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Review of Brazilian cave psocids of the families Psyllipsocidae and Prionoglarididae (Psocodea: 'Psocoptera': Trogiomorpha) with a key to the South American species of these families

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ISSN 0035-418
INTRODUCTION
All over the world, representatives of the psocid families
Psyllipsocidae and Prionoglarididae, belonging to the
suborder Trogiomorpha (Yoshizawa et al., 2006), are
regularly found in caves (Badonnel & Lienhard, 1994;
Lienhard, 2000, 2002, 2004a, 2007; Lienhard et al.,
2010a). But both families are poorly known from South
America (Lienhard & Smithers, 2002; Lienhard, 2004b,
2011). We present here a synthesis of the results of a study
of these families as they are represented in Brazilian
caves. This study is a part of a long-term research project
of the junior author on Brazilian cave invertebrates.
Since 1996 some 800 caves have been visited by the ju-
nior author and collaborating biospeleologists. Psocids of
the above mentioned families (especially Psyllipsocidae)
were observed in about 300 caves, mostly nymphs, but
sometimes also adults. While the identi cation of the
nymphs was not possible, the roughly 400 adult indivi-
duals of Psyllipsocidae (Fig. 1A) and Prionoglarididae
(Fig. 1BC) submitted to the senior author could be iden-
ti ed down to species level. This material was collected
in 124 Brazilian caves situated in 59 municipalities and
13 states (Appendix 2). A total of 23 species belonging
to three genera could be distinguished, 19 of them new
to science (Appendix 1). These new species were de-
scribed by Lienhard et al. (2010b) and Lienhard & Fer-
reira (2013a, b, 2014), whilst the four previously known
non-endemic species are treated below. All psyllipsocids
and prionoglaridids presently known from South America
are listed (Appendix 1) and keyed. General observations
on the distribution of the Brazilian cave species are also
presented and some evolutionary aspects are discussed.
This is the most important investigation on cave psocids
ever realized (for a bibliography on cave psocids see
Badonnel & Lienhard, 1994 and Lienhard, 2002, 2014).
It revealed an unexpected diversity for two families
which were almost unknown from South America,
not only from caves but also from non-subterranean
habitats. Some particularly interesting results of this
Revue suisse de Zoologie (March 2015) 122(1): 121-142
Review of Brazilian cave psocids of the families Psyllipsocidae and Prionoglarididae
(Psocodea: ‘Psocoptera’: Trogiomorpha)
with a key to the South American species of these families
Charles Lienhard1 & Rodrigo L. Ferreira2
1 Muséum d’histoire naturelle, c. p. 6434, CH-1211 Genève 6, Switzerland. Corresponding author.
E-mail: charleslienhard@bluewin.ch
2 Universidade Federal de Lavras, Departamento de Biologia (Zoologia), CP. 3037, CEP. 37200-000 Lavras (MG),
Brazil. E-mail: drops@dbi.u a.br
Abstract: Data on Brazilian cave psocids (Insecta) of the families Psyllipsocidae and Prionoglarididae are summarized,
as a synthesis of the results of the most important investigation on cave psocids ever realized. Prionoglarididae are
represented by 4 species of the endemic cavernicolous genus Neotrogla, Psyllipsocidae by 17 species of Psyllipsocus
(15 of them endemic) and 2 widely distributed species of Psocathropos. These 19 recently described Brazilian endemic
Neotrogla and Psyllipsocus were discovered in the course of the project, based on the examination of about 400 adult
psyllipsocids and prionoglaridids collected in 124 caves situated in 59 municipalities and 13 Brazilian states. Some
augmentations to the descriptions of the widely distributed Psyllipsocus ramburii, Psyllipsocus yucatan, Psocathropos
lachlani and Psocathropos pilipennis are given and the following new synonymies are proposed: Psyllipsocus ramburii
Selys-Longchamps (P. variabilis Badonnel n. syn., P. dubius Badonnel n. syn.), Psyllipsocus yucatan Gurney (P. collarti
Badonnel n. syn., P. decui Badonnel n. syn.), Psocathropos lachlani Ribaga (Vulturops termitorum Townsend n. syn.,
Dorypteryx astizi Brèthes n. syn.). The distribution of the 23 species of psyllipsocids and prionoglaridids known from
Brazilian caves is analysed and some evolutionary aspects are discussed. A key to the 25 South American species of
these families is given, including two other previously known species: Psyllipsocus delamarei from Argentina and the
troglobitic prionoglaridid Speleopsocus chimanta from Venezuela.
Keywords: Brazil - cave fauna - endemism - male genitalia - new synonymies.
