palmfruit berlese, 17.VII.1993, R. Leschen coll. (SEMC); 1
specimen, the same locality and collector, 20.VII.1993 (SEMC); 4 specimens, the same locality and collector, 21.VII.1993 (SEMC); 1 specimen, the same
locality and collector, 22.VII.1993 (SEMC); 1 specimen, the same locality and collector, 24.VII.1993 (SEMC); 1 specimen, the same locality and collector,
24.VII.1993 (SEMC); 3 specimens, the same locality and collector, 27.VII.1993 (SEMC); San Martín: 1 specimen, Prov. Huallaga, Tocache, 500 m, 10.
XI.1900, G. A. Baer coll. (IRSNB); Huancavelica: 1 specimen, Panguana, Rio Pachitea, Rio Yuyapichia, 9370 S 74560 W, 260 m, 1–14.IX.1986, Listabarch
coll. (NMW); Cusco: 1 specimen, Consuelo, Manu rd., km 165, 3.X.1982, L. E. Watrous and G. Mazurek coll. (FMNH); 1 specimen, the localidade and
collector, 6.X.1982 (FMNH); 1 specimen, the same locality and collector, 7–8.X.1982 (FMNH); 2 specimens, the same locality and collector, 8.X.1982
(FMNH); 1 specimen, the same locality and collector, 10.X.1982 (FMNH); Madre de Dios: 1 specimen, Tambopata, 25.X.1982, L. E. Watrous and G.
Mazurek coll. (FMNH); 1 specimen, the same locality and collector, 28.X.1982 (FMNH); 1 specimen, Tambopata Prov., 15 km NE Puerto, Maldonado
Reserva Cuzco Amazónico, 12330 S 69030 W, 200 m, 8.VI.1989, R. A. Leschen coll. (SEMC); 1 specimen, the same locality and collector, pitfall trap, 22.
VI.1989, J. S. Ashe and R. A. Leschen coll. (SEMC); 1 specimen, the same locality and collector, 25.VI.1989 (SEMC); 1 specimen, the same locality and
collector, 30.VI.1989 (SEMC); 1 specimen, the same locality and collector, 2.VII.1989 (MUSM); 1 specimen, Manu Prov., Parque Nac. Manu, Zona Res.,
Rio Manu, Cocha Juarez, trail nr. Manu, Lodge, flight intercept trap, 18–24.IX.1991, A. Hartman coll. (FMNH); 5 specimens, Cocha Cashu Bio. Stn., Manu
National Park, 350 m, 11530 4500 S 71240 2400 W, flight intercept trap, 17–19.X.2000, R. Brooks coll. (SEMC); 1 specimen, Cocha Salvador, Reserve Zone
Manu National Park, 310 m, 1200 1300 S 71310 3600 W, 20.X.2000, R. Brooks coll. (SEMC); 3 specimens, Pantiacolla Lodge, 8 km NW El Mirador Trail, Alto
Madre de Dios River, 800 m, flight intercept trap, 12380 3000 S 71160 4100 W, 23–26.X.2000, R. Brooks coll. (SEMC). BOLIVIA: 3 specimens, Yuracaris
(IRSNB); Cochabamba: 8 specimens, Cochabamba, 109 km E, Yungas, Cochabamba – Villa Tunari Rd., 1480 m, 1780 5000 S 65420 4900 W, flight intercept
trap, 1–6.II.1999, F. Génier coll. (SEMC); 2 specimens, the same locality, 8–12.II.1999, R. Hanley coll. (SEMC); 5 specimens, the same locality, 1400 m,
1780 5200 S 65420 5400 W, 6.II–8.VI.1999, F. Génier coll. (SEMC); 2 specimens, the same locality and collector, 1400 m, 1780 5200 S 65420 5400 W, 8–12.II.1999
(SEMC); 2 specimens, Cochabamba, 117 km E, Yungas Lagunitas, 1000 m, 1760 2200 S 65400 5700 W, flight intercept trap, 1–6.II.1999, F. Génier coll.
(SEMC); 4 specimens, Cochabamba, 124 km E, Yungas, Cochabamba – Villa Tunari Rd., 700 m, 1730 4500 S 65380 3800 W, carrion trap, 1–6.II.1999, F.
Génier coll. (SEMC); 1 specimen, Cochabamba, 67.5 km NE, Est. Biol. Valle del Sajita, Univ. de San Simon, 300 m, 1760 3300 S 64470 5200 W, flight intercept
trap, 9–13.II.1999, F. Génier coll. (SEMC); Santa Cruz: 1 specimen, 3.7 km SSE Buena Vista, Hotel Flora Y Fauna, 1729.950 S 6333.15W, 400–440 m,
primary forest, FIT, 4–9.XI.2002, R. Leschen coll. (SEMC); 1 specimen, the same locality and collector, Chiquitano forest, at light, 6.XI.2002, S. Lingafelter
coll. (SEMC). BRAZIL: 1 specimen, Chapada, no date and collector (FMNH); 1 specimen, Pebas, no date and collector (IRSNB); 2 specimens, no locality,
date and collector (1 NMW, 2 MNRJ); 1 specimen, Vassouras,18.IX.1933, Eldmann coll. (DZUP); Amazonas: 1 specimen, Ega, no date and collector
(IRSNB); 1 specimen, Reserva Ducke, 26 km. NE Manaus, 28.II.1977, B. C. Ratcliffe coll. (INPA); 4 specimens, 35 km NE Manaus, Res. Flor. Ducke, 25.
VII–08.VIII.[19]95, Arndt and Gröger coll. (ZMHB); Pará: 1 specimen, Aldeia Maracacume, Maranhao 80 km E. of Caninde, 22–23.V.1963, B. Malkin
coll. (FMNH); 7 specimens, Belém, Guamá, 22.III.1974, I. S. Gorayeb coll. (INPA); 16 specimens, Belém, Rio Guamá, 22.III.1974, R. T. Schuh coll.
(AMNH); Bahia: 1 specimen, San Antonio da Barra, no date and collector (IRSNB); 1 specimen, no locality and date, Bohm. coll. (FMNH); 6 specimens,
no locality and date, Fry coll. (FMNH); 1 specimen, no locality, date and collector (FMNH); Goiás: 1 specimen, Jatahy, no date and collector (IRSNB);
Distrito Federal: 1 specimen, Brasília, XI-1987, V. de Mello coll. (AMBC); Rio de Janeiro: 8 specimens, no locality and date, Fry coll. (1 ZMHB, 7
FMNH); 1 specimen, no locality and date, Boehm. coll. (FMNH); 4 specimens, Tijuca, XII.1884, E. Gounelle coll. (2 ZMHB, 2 IRSNB); 15 specimens,
Deodoro, 3.IV.[1]937, W. Zikán coll. (MNRJ); 1 specimen, Tijuca, 5.III.1959, H. Schubart coll. (MNRJ); 1 specimen, the same locality and collector, 18.
V.1959 (MNRJ); 2 specimens, Trapicheiro, 13.XII.1959, the same collector (MNRJ); 1 specimen, Estr. das Canoas, 3.XI.1962, H. Schubart coll. (INPA); 1
specimen, Rio de Janeiro, XII.1967, M. Alvarenga coll. (AMNH); 1 specimen, the same locality and collector, X.1969 (AMNH); 3 specimens, Represa Rio
Grande, III.1972, the same collector (AMNH); São Paulo: 2 specimens, São Paulo, Ypiranga, Ihering coll. (FMNH); Paraná: 2 specimens, nr. Porto de
Cima, 31.III–1.IV.2006, W. Rossi coll. (DZUP); Santa Catarina: 1 specimen, no locality, date and collector (NMW); 1 specimen, no locality and date
Lüderwaldt coll. (FMNH); 3 specimens, no locality and date, Klimsch coll. (2 FMNH, 1 NMW); 2 specimens, Blumenau, no date, Hetschko coll. (NMW); 1
specimen, no locality and date, Fry coll. (FMNH); 1 specimen, Blumenau, no date and collector (DZUP); 1 specimen, no locality, 1930, W. Ehrhardt coll.
(ZMHB); 1 specimen, Corupa, XI.1848, H. Humboldt coll. (AMNH); 1 specimen, Corupa, XII.1948, H. Humboldt coll. (AMNH). Undetermined country:
10 specimens, no locality, date and collector (1 NMW, 3 MNRJ, 2 ZMHB, 4 MZSP).
Revision of Piestus with remarks on related genera
Invertebrate Systematics
581
Piestus gounellei Fauvel, 1902
BRAZIL: Espírito Santo: 2 specimens, Pedra Azul, 1500 m, Domingos Martins, I.2000, A. Bello coll. (AMBC); Rio de Janeiro: 1 specimen, no locality,
XII.1899, ilegible coll. (FMNH); 1 specimen, no locality, date and collector (FMNH); 1 specimen, Nova Friburgo, 2–3.IV.1903, E. Gounelle coll. (IRSNB);
1 specimen, Guanabara, Represa Rio Grande, III.1972, M. Alvarenga coll. (AMNH); 1 specimen, Nova Friburgo, Macaé de Cima, 1500 m, 1–15.I.2006,
P. Grossi, coll. (DZUP); São Paulo: 1 specimen, no locality and date, B. Pohl coll. (AMNH); 1 specimen, no locality and date, B. Mráz coll. (FMNH);
1 specimen, Casa Grande, IX.1938, J. Guerin coll. (FMNH); 4 specimens, Serra da Bocaina, S. Jose Barreiro, 1650 m, XI.1968, M. Alvarenga coll. (AMNH);
Paraná: 1 specimen, Marumbi, Morretes, 500 m, 12.XI.1966, Laroca and O. Mielke coll. (DZUP); 1 specimen, Mananciais da Serra, Piraquara, III.2005,
A. J. C. Aguiar coll. (DZUP); 1 specimen, the same locality, 1000 m, XI.2005, P. Grossi, coll. (DZUP); 5 specimens, the same locality and collector,
30.XI.2005 (1 ZMHB, 4 DZUP); 1 specimen, the same locality and collector, 07.XII.2005 (DZUP); 1 specimen, the same locality and collector, XII.2005
(DZUP); 3 specimens, the same locality and collector, 25180 4000 S 48580 0400 W, 1000 m, 09.XII.2005 (1 ZMHB, 2 DZUP); 7 specimens, the same locality,
17.X.2007, E. Caron and P. Grossi coll. (DZUP); Santa Catarina: 1 specimen, no locality, date and collector (FMNH); 2 specimens, Blumenau, no date,
Hetschko coll. (NMW); 1 specimen, the same locality, no date and collector (FMNH); 1 specimen, Nova Teutonia, II.1933, B. Pohl coll. (MZSP); 1
specimen, the same locality, 27110 B 52230 L, VIII.1935, F. Plaumann coll. (MNRJ); 1 specimen, the same locality and collector, 24.IX.1938 (FMNH); 1
specimen, the same locality and collector, XII.1953 (FMNH). PARAGUAY: Caazapá: 1 specimen, Hermosa. Prop. Sosa family, San Rafael Reserve,
26190 1500 S 55440 5500 W, 90 m, flight intercept trap, 3–6.XII.2000, Z. H. Falin coll. (SEMC); Itapúa: 1 specimen, Yatai, prop. Hostettler family, San Rafael
Reserve, 100 m, 26380 1700 S 55390 5000 W, flight intercept trap, 21–25.XI.2000, Z. H. Falin coll. (SEMC). ARGENTINA: Misiones: 1 specimen, P.N. Iguzu,
Salto Macuco, 26.XII.1990, S. and J. Peck coll. (FMNH); 1 specimen, 15 km SE Puerto Iguazu, mature forest, 27.XII.1990–6.I.1991, S. and J. Peck (FMNH).
Undetermined country: 2 specimens, no locality, date and collector (NMW).
Piestus mexicanus Laporte, 1835
MEXICO: 31 specimens, no locality, date and collector (4 ZMHB, 2 IRSNB, 13 NMW, 11 FMNH, 1 AMNH); 1 specimen, no locality and collector, 1871
(NMW); 7 specimens, Jalapa, no date, Flohr coll. (ZMHB); 1 specimen, no locality an date, L. W. Schaufuss coll. (ZMHB); 1 specimen, no locality and date,
Hoege coll. (NMW); 1 specimen, no locality and collector, 1890 (NMW); 3 specimens, no locality and date, Truqui col. (FMNH); 3 specimens, Jalapa, no
date, Hoge (FMNH); 4 specimens, no locality and date, Reitter coll. (FMNH); Tamaulipas: 4 specimens, Mpo. Gomez Farias, Atlas Cimas, 1000 m, leaf
litter, 16.III.1989, R. W. Jones coll. (1 DZUP, 3 FMNH); Durango: 2 specimens, no locality, date and collector (1 ZMHB, 1 NMW); San Luís Potosí: 2
specimens, El Salto de Agua, 20.VI.1975, L. E. Watrous coll. (FMNH); Hidalgo: 2 specimens, 55 km NE Jacala, 1190 m, 4.VI.1987, R. Anderson coll.
(SEMC); Veracruz: 1 specimen, Jalapa, no date, F. Schneider coll. (ZMHB); 6 specimens, Sierra de Zongolica, no date and collector (5 ZMHB, 1 SEMC); 2
specimens, Cordoba, no date, A. Fenyes coll. (FMNH); 2 specimens, Fortin de las Flores, 28.VI.1975, Q. W. Wheeler coll. (FMNH); 1 specimen, Balzapote,
berlese, rain forest leaf litter, 7.VII.1976, A. Newton coll. (FMNH); 6 specimens, 4.4 mi N Huatusco, cloud forest, 4200ft., sifting litter at edge of stream, 24.
IV.1977, J. S. Ashe coll. (FMNH); 1 specimen, the same locality and collector, 25.IV.1977 (FMNH); 1 specimen, 33 km NE Catemaco, Los Tuxtlas Biol.
Sta., 160 m, ridge rainforest, FIT, 1.VII.1983–1.VIII.1983, S. and J. Peck coll. (FMNH); 1 specimen, the same locality, 20–27.VII.1984, H. and A Howden
coll. (FMNH); 1 specimen, 1.7 km W Huatusco, Hwy. 125, 1250 m, mushrooms, rotting vegetation, 16.VII.1990, I. Yarom coll. (SEMC); 8 specimens, 18.8
km S Huatusco, Hwy. 125 and 13.3 km E on Ixhuatlán Rd., 1130 m, sifting trash, 17.VII.1990, J. S. Ashe, K. J. Ahn and R. Leschen coll. (SEMC); 1
specimen, the same locality and collector, rotten banana trunk, 19.VII.1990 (SEMC); Oaxaca: 6 specimens, 7.3 mi N Chacalapa, 15490 N 96280 W, Hwy
175, tropical deciduous, 410 m, sifting litter along stream, 29.IV.1977, J. S. Ashe coll. (FMNH); Chiapas: 1 specimen, 2.5 mi SW El Bosque, 4000’, near
nest entrance Atta cephalotes, 25.VIII.1973, A. Newton coll. (FMNH); 1 specimen, 8 mi N Pueblo Nuevo Solistahuacán, 6000’, berlese cloud forest/leaf
litter, 26–27.VIII.1973, A. Newton coll. (FMNH); 2 specimens, 13.5 mi s. Mapastepec, under bark log. Lowland, rainforest, 120 m, 30.XII.1975, H. Frania
coll. (AMNH); 12 specimens, 4 km N of Escuintla, wet rooting cut vege. and leaves along stream lowland rainforest, 120 m, 4.I.1977, H. Frania coll.