Manuscript accepted 26.08.2014
DOI: 10.5281/zenodo.14579
122 C. Lienhard & R. L. Ferreira
Fig. 1. (A) Psyllipsocus yucatan, habitus in dorsal view (in alcohol), body length 1.5 mm (from head to abdominal tip) (after Lienhard
et al., 2012). (B) Neotrogla curvata, habitus in dorso-lateral view (alive, in cave), body length 3.5 mm. (C) Neotrogla aurora,
habitus in dorsal view (alive, in cave), body length 2.8 mm.
Brazilian cave psocids 123
study have already been presented in two special papers:
the enigmatic presence of microcrystals on the wings of
some specimens of Psyllipsocus yucatan by Lienhard
et al. (2012) and the spectacular coupling role reversal
(reversed intromittent organs) in the genus Neotrogla by
Yoshizawa et al. (2014).
MATERIAL AND METHODS
The material examined was collected by RLF (unless other
collector mentioned) by hand-collecting (for collecting
data, see Appendix 2). Dissection and slide-mounting
followed the methods described by Lienhard (1998). For
general remarks and morphological abbreviations used in
the descriptions, see Lienhard & Ferreira (2014).
The material examined is deposited in the following
institutions: Universidade Federal de Lavras, Depar-
tamento de Biologia (Coleção de Invertebrados
Subterrâneos), Lavras, Brazil (ISLA); Muséum d’histoire
naturelle, Geneva, Switzerland (MHNG); Systematic
Entomology, Hokkaido University, Sapporo, Japan
(SEHU) (some specimens of Neotrogla spp.).
Abbreviations used in the key (nomenclature of wing
veins according to Yoshizawa, 2005): A1 = rst anal
vein; AP = areola postica (a marginal cell in forewing
formed by veins CuA1 and CuA2); CuA1 = rst branch
of cubitus anterior; CuA2 = second branch of cubitus
anterior; CuP = cubitus posterior; FW = forewing
(length); FWw = forewing (greatest width); M = media;
M1-M3 = branches of media; m1 = medial cell anteriorly
delimited by M1; m2 = medial cell anteriorly delimited
by M2; P2 = second article of maxillary palp; R = radius;
Rs = radial sector (undivided basal part of R2-R5); R1 =
rst branch of radius; r1 = radial cell anteriorly delimited
by R1; R2+3 = rst branch of radial sector; r3 = radial
cell anteriorly delimited by R2+3; R4+5 = second branch
of radial sector; r5 = radial cell anteriorly delimited by
R4+5; Sc = subcosta; v3 = third (external) valvula of the
ovipositor.
Abbreviations for Brazilian states: AL = Alagoas, AM
= Amazonas, BA = Bahia, CE = Ceará, ES = Espírito
Santo, GO = Goiás, MG = Minas Gerais, MT = Mato
Grosso, PI = Piauí, RN = Rio Grande do Norte, RS = Rio
Grande do Sul, SP = São Paulo, TO = Tocantins.
TAXONOMIC PART
Non-endemic Psyllipsocidae from Brazil
All South American species of the families Psyllipsocidae
and Prionoglarididae are listed in Appendix 1, based on
previously published data (Badonnel, 1962; Lienhard et
al., 2010a, b; Lienhard & Ferreira, 2013a, b, 2014) and
on the following data concerning four widely distributed
Brazilian species, not treated in detail before. At present,
these four species are the only non-endemics known
from Brazil. The 15 endemic species of Psyllipsocus
were treated by Lienhard & Ferreira (2013b, 2014) and
the 4 endemic species of Neotrogla by Lienhard et al.
(2010b), Lienhard & Ferreira (2013a) and Yoshizawa et
al. (2014). The South American fauna of these families
comprises 25 species, 23 of which are known from Brazil
(Psyllipsocus delamarei only known from Argentina
and Speleopsocus chimanta from Venezuela). See also
the identi cation key for the South American species,
presented below.
Psyllipsocus ramburii Selys-Longchamps, 1872
Figs 2-3
Psyllipsocus ramburii Selys-Longchamps, 1872: 146.
Psyllipsocus variabilis Badonnel, 1986: 182. Syn. nov.
Psyllipsocus dubius Badonnel, 1987: 173. Syn. nov.
For further synonymy see Lienhard & Smithers (2002).
Material examined: ISLA and MHNG; numerous
females, most of them micropterous (slide-mounted
and in alcohol), collected by R. L. Ferreira in caves
situated in the following Brazilian municipalities. –
Castelo (ES), Gruta do Limoeiro cave, 7.i.2005. – Arcos
(MG), Caverna do Alinhamento cave, 1.vi.2002. –
Cordisburgo (MG), Gruta de Maquiné cave, 10.vii.2000.
– Cordisburgo (MG), Gruta Santo Amaro 1 cave,
28.ix.2010. – Coromandel (MG), Gruta João do Pó
cave, 5.x.2000. – Coromandel (MG), Gruta Ronan
cave, 3.x.2000. – Coromandel (MG), Gruta Ronan II
cave, 4.x.2000. – Diamantina (MG), Gruta do Salitre
cave, 13.xii.2007. – Itumirim (MG), Gruta Santo
Antônio cave, 25.x.2002. – João Pinheiro (MG), Gruta
do Sapecado cave, 15.x.2010. – Lagoa Santa (MG),
Gruta da Lapinha cave, 12.vi.2002. – Matozinhos (MG),
Gruta dos Irmãos Piriás cave, 28.vii.2000. – Matozinhos
(MG), Gruta Lavoura cave, 28.+29.v.1997, 26.vi.1997,
19.ii.2000. – Matozinhos (MG), Meandro Abismante
cave, 6.vii.2002. – Matutina (MG), Gruta 9 cave,
10.x.2010. – Moeda (MG), cave SMS 29, 11.xii.2005.
– Pains (MG), Gruta do Capão cave, 5.v.2001. – Pains
(MG), Gruta Paranoá cave, 15.i.2008. – Pains (MG),
Gruta Retiro cave, 27.xi.1999. – Paracatu (MG), Gruta
da Fazenda Tamanduá II cave, 14.x.2010. – Paracatu
(MG), Lapa do Brocotó cave, 16.ix.2010. – Paracatu
(MG), Lapa de Santo Antônio cave, 13.vii.2010. – Santa
Maria do Suaçuí (MG), Gruta do Rio Suaçuí cave,
19.vii.2002. – Sete Lagoas (MG), Gruta Rei do Mato
cave, 3.+4.xi.2011. – Vazante (MG), Lapa da Delza
cave, 12.vii.2010. – Torres (RS), Furna da Lagoa de
Itapeva cave, 20.v.2008. – Altinópolis (SP), Gruta do
Itambé cave, 27.ii.2006. – Altinópolis (SP), Gruta Olho
de Cabra cave, 2.iii.2006. – Altinópolis (SP), Gruta do
Paraná cave, 1.iii.2006. – Itirapina (SP), Gruta da Toca
cave, 22.x.2004.
Further description: P. ramburii is the type species
of the genus Psyllipsocus; it is polymorphic and
occurs in three forms: macropterous, brachypterous
124 C. Lienhard & R. L. Ferreira
Fig. 2. Psyllipsocus ramburii, from Brazilian caves, macropterous female MHNG 7929 from Gruta Santo Antônio (A-D, H),
micropterous female MHNG 7954 from Gruta do Rio Suaçuí (E-G). (A) Forewing. (B) Hindwing. (C) P2-P4 of maxillary
palp. (D) P2-chaetotaxy. (E) Habitus (body length 1.7 mm). (F) Pedicel, showing absence of microspades organ (compare with
Fig. 7D). (G) Spermapore plate and spermatheca. (H) Spermatheca.
Brazilian cave psocids 125
and micropterous; the existence of an apterous form is
somewhat doubtful and has yet to be con rmed (see
Key and Discussion, below). For detailed descriptions
see Mockford (1993) and Lienhard (1998). Two
characters are also especially noted by Mockford
(2011): “macropterous form lacking distal closed cell
in forewing; setal organ at base of paraproctal spine
consisting of one very long and one short seta.” Here
we note the following additional characters (Figs 2-3,
see also Key): pedicel lacking microspades organ (Fig.
2F); P2 lacking stout sensillum (Fig. 2D); spermatheca
small and crumpled (due to parthenogenesis), with 3-4
small and irregularly shaped sclerites near duct (Fig.
2GH; see also Lienhard, 1998: g. 37e and Badonnel,
1986: g. 22); spermapore region with a characteristic
pair of more or less sclerotized longitudinal straps (Fig.
2G; see also Mockford, 1993: g. 62 and Badonnel,
1986: g. 22); macropterous form occasionally with
slightly shortened wings (not as short as in the typical
brachypterous form, see Lienhard, 1998: gs 37c, 38c)
and rather variable venation (Fig. 3, see also Badonnel,
1986: gs 14-18); in micropterous form length of
minute wing rudiments variable, relatively long and
bearing some vein rudiments (Lienhard, 1998: g. 37a,
g) or very short and veinless (Fig. 2E).
Biology: P. ramburii is a parthenogenetic species
(thelytoky) and only one single (accidental?) male is
known (Lienhard, 1977, 1998). The phallosome of the
male is equipped with a pair of well-developed basal
struts, as they are present in all psyllipsocids (Mockford,
2011) except for most of the Brazilian cave endemics
(Lienhard & Ferreira, 2014; see also Evolutionary
Considerations, below).