(AMNH); 25 specimens, 9.7 mi S Solusuchiapa, Hwy 195, montane tropical, 530 m, sifting litter along stream, 5.V.1977, J. S. Ashe coll. (FMNH); 1
specimen, 4.7 km N Finca Prucia, 24.6 km S Jaltinango de la pas., 1050 m, oak-pine-montane, trop. transition, 3.VII.1979, J. S. Ashe coll. (FMNH); 3
specimens, Bonampak Rd., 100 km SE Palenque, berlese rainforest, 8.VII.1983, S. and J. Peck coll. (FMNH); 16 specimens, the same locality and collector,
230 m, 24.VII.1983 (FMNH); 11 specimens, Parque Laguna Belgica, 19.3 km N Ocozocoautla, 970 m, 8.VI.1991, J. Ashe coll. (SEMC); 36 specimens, 5.9
km E Bochil, 1300 m, riparian mesophytic forest litter, 15.IX.1992, R. S. Anderson coll. (SEMC); 4 specimens, 10 km W El Bosque 1475 m, 15.IX.1992, R.
S. Anderson coll. (SEMC). BELIZE: Cayo: 1 specimen, District Chiriqui N.P., 16290 3500 N 89020 4900 W, Doyles Delight, nr. Campground, 1100 m, 19–28.
VIII.2007, P. W. Kovarik coll. (FMNH). GUATEMALA: 8 specimens, no locality, date and collector (7 ZMHB, 1 IRSNB, 2 FMNH); Zacapa: 1 specimen,
3.5 km SE La Union, 1500 m, 23.VI.1993, J. Ashe and R. Brooks (SEMC); 1 specimen, the same locality and collector, 23–25.VI.1993 (SEMC); 1
specimen, the same locality and collector, 25–27.VI.1993 (SEMC); Escuintla: 1 specimen, Finca San Victor, 600ft., decaying bananas, 10.IX.[19]48, R. D.
Mitchell coll. (FMNH). NICARAGUA: Granada: 1 specimen, Res. Nat. Volcan Mombacho, 1150.050 N 8558.830 W, 1210 m, flight intercept trap, 1–5.
VI.2002, R. Brooks, Z. Falin and S. Chatzimanolis coll. (SEMC). COSTA RICA: 3 specimens, San Juan, no date, E. Schimidt coll. (AMNH); 8 specimens,
Vara Blanca, 2000 m, H. Schmidt coll. (4 AMNH, 4 FMNH); 3 specimens, no locality, date and collector (FMNH); 1 specimen, Volcan Irazu, no date and
collector (IRSNB); 1 specimen, Puente de las Mulas, III.1916, no collector (FMNH); 1 specimen, Rio Virilla, 26.XII.[19]31, H. Schmidt coll. (SEMC); 2
specimens, Río Virillo, II.[19]35, no collector (FMNH); 1 specimen, Vara Blanca, II.1936, no collector (FMNH); 1 specimen, no locality, 12–16.II.[19]40,
no collector (FMNH); 3 specimens, El Poró, 800 m, 18.II.1941, no collector (FMNH); 2 specimens, Sardinal, 23.II.1941, no collector (FMNH); Guanacaste:
6 specimens, Lomas Bardubal Biological Reserve, 10300 2200 N 85220 1500 W, 17 m, dead log, no date and collector (SEMC); 2 specimens, Maritza Biol. Stn.,
550 m, flight intcpt. Trap, 22.V.1993, J. and A. Ashe coll. (SEMC); 13 specimens, 14 km WNW Bagaces, Lomas Barbudal, Berlese litter, 29.VI.1994, S.
O’Keefe coll. (FMNH); Alajuela: 1 specimen, Sn. Mateo, 500 m, 28.II.[19]39, no collector (FMNH); 1 specimen, Hiquito, San Mateo, 5.IX.[19]40, Sales
coll. (FMNH); 3 specimens, Escobal,4.IV.[19]43, no collector (FMNH); 1 specimen, 155 km N Jet.Trs. 126(9) and 120, 1.2 km E on Rd to Virgen de
Socorro, Río Sarapiquí, 2300ft., under bark of log, 28.III.1991, L. Herman coll. (AMNH); Heredia: 69 specimens, 9.7 km N Heredia, Rt. 9, 4200ft., litter
near stream, 9.II.1985, L. Herman coll. (AMNH); 2 specimens, 22.8 km N Heredia, Rt.9, 5700ft., litter from stream, 9.II.1985, L. Herman coll. (AMNH); 2
specimens, Sto. Domingo del Heredia, INBio Cafetal, 1100 m, flight intercept trap, 25–28.VI.1997, S. and J. Peck coll. (SEMC); San José: 3 specimens, La
Uruca, 1100 m, no date and collector (IRSNB); 1 specimen, La Caja, 8 Kil., 1934, Schmidt coll. (FMNH); 9 specimens, no locality, date and collector
(AMNH); 32 specimens, no locality, date and E. Schmidt coll. (AMNH); 3 specimens, La Caja, no date, Nevermann coll. (2 NMW, 1 DZUP); 5 specimens,
the same locality, no date, Reimoser coll. (NMW); 2 specimens, the same locality, no date, Schmidt coll. (FMNH); 1 specimen, Finos La Caja, 1930,
582
Invertebrate Systematics
E. Caron et al.
H. Schmidt coll. (FMNH); 12 specimens, La Caja: 8 kil., 1931, Schmidt coll. (FMNH); 4 specimens, no locality and collector, 1935 (FMNH); 1 specimen,
III.[19]37, no collector (FMNH); 5 specimens, Caja, VII-[19]38, no collector (FMNH); 3 specimens, 8.5 km S Orotina, 300 ft., 16.II.1985, L. Herman coll.
(AMNH); 1 specimen, Escazu, 25–29.VI.1988, F. D. Parker coll. (SEMC); 1 specimen, the same locality and collector, 28–31.V.1989 (SEMC); Cartago: 2
specimens, Irazu, no date and collector (NMW); Puntarenas: 9 specimens, Las Cruces Botanical Garden near San Vito, 3500 ft., 27–28.II.1985, L. Herman
coll. (AMNH); 25 specimens, 65 km, W Rincon, nr. Fundacion Neotropica, litter nr. stream, 25.III.1991, the same collector (AMNH); 1 specimen,
Corcovado National Park, Sirena Stn., upper Rio Claro Trail, 8280 2900 N 83350 800 W, 100 m, flight intercept trap, 28.VI–1.VII.2000, Z. H. Falin coll.
(SEMC). PANAMA: 1 specimen, Canal Zone, Barro Colorado Island, Zetek Trail, fibrous debris in tree trunk, 4.IV.1959, H. S. Dybas coll. (FMNH); 1
specimen, Canal Zone, Albrook Forest Site, 100ft., on ground, 20.I.1968, R. Hutton coll. (AMNH); 1 specimen, the same locality, ground black light trap,
30–31.I.1968, R. S. Hutton coll. (AMNH); Chiriquí: 3 specimens, no locality and collector, 1930 (FMNH); 1 specimen, 9.7 road miles S. of El Volcan, alt.
2750ft., 10.III.1959, G. A. Solem coll. (FMNH); 1 specimen, Cordillera Abajo, 08400 5600 N 82360 1000 W, 920 m, litter near stream, 21.V.[20]01, L. Herman
and W. Opitz coll. (AMNH); 1 specimen, near Dolega, 380 m, 8360 1000 N 82250 2000 W, litter near stream, 23.XII.[20]01, L. Herman coll. (AMNH); 1
specimen, 1.7 km E. Rio Sereno, 8490 2300 N 82500 2900 W, 890 m, litter nr stream, 26.XII.[20]01, L. Herman coll. (AMNH); Panamá: 1 specimen, Altos de
Majé, berlese litter in young forest, 6–15.X.1975, D. S. Chandler coll. (FMNH); 12 specimens, Las Cumbres, sift forest litter, 24.X.1975, the same collector
(FMNH); 3 specimens, Cerro Azul, 2000’, wet debris small forest stream, 21.II.1976, A. Newton coll. (1 DZUP, 2 FMNH); 25 specimens, Parque Nac.
Soberamia, Old Gamboa Rd., 4 km E Gamboa, leaf-fruitfall, 11.VI.1993, Jameson coll. (SEMC); 19 specimens, road to Cerro Campana, 8420 3400 N
79530 0600 W, 280 m, litter nr stream, 2.I[20]02, L. Herman coll. (AMNH). COLOMBIA: Boyacá: 1 specimen, no locality, date and collector (IRSNB).
Undetermined country: 5 specimens, no locality, date and collector (2 ZMHB, 3 NMW).
Piestus foveolatus, sp. nov.
COSTA RICA: 10 specimens, Farm Castilla, VI.[19]38, no collector (FMNH); 4 specimens, Finca Castilla, VI.[19]38, no collector (FMNH); 1 specimen,
ilegible locality,1.V.1939, no collector (FMNH); 1 specimen, ilegible locality, 3.VIII.[19]39, no collector (FMNH); 2 specimens, Tapauté?, 9.IX.[19]39,
no collector (FMNH); 1 specimen, Tapauté?, 15.III.[19]40, no collector (FMNH); 2 specimens, Tapauté?, 22–28.I.1941, no collector (FMNH); 1 specimen,
Tapauté?, 27.I.1941, no collector (FMNH); 1 specimen, Peralta, 21.II.1943, no collector (FMNH); 1 specimen, Capellades, 1650m, 19–21.III.[19]40,
no collector (FMNH); 3 specimens, Cervantes, 8.IV.1940, no collector (FMNH); Alajuela: 1 specimen, 96 km. N Jct Rts. 9 and 120, Rd. to Puerto Viejo,
4600’, litter near stream, 17.III.1991, L. Herman coll. (AMNH); 8 specimens, 7.7 km, N Jet. Rt./126(9) and 120, Rd. to Puerto Viejo, 4600’, litter near
stream, 17.III.1991, L. Herman coll. (AMNH); 1 specimen, 14 km S. Volcan Arenal, 10200 N, 84430 W, 1000m, sifted leaf litter, 29.IV.1968, J. Longino
coll. (SEMC); 1 specimen, Eladio’s River trail, 800m, flight intercept, 19.V.1989, J. Ashe, R. Leschen and R. Brooks coll. (SEMC); 1 specimen, Peñas
Blancas, 970m, flight intercept trap, 20.V.1989, the same collector (SEMC); 2 specimens, the same locality and collector, 1190m, flight intercept trap, 20.
V.1989, J. Ashe, R. Leschen and R. Brooks coll. (SEMC); Heredia: 4 specimens, La Selva, 3.2 km SE, Puerto Viejo, 100m, flight intercept trap, 14.II.1992,
W. Bell coll. (SEMC); 1 specimen, the same locality and collector, 21.II.1992 (SEMC); 1 specimen, the same locality and collector, 17.III.1992 (SEMC);
2 specimens, the same locality and collector, 24.III.1992 (SEMC); 1 specimen, the same locality and collector, 27.III.1992 (SEMC); 2 specimens, La Selva
Biol. Res. Sta. 3 km. S. Puerto Viejo, 80m, 10250 000 N, 8400 000 W, flight intercept trap, 2–15.VI.1996, R. Hanley coll. (SEMC); 14 specimens, Finca Zurqui,
Tajo, 10020 300 N, 84010 2600 W, 1575m, wet cloud forest litter, 27.VI.2000, R. Anderson coll. (SEMC); San José: 29 specimens, Cerros de Escazu, 2 km S.
San Antonio, 1650m, 9530 3000 N, 8490 000 W, berlese forest litter, 13.VI.1997, R. Anderson coll. (SEMC); 13 specimens, Cerro Chompipe 2 km. N. Monte de
la Cruz, 2000m, 9530 3000 N, 8490 000 W, berlese forest litter, 13.VI.1997, R. Anderson coll. (SEMC); Puntarenas: 3 specimens, 35 km NE San Vito nr. Las
Alturas, Rio Bella Vista, Rd to Gravel Pit, 4300’elev., dry leaf litter near river, 22.III.1991, L. Herman coll. (AMNH); 1 specimen, the same locality and
collector, wet leaf litter, 23.III.1991 (AMNH); 8 specimens, the same locality, date and collector, moist leaf litter (AMNH); 6 specimens, the same locality,
date and collector, dry leaf litter (AMNH); 5 specimens, San Vito, Estac. Biol. Las Alturas, 2 km NE, 8560 5600 N, 82500 100 W, 1520m, berlese leaf litter,
20.VI.1998, R. Anderson coll. (SEMC); 1 specimen Las Cruces Biol. Sta., San Vito 5 km SW, 1425m, 8460 5900 N, 82590 1800 W, berlese leaf litter,
22.VI.1998, the same collector (SEMC); 15 specimens, Las Alturas Biol. Sta., 1660m, 0856.170 N, 8250.010 W, flight intercept trap, 31.V–3.VI.2004,
J.S. Ashe, Z. Falin and I. Hinojosa coll. (SEMC); 4 specimens, Altamira Biol. Sta., 1510–1600m, 0901.760 N, 8300.490 W, flight intercept trap, 4–7.VI.2004,
the same collector (SEMC). PANAMA: Chiriquí: 1 specimen, Finca Palo Santo, nr. Nueva Califórnia, alt. 4200 ft., at light, 9–10.III.1959, H.S. Dybas coll.
(FMNH); 1 specimen, nr. Nueva California, W. of Finca Palo Santo, 5000 ft., 10.III.1959, H. Dybas coll. (FMNH); 3 specimens, NW of Volcán, 1410m,
08490 1600 N, 82400 2600 W, litter near stream, 17.V.[20]01, L. Herman and W. Opitz coll. (AMNH); 2 specimens, La Fortuna, Cont. Divide Trail, 08460 N,
82120 W, 1150m, flight intercept trap, 23.V–9.VI.1995, J. Ashe and R. Brooks coll. (SEMC); 1 specimen, the same locality and collector, slash, 9.VI.1995;
1 specimen, the same locality and date, berlese forest litter, R. Anderson coll. (SEMC); 1 specimen, 20 km N Gualaca, Finca La Suiza, 1350m, 8390 000 N,
82120 000 W, flight intercept trap, 24.V–9.VI.1995, J. Ashe and R. Brooks coll. (SEMC); 3 specimens, the same locality, 1450m, oak forest litter, 12.VI.1995,
R. Anderson coll. (SEMC); 1 specimen, 27.7 km W. Volcan Hartmann’s Finca, 08450 N, 82480 W, 1450m, flight intercept trap, 14–17.VI.1995, J. Ashe and
R. Brooks coll. (SEMC); 1 specimen, the same locality, 1500m, 8510 4200 N, 82440 3600 W, oak forest litter, 16.VI.1995, R. Anderson coll. (SEMC);
2 specimens, the same locality, 1600m, 8510 4200 N, 82440 4800 W, treefall litter, 17.VI.1996, J. Ashe and R. Brooks coll. (SEMC). COLOMBIA: Valle Del
Cauca: 1 specimen, PNN Farallones de Cali Anchicaya, 3260 N, 76480 W, 730m, Winkler, 21–23.I.2001, S. Sarria coll. (SEMC).
Piestus sulcipennis Scheerpeltz, 1952
BRAZIL: 1 specimen, no locality and date, Bang coll. (FMNH).
Piestus costatus Sharp, 1887
COSTA RICA: 1 specimen, Coronado, 1400–1500 m, 15.VIII.[19]31, Nevermann coll. (FMNH); 1 specimen, no locality and collector, 5.V.[19]40
(FMNH); 1 specimen, Puntarenas-Guanacaste, border Monte Verde, Cerro Amigos, 1760 m, flight intercept trap, 12.V.1989, J. Ashe, R. Brooks and R.
Leschen coll. (SEMC); San José: 7 specimens, km 113 Pan-Am. Hwy, 23 km. N. San Isidro, 2000 m, 9290 000 N 83420 2000 W, berlese forest litter, 20.
VI.1997, R. Anderson coll. (SEMC); 4 specimens, km 117 Pan-Am. Hwy, 19 km. N. San Isidro, 1800 m, 9280 000 N 83420 2000 W, berlese forest litter, 20.