Distribution: P. ramburii is a widespread species
(Lienhard & Smithers, 2002), often domicole but also
regularly found in caves all over the world (Badonnel &
Lienhard, 1994). Previously published South American
records can be found in the following references:
Strinati, 1971 (Uruguay); Badonnel, 1972 (Chile);
Badonnel, 1986 (Colombia, as P. variabilis); Badonnel,
1987 (Venezuela, as P. dubius); Cammousseight & New,
1994 (Chile).
Discussion: The presence, in the macropterous
specimens from Brazil, of a variability of wing venation
similar to that described by Badonnel (1986) for
P. variabilis from Colombia, and the absence of other
distinctive characteristics, support synonymizing the
latter with P. ramburii.
The case of P. dubius is more complex. This apterous
species was described from Venezuela by Badonnel
(1987), based on the holotype female from cacao ground
litter and a second (non-paratype) female from a cave.
The microscopical slides of both specimens, mounted by
André Badonnel himself, are deposited in the MHNG and
were examined. Unfortunately the thorax of the holotype
had been damaged and only the prothorax is present on
the slide. Thus, the absence of wing rudiments could not
be con rmed for this specimen. The second specimen
is better preserved; it is complete, though its thorax
was deformed by slide-mounting. Surprisingly it bears
wing rudiments which are very similar to those of the
micropterous form of P. ramburii; in this female, they are
particularly small and completely veinless (as in the female
from a Brazilian cave gured in Fig. 2E). This specimen
of P. dubius has clearly to be assigned to P. ramburii. In
the original description of P. dubius, Badonnel insists
on the complete apterism of this species and does not
mention any variability of wing development. He also
mentions that the only morphological difference between
P. ramburii and P. dubius is the complete absence of wing
rudiments in the latter. Even though the holotype might
really be apterous, we are convinced that synonymy of
P. dubius with P. ramburii is justi ed. The terminalia
Fig. 3. Psyllipsocus ramburii, macropterous female (MHNG 8125) with slightly shortened wings, from Gruta João do Pó cave (Brazil,
MG, Coromandel). Forewings and hindwings (pilosity not shown).
126 C. Lienhard & R. L. Ferreira
of the holotype correspond exactly to P. ramburii; in
particular the spermapore region with the typical pair of
sclerotized longitudinal straps clearly visible. However,
the quite plausible existence of an apterous form of this
polymorphic species is not yet de nitively con rmed
with this synonymization. It is possible that Badonnel
did not realize that the part of the broken thorax of the
holotype present on the slide was only the prothorax and
misinterpreted the absence of wing rudiments on this part
as apterism, while he probably overlooked the small wing
rudiments present on the deformed slide-mounted thorax
of the second female. Alternatively, it is also possible that
he had already observed the absence of wing rudiments in
the holotype before dissecting the specimen; impressed
by this unusual character he might have overlooked their
presence in the second female.
Psyllipsocus yucatan Gurney, 1943
Figs 1A and 4-6
Psyllipsocus yucatan Gurney, 1943: 212.
Psyllipsocus collarti Badonnel, 1946: 140. Syn. nov.
Psyllipsocus decui Badonnel, 1977: 340. Syn. nov.
Material examined: ISLA and MHNG; numerous
males and females (slide-mounted and in alcohol),
collected by R. L. Ferreira (unless other collector
mentioned) in caves situated in the following Brazilian
municipalities. – Murici (AL), Toca da Raposa 1 cave
(granite), 13.i.2007. – Murici (AL), Toca da Raposa 2
cave (granite), 13.i.2007. – Campo Formoso (BA), Toca
da Tiquara cave, 8.i.2008, i.2009, xii.2010. – Santa
Maria da Vitória (BA), cave PEA 343, 15.v.2011, leg.
S. S. Salgado. – Ubajara (CE), Gruta dos Mocós cave,
3.i.2007. – Ubajara (CE), Gruta do Morcego Branco
cave, 3.i.2007. – Ubajara (CE), Gruta de Ubajara
cave, 30.xii.2006. – Castelo (ES), Gruta do Limoeiro
cave, 7.i.2005. – Arinos (MG), Lapa do Salobo cave,
18.vii.2010. – Santa Maria do Suaçuí (MG), Gruta do
Rio Suaçuí cave, 19.vii.2002. – T lo Otoni (MG),
Lapa da Vaca Parida cave, 26.i.2005. – Felipe Guerra
(RN), Caverna Trapiá cave, 6.i.2010, 4.viii.2010, leg.
D. M. Bento. – Ilha Bela (SP), Gruta da Serraria cave,
22.iv.2006.