VI.1997, R. Anderson coll. (SEMC); 1 specimen, the same locality and collector, 25.VI.1997 (SEMC); 1 specimen, PanAmerican Hwy Km 80.5, 7 km SSW,
Cabinas de Quetzal, 9330 5300 N 83480 500 W, 2150 m, fogging fungus covered log, 21.VII.2000, J.Ashe, R. Brooks and Z. Falin coll. (SEMC); Cartago: 2
Revision of Piestus with remarks on related genera
Invertebrate Systematics
583
specimens, Tapantí, 22–28.I.1944, no collector (FMNH); 3 specimens, P.N. Tapantil, 1150 m, 9450 4100 N 83470 500 E, flight intercept trap, 17–20.VII.2000, J.
Ashe, R. Brooks and Z. Falin coll. (SEMC); Puntarenas: 7 specimens, OTS Sta. Finca Las Cruces, 4000ft., San Vito, 82580 W 8460 N, Berlese 1500cc. leaf
litter in stream bed, away from flowing water steep banks, Virgin forest cover, 18.III.1973, J. Wagner and J. Kethley coll. (FMNH); 9 specimens, the same
locality and collector, 20.III.1973 (FMNH); 3 specimens, Prov. 35 km. NE San Vito nr. Las Alturas, Rio Bella Vista, Rd. To Gravel Pit, 4300’elev., dry leaf
litter near river, 22.III.1991, L. Herman coll. (1 DZUP, 2 AMNH); 15 specimens, the same locality and collector, moist leaf litter, 23.III.1991 (AMNH); 3
specimens, the same locality and collector wet leaf litter, III-23–1991 (AMNH); 19 specimens, Las Cruces Botanical Garden near San Vito, 3500ft., 27–28.
II.1985, L. Herman coll. (AMNH); 1 specimen, the same locality and collector, 27.II.1985 (AMNH); 1 specimen, Monte Verde, 1400 m, flight intercept trap,
21.V.1989, J. Ashe, R. Brooks and R. Leschen (SEMC); 7 specimens, Altamira Biol. Sta., 1510–1600 m, 0901.760 N 8300.490 W, inside hepialid webs on
trees, 6.VI.2004, J. S. Ashe, Z. Falin and I. Hinojosa coll. (SEMC). PANAMA: Bocas del Toro: 1 specimen, 25 km NNE San Felix, 81500 8340 , 1500 m,
Berlese floor litter and root mat. Quebrada Alicia cloud forest, 11.VI.1980, J. Wagner coll. (FMNH); Chiriquí: 3 specimens, nr. Nueva California, Finca Palo
Santo, 4900’, 6.III.1959, H. S. Dybas coll. (FMNH); 2 specimens, Finca Lerida, nr. Boquete, 5650ft., under bark of log on ground, 12.III.[19]59, H. Dybas
coll. (FMNH); 5 specimens, 4 km NW Volcán, 1450 m, 8490 1900 N 82400 3100 W, litter near stream, 24.XII.[20]01, L. Herman coll. (1 DZUP, 4 AMNH);
3 specimens, Tizingal Rd. NW of Volcán, 8480 3700 N 82390 5300 W, 1350 m, litter nr stream, 25.XII.[20]01 L. Herman coll. (AMNH).
Piestus chiriquensis Sharp, 1887
NICARAGUA: Río San Juan: 1 specimen, 60 km SE San Carlos, Refúgio Bartola, 100 m, 1058,400 N 8420.300 W, 25.V.2002, R. Brooks, Z. Falin and
S. Chatzimanolis coll. (SEMC). COSTA RICA: 2 specimens, Farm Castilla, VI.[19]38, no collector (FMNH); 1 specimen, Finca Castilla, VI.[19]38, no
collector (FMNH); 1 specimen, Chitaría, 23–25.VII.1941, no collector (FMNH); Guanacaste: 1 specimen, Heliconias Biol. Sta., 1042.920 N 8502.380 W,
600 m, flight intercept trap, 20–23.XI.2001, R.Brooks coll. (SEMC); Alajuela: 1 specimen, 9.6 km N. Jct Rts. 9 and 120, Rd. to Puerto, Viejo 4600’, litter
near stream, 17.III.1991, L. Herman coll. (AMNH); 1 specimen, E.B. Sam Ramon, R.B. San Ramon, 27 km. N. and 8 km. W. San Ramon, 10130 400 N
84350 4600 W, flight intercept trap, 8.VII.2000, J. Ashe, R. Brooks and Z. Falin coll. (SEMC); Heredia: 15 specimens, OTS La Selva Field Sta. Puerto Viejo
de Sarapiqui, Rio Puerto Viejo, 10260 N 83590 W, Ber. 100cc conc. Epiphytic leaf mould w. Peripatus, 5–11.III.1973, J. Wagner and J. Kethley coll. (1
DZUP, 14 FMNH); 1 specimen, La Selva Biol. Station nr. Puerto Viejo de Sarapiqui, 18.II.1985, L. Herman coll. (AMNH); 1 specimen, La Selva, 3.2 km SE
Puerto Viejo, 100 m, flight intercept trap, 21.III.1992, W. Bell coll. (SEMC); 1 specimen, 3 km S Puerto Viejo OTS, La Selva, 100 m, X.1992, P. Hanson
coll. (SEMC); 2 specimens, Porrosati, 6 km N. San José de la Montana, 1900 m, 1050 3000 N 8470 000 W, berlese forest litter, 27.VI.1997, R. Anderson coll.
(SEMC); San José: 2 specimens, 117 km. Pan American Highway 19 km N. San Isidro, 9280 000 N 83420 2000 W, 1800 m, cloud forest litter, 15.II.1998,
R. Anderson coll. (SEMC); 1 specimen, PanAmerican Hwy Km 80.5, 7 km SSW, Cabinas de Quetzal, 9330 5300 N 83480 500 W, 2150 m, fogging fungus
covered log, 21.VII.2000, J.Ashe, R. Brooks and Z. Falin coll. (SEMC); 1 specimen, PanAmerican Hwy, Km 80.5, 7 km SSW, Cabinas de Quetzal,
9330 5300 N 83480 500 W, 2150 m, fogging fungus covered log, 22.VII.2000, J. Ashe, R. Brooks and Z. Falin coll. (SEMC); 1 specimen, PanAmerican Hwy. km
80.5, 9.5 km SSW, on San Gerado Rd., Cataraia Trail, 9320 4700 N 83480 4000 W, 2020 m, 22.VII.2000, J. Ashe, R. Brooks and Z. Falin coll. (SEMC);
2 specimens, Genesis II Reserve, 0942.570 N 8354.640 W, 2360 m, 15.VI.2004, J. S. Ashe, Z. Falin and I. Hinojosa coll. (SEMC); Cartago: 1 specimen,
Tapantí, 1100 m, 9.IX.[19]39, no collector coll. (FMNH); 1 specimen, Capellades, 19–21.III.[19]40, no collector (FMNH); 1 specimen, Cañon, 2450 m,
28.XI.1994, H. Forster coll. (NMW); Limón: 1 specimen, Hamburgfarm, 15.XII.[19]24, Nevermann coll. (FMNH); 1 specimen, the same locality and
collector, 4.X.[19]28 (FMNH); 3 specimens, the same locality and collector, 1.X.1936 (FMNH); 3 specimens, Finca Castilla, VI.[19]38, no collector
(FMNH); Puntarenas: 1 specimen, OTS Sta. Finca Las Cruces, San Vito, 82580 W 8460 N, 4000ft., Ber. 1500cc. conc. litter, base of lrg. banana cump,
15.III.1973, J. Wagner and J. Kethley coll. (FMNH); 4 specimens, the same locality and collector, 16.III.1973 (FMNH); 3 specimens, the same locality and
collector, Berlese 1500cc. leaf litter in stream bed, away from flowing water steep banks, Virgin forest cover, 18.III.1973 (FMNH); 1 specimen, the same
locality and collector, 19.III.1973 (FMNH); 6 specimens, the same locality and collector, Berlese leaf litter in stream bed Rio Jaba, rocky crevices 800cc.