Further description: This species is always macro-
pterous. For detailed descriptions see Gurney (1943)
and the augmented description given by Mockford
(2011). Here we note the following characters (Figs 4-6,
see also Key). Forewing usually with a subtle but typ-
ical colouration (Fig. 4G), as described by Gurney
(1943: p. 212): “membrane faintly tinged with fuscous,
a clear spot at posterior margin of wing between apex
of Cu2 [= CuA2] and apices of anal veins [= CuP and
A1].” P2 lacking clearly differentiated stout sensil-
lum (Fig. 4D). Epiproct characteristically pigment-
ed (Fig. 4BF). Paraproctal setal organ consisting of
a minute hair and a longer and thicker seta; anal spine
replaced by a stout, relatively short and basally arti-
culated seta (not much longer than long seta of setal
organ) (Fig. 4BF); female paraproct simple, male para-
proct ventrally with a small concave protuberance
(“fossette inférieure” of Badonnel, 1946) (Fig. 4F).
Pigmentation of hypandrium interrupted in middle of
anterior margin (Fig. 6AB); aspect of distal phallosome
sclerites somewhat variable, depending on view (dorsal/
ventral) and on position after slide-mounting (Fig. 6A
similar to g. 5 in Badonnel, 1946; Fig. 6B similar to
g. 107 in Mockford, 2011). Sclerotization of oviposi-
tor v1 and v2 variable; sclerotized median axis of v1
weakly developed or absent; median axis of v2 usual-
ly well-sclerotized (Fig. 5A), but weakly developed
or completely absent in pale indivi duals (so observed
in several pale specimens from Brazil, Antigua and
Thailand). Spermapore plate weakly sclerotized but
characteristic (Fig. 5B); spermathecal sac with complex
sclerotizations near origin of duct (Fig. 5DE); spermato-
phores not sclerotized, somewhat variable in shape, but
always similar to that gured in Fig. 5C; the spermathe-
ca of a female may contain up to about a dozen sper-
matophores (see legend to Fig. 5B).
Biology: In general, P. yucatan seems to be a cave
species and it was explicitly indicated as living on
vegetation only once (Badonnel, 1948: on leaves of an
orange tree), with no domicole population known. The
presence of up to about a dozen spermatophores in the
spermatheca of one single female indicates that the
species is polyandrous.
Distribution: P. yucatan was originally described from
a Mexican cave by Gurney (1943). Previously published
South American records can be found in the following
references: García Aldrete & Mockford, 2009 (Brazil,
probably same record as that mentioned by Mockford,
2011); Mockford, 2011 (Brazil, in an arti cial cave);
Lienhard et al., 2012 (Brazil, in caves). The species is
also known from a cave in Cuba (Badonnel, 1977, as
P. decui) and from caves in Jamaica, Aruba and Antigua
(MHNG, unpublished). African specimens are known
from Congo (Badonnel, 1946, 1948, as P. collarti, on
vegetation), Kenya and Senegal (MHNG, unpublished).
Some specimens of P. yucatan have also been collected
in a cave in Thailand (MHNG, unpublished). The
species seems to be widely distributed in the tropics.
Discussion: The only difference between P. yucatan and
P. decui indicated by Mockford (2011) is the absence of
the sclerotized median axis of v2. The above mentioned
variability shows that this character cannot be used
to separate these species. The absence of a sclerotized
v2-axis in the palest specimens examined corresponds
to the situation described by Badonnel (1977) for the
very pale holotype of P. decui from Cuba, where v1 and
v2 are completely hyaline. Therefore P. decui is here
considered as a junior synonym of P. yucatan.
Brazilian cave psocids 127
Fig. 4. Psyllipsocus yucatan, from Brazilian caves, general morphology. (A) Habitus, male. (B) Epiproct and right paraproct, female.
(C) Lacinial tip, female. (D) P2-chaetotaxy, female. (E) P2-P4 of maxillary palp, female. (F) Epiproct and left paraproct, male.
(G) Forewing, male. (H) Hindwing, male.
128 C. Lienhard & R. L. Ferreira
Fig. 5. Psyllipsocus yucatan, from Brazilian caves, female genitalia. (A) Subgenital plate, right ovipositor valvulae and right hind
corner of clunium. (B) Spermapore plate and spermatheca (the latter contains about a dozen spermatophores, only one of them
shown). (C) Spermatophore (not same female as B). (D) Spermathecal sclerotizations and proximal part of spermathecal duct,
specimen MHNG 7990 from Toca da Raposa 2, AL. (E) Ditto, specimen MHNG 8000 from Gruta de Ubajara, CE.
Brazilian cave psocids 129
Badonnel (1948, 1977) and Mockford (2011) noted the
close proximity of P. yucatan and the African P. collarti.
CL has examined the series of syntypes of P. collarti
(1 female and 1 male slide-mounted by Badonnel, the
latter on two separate slides, and several specimens in
alcohol, some of them badly damaged, all deposited
in the Musée Royal de l’Afrique Centrale, Tervuren,
Belgium). No signi cant difference between the African
and the Brazilian specimens could be observed. The
structure gured by Badonnel (1946: g. 10, labeled vv)
as a sclerotized distal appendage of v1 is only visible on
one side of the slide-mounted terminalia; it is evident that
this is an artefact (probably a particle of detritus adhering
to the membranous v1). P. collarti is here considered as a
junior synonym of P. yucatan.
Fig. 6. Psyllipsocus yucatan, from Brazilian caves, male genitalia. (A) Hypandrium and phallosome, ventral view. (B) Ditto, dorsal
view, other specimen than in A (hypandrium schematically shown by interrupted lines).
130 C. Lienhard & R. L. Ferreira
Psocathropos lachlani Ribaga, 1899
Fig. 7
Psocathropos lachlani Ribaga, 1899: 157.
Vulturops termitorum Townsend, 1912: 269 (type species of
Vulturops). Syn. nov.
Psocathropos termitorum (Townsend); Roesler, 1944:
134 (Vulturops considered as a junior synonym of
Psocathropos).
Dorypteryx astizi Brèthes, 1923: 117. Syn. nov.
Psocathropos astizi (Brèthes); Lienhard, 1977: 435 (new
combination suggested).
Psocathropos astizi (Brèthes); Lienhard & Smithers, 2002: 43.
For further synonymy see Lienhard & Smithers (2002).
Material examined: ISLA and MHNG; several males
and females (slide-mounted and in alcohol), collected by
R. L. Ferreira in caves situated in the following Brazilian
municipalities. – Delmiro Gouveia (AL), Gruta Morcego
cave, 23.iv.2006. – Curaçá (Patamuté) (BA), Toca d’agua
de Patamuté cave, 6.i.2008. – Ubajara (CE), Gruta de
Ubajara cave, 30.xii.2006. – Santa Teresa (ES), Gruta
do André Huscki cave, 4.i.2005. – Cordisburgo (MG),
Gruta de Maquiné cave, 10.vii.2000 and 18.v.2010. –
Montalvânia (MG), Gruta Nossa Senhora do Perpétuo
Socorro cave, 14.vii.2007. – Nacip Raydan (MG), Gruta
Manga de Pedra cave (granito), 21.vii.2002. – T lo
Otoni (MG), Lapa da Vaca Parida cave, 26.i.2005. –
Altinópolis (SP), Gruta do Itambé cave, 27.ii.2006.
Further description: P. lachlani is the type species of
the genus Psocathropos; it is usually brachypterous. The
wings of the rare macropterous form (see Key) were
rst described by Mockford (1993); two macropterous
females from a laboratory culture (Jamaica: Kingston,
leg. B. D. Turner, 1971) are deposited in the MHNG.
For detailed descriptions see Mockford (1993) and
Lienhard (1998); some important characters are also
mentioned by Badonnel (1967, see Discussion below)
and in the key given by Mockford (1991). Here we note
the following characters (Fig. 7, see also Key). Length
of forewing of brachypterous form variable, usually
not reaching abdominal tip; venation of forewing also
somewhat variable, but Rs always simple (Fig. 7A).
Hindwing of brachypterous form always reduced to a
minute bare and veinless ap (Fig. 7B). P2 lacking stout
sensillum. Pedicel with well-developed microspades
organ (Fig. 7E). Paraproctal trichobothria with weakly
differentiated basal orets; setal organ consisting of one
minute hair and one somewhat thicker and longer seta;
anal spine not differentiated, replaced by a normal seta
which is not much longer than the ventral seta of the
setal organ (Fig. 7FG). Epiproct and paraproct of female
with some conspicuous cylinder setae (Fig. 7F). Setae
of male epiproct and paraproct normal, but paraproct
ventrally pointed due to the presence of a conspicuous
posteroventral sclerotized ridge which is ventrally
linked to ne membranes (Fig. 7G); depending on slide-
mounting this structure may appear like a free-ending
appendix (see Badonnel, 1967: g. 58; Mockford,
1993: g. 467; Lienhard, 1998: g. 36j). Hypandrium
and phallosome as in Fig. 7H, hypandrium shorter
than its basal width, phallic cradle well differentiated;
longitudinal struts in middle connected to each other by
a V-shaped sclerite, pores on apical part of longitudinal
struts present but not visible in Fig. 7H (only visible
in strongly squashed genitalia; see Badonnel, 1967:
g. 57). Spermatheca as in Fig. 7D, with a sclerotized
globular vesicle near duct, spermapore lacking
sclerotization, spermathecal sac with a narrowed zone
towards opening of duct, spermatophore about pear-
shaped.