conc., 20.III.1973 (FMNH); 2 specimens, Las Cruces Botanical Garden nr. San Vito, 3500ft., 27–28.II.1985, L. Herman coll. (AMNH); 1 specimen, Monte
Verde, 1240 m, berlese of upper streambed, in sand, 10.V.1989, J. Ashe, R. Brooks and R. Leschen coll. (SEMC); 2 specimens, 35 km NE San Vito nr. Las
Alturas, Río Bella Vista, Rd. to Gravel Pit, 4300., wet leaf litter, 23.III.1991, L. Herman coll. (AMNH); 1 specimen, San Vito, Estac. Biol. Las Alturas, 1500
m, V.1992, P. Hanson coll. (SEMC); 1 specimen, Rincon, Cerro Helado, flight intercept trap, 21–25.VI.1997, S. and J. Peck coll. (SEMC); 1 specimen,
Rincon de Osa, 841.1410 N 8331.1170 W, 50 m, 23–26.VI.2001, S. and J. Peck coll. (SEMC); 1 specimen, Las Cruces Biol. Sta., 0847.140 N 8257.580 W,
1330 m, rotting banana plant stems, 29.V.2004, J. S. Ashe, Z. Falin and I. Hinojosa coll. (SEMC). PANAMA: Bocas del Toro: 1 specimen, Almirante, trail
to dam on Nigua Creek, Berlese: fibrous centre of decayed palm log, 25.III.1959, H. S. Dybas coll. (FMNH); Chiriquí: 1 specimen, La Fortuna, Vivero,
08420 N 82140 W, 1150 m, flight intercept trap, 14–18.VI.1994, A. R. Gillogly coll. (SEMC); 3 specimens, 27.7 km W Volcan, Hartmann’s Finca, 8450 000 N
82480 000 W, 1800 m, oak forest litter, 16.VI.1995, R. Anderson coll. (SEMC); Panamá: 6 specimens, Cerro Campana, 3200’, wet debris small forest stream,
23.II.[19]76, A. Newton coll. (1 DZUP, 5 FMNH); Darién: 1 specimen, Cana Biological Station, 7450 1800 N 77410 600 W, 530 m, 09.VI.1996, J. Ashe and
R. Brooks coll. (SEMC). COLOMBIA: 1 specimen, Aguatal, no date and collector (FMNH); Cundinamarca: 1 specimen, Finca San Pablo, 8 km N. Alban,
1800 m, 1–12.VIII.1987, P. and B. Wygodzinsky coll. (AMNH); Nariño: 1 specimen, R.N. La Planada Parcela Permanente, 01150 N 78150 W, 1885 m,
Pitfall, 04–06.XI.2000, G. Oliva coll. (SEMC). VENEZUELA. Lara: 1 specimen, Sanaré, 14.2 km SE Yacambú N.P., 9410 4500 N 69360 4800 W, 1650 m,
flight intercept trap, 18.V–1.VI1998, J. Ashe, R. Brooks and R. Hanley coll. (SEMC). ECUADOR: Esmeraldas: 16 specimens, Bilsa, 0200 000 S 79430 000 W,
flight intercept trap, 28.IV–10.V.1996, P. Hibbs coll. (SEMC); 2 specimens, Bilsa Biological Station, 0200 2400 N 79420 3600 W, 500 m, Malaise trap,
10.V–4.VI.1996, P. Hibbs coll. (SEMC); 19 specimens, Bilsa, 0200 000 S 79430 000 W, flight intercept trap, 10.V–5.VI.1996, P. Hibbs coll. (SEMC);
2 specimens, Bilsa, 0200 000 S 79430 000 W, flight intercept trap, 5.VI–7.VII.1996, P. Hibbs coll. (SEMC); 1 specimen, Bilsa Biological Station, 0200 2400 N
79420 3600 W, 500 m, flight intercept trap, 7–19.VII.1998, P. Hibbs coll. (SEMC); 1 specimen, the same locality and collector, 19.VII.1996 (SEMC);
Sucumbios: 1 specimen, Sacha Lodge, 0280 1400 S 76270 3500 W, 270 m, flight intercept trap, 21–24.III.1999, R. Brooks coll. (SEMC); Pichincha: 1
specimen, Prov., 47 km(S). Sto. Domingo, Rio Palenque Sta, Ber. leaf litter with palm fruits, 260 m, 21–24.II.1976, S. Peck coll. (FMNH); 3 specimens, the
same locality and collector, Ber. litter under decaying fruit, 250 m, 25.II.1976 (FMNH); 1 specimen, 16 kmE Sto. Domingo Tinalandia, 680 m, malaise
rainforest, 25.VII.1985, S. and J. Peck coll. (FMNH); 1 specimen, 17 km SE Sto. Domingo de Colorados, Tinalandia 3000’, light, 16–21.X.[19]88, L.
Herman coll. (AMNH); 1 specimen, W of Alluriquin, Tinalandia, 2600–2800ft., litter nr. stream, 19–20.V.1993, L. Herman coll. (AMNH); 3 specimens, 45
km NNW Quito Macquipucuna Station, 1600–1650 m, flight intercept trap, 3–18.IV.1996, P. Hibbs coll. (SEMC); 2 specimens, Rio Palenque Science
Center, 0360 S 79210 000 W, 200 m, malaise trap, 25.V–6.VII.1996, P. Hibbs coll. (SEMC); 2 specimens, Tinalandia, Santo Domingo, 16 km E, 0160 5300 S
7930 3900 W, 750 m, flight intercept trap, 26–27.III.1999, R. Brooks and D. Brzoska coll. (SEMC); Napo: 1 specimen, 33 km N Tena, 59 km E on Loreto Rd,
584
Invertebrate Systematics
E. Caron et al.
2,800’, 4.XI.[19]88, blk. Light, L. Herman coll. (AMNH); 1 specimen, Baeza, nr. Rio Papallacta, 5800ft., litter on bark, 24.V.1993, L. Herman coll.
(AMNH); 1 specimen, Sierra Azul, Hacienda Aragon, 0400 000 S 77550 000 W, 2200 m, flight intercept trap, 17.II–26.III.1996, P. Hibbs coll. (SEMC); 1
specimen, Cosanga, 4.2 km S on Baaza-Tena, Road then 1.5 km W on pipeline access road, 0370 1900 S 77500 100 W, 2150 m, flight intercept trap, 5–7.
XI.1999, Z. H. Falin coll. (SEMC); Cotopaxi: 2 specimens, N of San Francisco de las Pampas, visc. Rio Esmeraldas, litter and debris, 4400–5000ft., 14–15.
V.[19]93, L. Herman coll. (AMNH); 1 specimen, Canton Sigchos Las Pampas, Otonga Natural Reserve, 25–28.VII.2005, W. Roosi coll. (DZUP);
Tungurahua: 1 specimen, 12.2 km E Baños, 5000ft., litter near stream, 22.V.[19]93, L. Herman coll. (AMNH). PERU: Huánuco: 1 specimen, Divisoria,
1700 m, IX.1946, F. Woytkowski coll. (AMNH); 5 specimens, Cordillera Azul, 39 km NE Tingo María, 1700 m, montane rainforest, window trap, 11–14.
I.1983, A. Newton and M. Thayer coll. (FMNH); Cusco: 1 specimen, pillahuata, Manu rd., km 128, leaf litter, 22.IX.1982, L. E. Watrous and G. Mazurek
coll. (FMNH). BOLIVIA: Cochabamba: 1 specimen, 105 km E Yungas, nr. Río Carmen Mayu, Cochabamba-Villa Tunari Rd., 1780 5100 S 65430 5000 W,
1750 m, flight intercept trap, 8–12.II.1999, R. Hanley coll. (SEMC).
Piestus nevermanni Scheerpeltz, 1952
COSTA RICA: 1 specimen Vara Blanca, VIII.[19]38, no collector (FMNH); Cartago: 1 specimen, P.N. Tapantil, 9450 4100 N 83470 500 E, 1150 m, flight
intercept trap, 17–20.VII.2000, J. Ashe, R. Brooks, Z. Falin coll. (SEMC). PANAMA: Chiriquí: 2 specimens, La Fortuna, Hydro. Trail, 08420 N 82140 W,
1150 m, flight intercept trap, 23.V–9.VI.1995, J. Ashe and R. Brooks coll. (1 DZUP, 1 SEMC); 2 specimens, 5.9 km NE Cerro Punta, 08220 N 82340 W,
2400 m, Par. Nac. Volcan Banu, under bark, 14.VI.1995, J. Ashe and R. Brooks coll. (SEMC); 2 specimens, La Fortuna, ‘Hydro Trail’, 08420 N 82140 W,
1150 m, treefall litter, 22.V.1995, J. and A. Ashe coll. (SEMC).
Piestus paradoxus Bernhauer, 1917
VENEZUELA: Aragua: 15 specimens, 16 km N. Maracay, 10240 N 67290 W, 1500 m, under bark, 7.III.1995, R. W. Brooks coll. (1 DZUP, 14 SEMC);
6 specimens, Rancho Grande Biol. Stn., 10210 000 N 67410 000 W, 1200–1300 m, fungusy log, 12.V.1998, J. Ashe, R. Brooks and R. Hanley coll. (1 AMNH,
1 DZUP, 4 SEMC).
Piestus aper Sharp, 1876
PANAMA: 2 specimens, Barro Colo. Is., 18.VIII.[19]38, E. Williams coll. (FMNH); Bocas del Toro: 1 specimen, Fortuna, Grande road, 8470 N 82120 W,
1050 m, premontane rain forest, sifting litter, 12–14.VII.1987, D. M. Olson coll. (FMNH); Colón: 1 specimen, Parque Nac. Soberania, Pipeline Rd. Km 6.1,
09070 N 79450 W, 40 m, flight intercept trap, 7–21.VI.1995, J. Ashe and R. Brooks coll. (SEMC). COLOMBIA: Amazonas: 1 specimen, PNN Amacayacu
Matamata, 03410 S 70150 W, 150 m, Winkler, 13–15.IX.2000, A. Parente coll. (SEMC). SURINAME: Marowijne: 1 specimen, Perica, 70 km E
Paramaribo on East–west Road, 5400 2800 N 54360 3100 W, 5 m, flight intercept trap, 31.V–5.VI.1999, Z. H. Falin and B. DeDjin coll. (SEMC); 1 specimen,
Palumeu, 3200 5600 N 55260 1800 W, 160 m, flight intercept trap, 5–9.VII.1999, Z. H. Falin and D. Konoe coll. (DZUP); 1 specimen, the same locality and
collector, 7–8.VII.1999 (SEMC). FRENCH GUIANA: Saint-Laurent-du-Maroni: 2 specimens, Eaux Claires, 3.5 mi N Saul, 338–400 N 5313–140 W,
155–260 m, fig fruit fall, 5–13.X.[19]95, L. Herman coll. (AMNH); 5 specimens, Saül, Sifting, IV.1999, A. Berkov coll. (AMNH); Cayenne: 1 specimen,
Roura, 27.4 km SSE, 4440 2000 N 52130 2500 W, 280 m, flight intercept trap, 10.VI.1997, J. Ashe and R. Brooks coll. (SEMC). PERU: Loreto: 7 specimens,
Campamento San Jacinto, 218.750 S 7551.770 W, 175–215 m, flower fall berlese, 2.VII.1993, R. Leschen coll. (SEMC); 4 specimens, the same locality and
collector, 10.VII.1993 (SEMC); 2 specimens, the same locality and collector, 11.VII.1993 (SEMC); 9 specimens, 1.5 km N. Teniente Lopez, 235.660 S
7606.920 W, fruit fall, 17.VII.1993, 210–240 m, R. Leschen coll. (1 DZUP, 8 SEMC); Cusco: 2 specimens, Consuelo, Manu rd. km 165, litter under rotten
palm, 1.X.1982, L. E. Watrous and G. Mazurek coll. (FMNH); 1 specimen, the same locality and collector, rotten palm, 3.X.1982 (FMNH); 2 specimens, the
same locality and collector, 5.X.1982 (FMNH); 3 specimens, the same locality and collector, 12.X.1982 (FMNH); 6 specimens, the same locality and
collector, leaf litter, 13.X.1982 (FMNH); 6 specimens, the same locality and collector, 14.X.1982 (FMNH); Madre de Dios: 1 specimen, Tambopata Lago
Sandoval, 1236.63S 692.24W, 936ft., leaf litter winkler, primary rain forest, 18.V.1998, P. Parrillo coll. (FMNH); 1 specimen, Pantiacolla Lodge, 8 km
NW, El Mirador Trail, Alto Madre de Dios River, 12380 3000 S 71160 4100 W, 800 m, flight intercept trap, 23–26.X.2000, R. Brooks coll. (SEMC). BOLIVIA:
Santa Cruz: 3 specimens, 3.7 km SSE Buena Vista, Hotel Flora Y Fauna, 1729.950 S 6333.150 W, 400–440 m, Chiquitano for. leaf litter, 5.XI.2002, R.
Leschen coll. (SEMC). BRAZIL: Amazonas: 1 specimen, 35 km NE Manaus, Res. Flor. Ducke, 25.VII– 08.VIII.[19]95, Arndt and Gröger coll. (ZMHB);
Bahia: 1 specimen, Ilheus, Mata da Esperanca, 14470 200 S 3930 4500 W, 50 m, atlantic forest, 29.I.1995, D. Agosti coll. (AMNH); Mato Grosso: 1 specimen,
Barra do Tapirape, forest stream near shore, 10.VIII.[19]62, B. Malkin coll. (FMNH). PARAGUAY: Guairá: 2 specimens, Villa Rica, 8.XII.1924, Fr.
Schade coll. (NMW); 1 specimen, Melgarejo, Tucuara Creek, flood detritus, 20.X.1994, U. Drechsel coll. (SEMC); Itapúa: 1 specimen, San Pedro Mi, San
Rafael Reserve, 26310 2400 S 55480 1800 W, 90 m, fruitfall, 27.XI.2000, Z. H. Falin coll. (SEMC).
Piestus angularis Fauvel, 1864
MEXICO: Veracruz: 2 specimens, Cordoba, no date, F.A. Fenyes coll. (1 NMW, 1 FMNH); 2 specimens, El Fortin, 8.VIII.1941, H. Dybas coll. (FMNH).
BELIZE: Cayo: 1 specimen, Mountain Pine Ridge Area, Rio Frio Cave, riparian litter, 16580 18.900 N 88590 46.600 W, 28.V.1997, C. Carlton coll. (SEMC);
1 specimen, Cayo District Chiriqui N.P., Doyle’s Delight nr. Campground, 16290 3500 N 89020 4900 W, 1100 m, 19–28.VIII.2007, P. W. Kovarik coll.
(FMNH). GUATEMALA: Zacapa: 3 specimens, 3.5 km SE La Union, 1500 m, flight intercept trap, 23–25.VI.1993, R. Brooks and J. Ashe coll. (SEMC).
COSTA RICA: 2 specimens, San Pedro Poás, 10.II.1942, no collector (FMNH); 1 specimen, Carrizal, 9–13.IV.[19]44, no collector (FMNH); Alajuela: 1
specimen, Atenas, 13.X.[19]40, no collector (AMNH); 1 specimen, San Ramon, 5 km W, 1220 m, 1–30.XI.1997, P. Hanson coll. (SEMC); Heredia: 1
specimen, OTS, La Selva Field Sta., Puerto Viejo de Sarapiqui, Rio Puerto Viejo, 10260 N 83590 W, 5–11.III.1973, J. Wagner and J. Kethley coll. (FMNH);
2 specimens, La Selva, E. River Trail, flight interrcept trap, 11.III.1992, W. Bell coll. (SEMC); San José: 1 specimen, Puriscal, IX.[19]39, ilegible collector
(FMNH); Limón: 1 specimen, Reventazon, Hamburg Farm 27.XI.1936, F. Nevermann coll. (FMNH); Puntarenas: 6 specimens, OTS Sta. finca Las Cruces,
4000ft., San Vito, 82580 W 8460 N, leaf litter in stream bed, away from flowing water steep banks, virgin forest cover, 19.III.1973, J. Wagner and J. Kethley
coll. (FMNH); 2 specimens, the same locality and collector, 20.III.1973 (FMNH); 17 specimens, Las Cruces Botanical, Garden nr. San Vito, 3500ft., 27–28.