Biology: In relatively warm regions, P. lachlani is
often domicole, as was the type population from
southern Italy (Ribaga, 1899); in the tropics it is also
regularly found in ground litter and sometimes in
caves (Lienhard, 2002). Brèthes (1923) and Townsend
(1912) recorded the species indoors (as P. astizi and
P. termitorum respectively; see Discussion, below), the
latter especially in houses infested with termites. The
presence of one “fresh” spermatophore and debris of
about three “old” spermatophores in the female gured
(Fig. 7D) indicates that the species is polyandrous.
Distribution: P. lachlani is widespread (Lienhard &
Smithers, 2002). Previously published South American
records can be found in the following references:
Townsend, 1912 (Peru, as P. termitorum); Brèthes,
1923 (Argentina, as P. astizi); New, 1984 (Brazil);
Mockford, 1993 (Colombia); García Aldrete &
Mockford, 2009 (Brazil). In adddition to the Brazilian
specimens mentioned above, some unpublished
material from Paraguay (MHNG) was also examined.
Other specimens examined (MHNG) are from the
following countries: Guatemala, Guadeloupe, Jamaica,
Ascension Island, Cabo Verde, Madeira, Liberia, Togo,
Rwanda, Mozambique, Morocco, Israel, Reunion, India,
Malaysia, Singapore, Indonesia, Thailand, Fiji.
Discussion: The detailed descriptions and the forewing
gures given by Townsend (1912) and Brèthes (1923)
for P. termitorum and P. astizi respectively t very well
the widely distributed P. lachlani; therefore we propose
here to consider these two names as junior synonyms of
the latter.
Under the synonym P. microps (Enderlein), Badonnel
(1967: p. 27) gives excellent gures of the forewing
and the male terminalia of P. lachlani, but it has to be
noted that in the legend (not in the text!) his gure 58 is
erroneously assigned to P. pilipennis.
Psocathropos pilipennis (Enderlein, 1931)
Fig. 8
Gambrella pilipennis Enderlein, 1931: 221.
Psocathropos pilipennis (Enderlein); Menon, 1942: 35.
Brazilian cave psocids 131
Fig. 7. Psocathropos lachlani, from Brazilian caves. (A) Forewing, female. (B) Hindwing, female. (C) Left ovipositor valvulae and
left hind corner of clunium. (D) Spermatheca containing one “fresh” spermatophore and debris of three “old” spermatophores.
(E) Pedicel with microspades organ, female. (F) Right paraproct, female. (G) Epiproct and paraprocts, male. (H) Hypandrium
and phallosome, ventral view (pilosity not shown).
132 C. Lienhard & R. L. Ferreira
Material examined: ISLA and MHNG; 4 males and
4 females (slide-mounted and in alcohol), collected
by R. L. Ferreira in caves situated in the following
Brazilian municipalities. – Ubajara (CE), Gruta do
Araticum cave, 1.i.2007. – Coronel José Dias (PI), Toca
do Garrincho cave, 11.ix.2008. – Felipe Guerra (RN),
Gruta Roncador cave, 1.vi.2006. – Jandaíra (RN), Gruta
Aroeira cave, 11.i.2006.
Further description: This species is always
brachypterous. For general descriptions see Enderlein
(1931: female) and Badonnel (1967: male, see also
Discussion below); some important characters are also
mentioned in the key given by Mockford (1991) and
in Mockford (1993: p. 51). Here we note the following
characters (Fig. 8, see also Key). Body length 1.4-
1.6 mm. Colouration similar to P. lachlani, but usually
less pigmented, sometimes hypodermal pigment almost
absent. Forewing slightly tapering towards apex, usually
almost reaching tip of abdomen or slightly surpassing it
(especially in males); venation of forewing somewhat
Fig. 8. Psocathropos pilipennis, from Brazilian caves. (A) Forewing, male (pilosity not shown). (B) Hindwing, male, showing venation
and insertion points of setae. (C) Right paraproct, female. (D) Spermathecal sclerotizations and proximal part of spermathecal
duct, specimen MHNG 8148 from Toca do Garrincho, PI. (E) Spermatheca (duct incomplete) containing one spermatophore,
specimen MNHG 8150 from Gruta do Araticum, CE (sclerotizations less squashed than in D). (F) Hypandrium and phallosome,
ventral view (pilosity not shown).