II.1985, L. Herman coll. (AMNH); 1 specimen, Las Alturas Biol. Sta., 0856.170 N 8250.010 W, 1660 m, flight intercept trap, 31.V–3.VI.2004, J. S. Ashe,
Revision of Piestus with remarks on related genera
Invertebrate Systematics
585
Z. Falin and I. Hinojosa coll. (SEMC); 2 specimens, Altamira Biol. Sta., 0901.760 N 8300.490 W, 1510–1600 m, 7.VI.2004, J. S. Ashe, Z. Falin, and I.
Hinojosa coll. (SEMC). PANAMA: 1 specimen, Canal Zone, Barro Colorado Is., wet leaves and flood debris forest stream, 6.II.1976, A. Newton coll.
(FMNH); 1 specimen, the same locality and collector, 8.II.1976 (FMNH); Chiriquí: 1 specimen, 4 km NW Volcán, 8490 1900 N 82400 3100 W, 1450 m, litter
near stream, 24.XII.[20]01, L. Herman coll. (AMNH); Colón: 1 specimen, Parque Nac. Soberania, Pipeline Rd., 09070 N 79450 W, 17–18.V.1995, J. Joly
and C. Chaboo coll. (SEMC). SURINAMEE: Marowijne: 1 specimen, Nassau Mountain, 4480 3600 N 54310 1600 W, 500 m, beating treefall, 2.VI.1999, Z. H.
Falin coll. (SEMC); Para: 1 specimen, Carolina Creek, 11 km SE, Zanderij Airport, 5230 3600 N 5590 2900 W, 30 m, flight intercept trap, 18–20.VI.1999, Z. H.
Falin and A. Gangadin coll. (SEMC). FRENCH GUIANA: Saint-Laurent-du-Maroni: 6 specimens, Eaux Claires, N of Saül, 3390 4400 N 53130 1700 W, 160
m, 5.X.1995, L. Herman coll. (AMNH); 1 specimen, nr. Eaux Claires, 3.5 mil N Saul, 338–400 N 5313–140 W, 155–260 m, leaf litter near stream, 5–13.X.
[19]95, L. H. Herman coll. (AMNH); 1 specimen, Saül, 7 km N, Les Eaux Claires, 3390 4600 N 53130 3900 W, 220 m, treefall litter, 4.VI.1997, J. Ashe and R.
Brooks coll. (SEMC); Cayenne: 1 specimen, Roura, 39.4 km SSE, 4320 4300 N 5280 2600 W, 270 m, treefall litter, 10.VI.1997, J. Ashe and R. Brooks coll.
(SEMC). ECUADOR: Esmeraldas: 2 specimens, Bilsa Biological Station, 0200 2400 N 79420 3600 W, 500 m, Malaise trap, 28.IV–10.V.1996, P. Hibbs coll.
(SEMC); Napo: 1 specimen, Jatun Sacha Biol. Station, 21 km E. Puerto Napo, 400 m, lowland rain for., 7.VII.1994, F. Génier coll. (SEMC); 1 specimen,
Sierra Azul, Hacienda Aragon, 0400 000 S 77550 000 W, 2300 m, flight intercept trap, 17.II–26.III.1996, P. Hibbs coll. (SEMC); Zamora-Chinchipe: 1
specimen, Río Bombuscaro, 470 000 S 78590 000 W, flight intercept trap, 26.VI–4.VII.1996, P. Hibbs coll. (SEMC). PERU: Cusco: 1 specimen, Consuelo,
Manu rd. km 165, litter under rotten palm, 1.X.1982, L. E. Watrous and G. Mazurek coll. (FMNH); 2 specimens, the same localiy and collector, 2.X.1982
(FMNH); 16 specimens, the same locality and collector, 4.X.1982 (FMNH); 26 specimens, the same locality and collector, 5.X.1982 (2 DZUP, 24 FMNH);
11 specimens, the same locality and collector, 6.X.1982 (FMNH); 2 specimens, the same locality and collector, 7.X.1982 (FMNH); 1 specimen, the same
locality and collector, 7–8.X.1982 (FMNH); 6 specimens, the same locality and collector, 8.X.1982 (FMNH); 1 specimen, the same locality and collector,
9.X.1982 (FMNH); 7 specimens, the same locality and collector, 12.X.1982 (FMNH); 1 specimen, the same locality and collector, 13–15.X.1982 (FMNH).
BOLIVIA: Cochabamba: 3 specimens, Cochabamba, 109 km E Yungas, Cochabamba-Villa Tunari Rd., 1780 5000 S 65420 2900 W, 1480 m, flight intercept
trap, 1–6.II.1999, F. Génier coll. (SEMC); 26 specimens, Cochabamba, 109 km E Yungas, Cochabamba-Villa Tunari Rd., 1780 5000 S 65420 2900 W, 1480 m,
flight intercept trap, 6–8.II.1999, R. Hanley coll. (SEMC); 3 specimens, 109 km E Yungas, Cochabamba-Villa Tunari Rd., 1780 5000 S 65420 2900 W, 1480 m,
flight intercept trap, 8–12.II.1999, R. Hanley coll. (SEMC). BRAZIL: Santa Catarina: 1 specimen, Blumenau, no date and collector (NMW); 1 specimen,
no locality, date and collector (FMNH). PARAGUAY: Alto Paraná: 1 specimen Alto Piray, VII-[1]900, Silvestri coll. (FMNH).
Piestus rugosus Sharp, 1876
GUYANA: 1 specimen, Kartabo, Bartica, 14.V.1924, no collector (AMNH). ECUADOR: Sucumbios: 3 specimens, Sacha Lodge, 0280 1400 S 76270 3500 W,
270 m, under bark, 24.III.1999, R. Brooks coll. (1 DZUP, 2 SEMC). PERU: Cusco: 1 specimen, Marcapata, no date and collector (IRSNB). BOLIVIA:
2 specimens, Yuracaris, no date and collector (IRSNB); Cochabamba: 1 specimen, Cochabamba, 117 km E Yungas, Cochabamba-Villa Tunari Rd.,
1760 3200 S 65410 1200 W, 1040 m, flight intercept trap, 1–6.II.1999, R. Hanley coll. (SEMC); 2 specimens, 67.5 km NE, Est. Biol. Valle del Sajita, Univ. de
Sam Simon, 1760 3300 S 64470 5200 W, 300 m, fungus covered log, pyrethrum insecticide fogging, 7.II.1999, R. Hanley coll. (SEMC). BRAZIL: Pará: 1
specimen, no locality and collector, Fry coll. (FMNH); 1 specimen, Aldeia Aracu, Igarape, Gurupu-Umu, Maranhao, 50 km E of Caninde, under bark,
V.1963, B. Malkin coll. (FMNH).
Hypotelus laevis (Solsky, 1872)
PERU: Cusco: 3 specimens, Consuelo, Manu rd. km 165, palm fruit, winkler, 1.X.1982, L. E. Watrous and G. Mazurek coll. (FMNH); 2 specimens, the
same locality and collector, 3.X.1982 (FMNH); 2 specimens, the same locality and collector, 4.X.1982 (FMNH); 2 specimens, the same locality and collector,
5.X.1982 (FMNH).
Hypotelus andinus (Bernhauer, 1917)
COLOMBIA: Boyacá: 1 specimen, SFF Iguague La Planada, 5250 N 73270 W 2850 m, Malaise, 11.X–01.XI.2000, P. Reina coll. (SEMC); 1 specimen, the
same locality and collector, 01–23.XI.2000 (SEMC). ECUADOR: 1 specimen, Banos, 1500’, 6.V.[19]39, W. C. Macintyre coll. (AMNH).
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