Brazilian cave psocids 133
variable, but Rs always bifurcate (Fig. 8A). Hindwing
strongly shortened but always with some clearly
differentiated veins and several long marginal setae
(Fig. 8B). P2 and pedicel as in P. lachlani. Epiproct and
paraproct simple in both sexes (i.e. no cylinder setae in
female and no paraproctal posteroventral ridge in male),
anal spine replaced by a normal seta, setal organ as in
P. lachlani (Fig. 8C). Hypandrium and phallosome as in
Fig. 8F, hypandrium longer than its basal width, apically
truncate, phallic cradle not clearly differentiated;
longitudinal struts lacking V-shaped connecting sclerite,
apically with clearly visible pores. Spermatheca as in
Fig. 8E, sclerotizations near duct more complicated than
in P. lachlani and vesicle slightly elongate (Fig. 8DE).
Spermapore lacking sclerotization, spermathecal sac
oval, lacking narrowed zone towards opening of duct,
spermatophore elongately pear-shaped (Fig. 8E).
Biology: In houses (Menon, 1942; Badonnel, 1967;
Mockford, 1991), in ground litter and on low vegetation
(New, 1977), in caves.
Distribution: Published records are from the Seychelles
(Enderlein, 1931), India (Menon, 1942), Madagascar
(Badonnel, 1967) and the Aldabra Islands (New, 1977).
In addition to the Brazilian material, some specimens
from the following countries have also been examined
(MHNG): Honduras (in cave), Curaçao (in cave),
Singapore (ground litter and/or in houses), Malaysia
(on fungi in forest), Vietnam (ground litter), Thailand
(domicole?). The species seems to be widespread
in the tropics, though less common than P. lachlani.
However, it is not excluded that some published records
of the latter comprise also undiscovered material of
P. pilipennis (e.g. Menon, 1942; see Discussion below).
Badonnel (1967) also indicates that both species were
taken together in a house in Madagascar.
Discussion: Menon (1942) synonymizes the genus
Gambrella Enderlein with Psocathropos Ribaga and
gives a description of its type species, P. pilipennis.
However, his material apparently consisted of a
mixture of both species, P. pilipennis and P. lachlani.
Therefore he mentions a variability of wing morphology
(hindwing sometimes “totally wanting” and forewing
sometimes relatively short with simple Rs, see gures
25-26) which we have never observed in P. pilipennis.
In our opinion, his gures 25 and 26 illustrate forewings
of P. lachlani, while his gures 23, 24 and 27 illustrate
wings of P. pilipennis, and his gure 28 the male
genitalia of the latter. Badonnel (1967) gives excellent
gures of the wings and the male genitalia, but it has to
be noted that his gure 58 does not illustrate epiproct
and paraproct of P. pilipennis, as erroneously indicated
in the legend, but of P. lachlani (= P. microps), as
correctly indicated in the text on the latter species
(p. 27).
Key to the South American species of Psyllipsocidae and Prionoglarididae
This key allows species identi cation of adult South American psocids belonging to the families Psyllipsocidae and
Prionoglarididae. In addition to the characters mentioned in the key, these species are characterized among Psocoptera
by their 3-segmented tarsi, the absence of scales on body and wings, the absence of a thickened and distinctly opaque
pterostigma (sometimes slightly opaque in Prionoglarididae) and the presence, in the forewing, of a simple anal vein and
a nodulus (i.e. veins CuP and A1 joining at wing margin). Their body is never dorsoventrally attened and, if present, their
forewings are membranous, never coriaceous, though sometimes slightly vaulted (elytriform).
For abbreviations see Material and Methods, above.
1 Forewing normally developed (macropterous) or slightly shortened, usually extending well beyond tip of abdomen,
not tapering towards apex but broadly rounded, with normal venation ...................................................................2
Apterous, micropterous or clearly brachypterous (forewing usually not reaching tip of abdomen or tapering towards
apex), venation at least slightly reduced ...................................................................................................................3
2 Forewing: basal portion of Sc developed as a slightly curved vein joining R1 just basally of pterostigma (Lienhard
& Ferreira, 2013a: g. 1a). Body length: 2.5 to 3.7 mm ......................................... Prionoglarididae: Neotrogla 11
Forewing: basal portion of Sc rudimentary, ending in membrane, not joining R and usually not extending beyond
base of basal closed cell (Fig. 2A). Body length: 1.0 mm (Psyllipsocus didymus) to 2.7 mm (P. serrifer), mostly
about 1.5 mm .................................................................................................................................Psyllipsocidae 14
3 Apterous, without wing rudiments; ocelli completely reduced ................................................................................ 4
Micropterous or brachypterous ................................................................................................................................. 5
4 Pretarsal claw without preapical tooth. Eye minute, consisting of a single ommatidium. Paraproct without anal
spine ..........................................................................................................Prionoglarididae: Speleopsocus chimanta
Note: This species is a troglobite known from a cave in south eastern Venezuela (see Lienhard et al., 2010a).
Pretarsal claw with a small preapical tooth. Eye differentiated as a small compound eye, with several ommatidia.
Paraproct with a long anal spine ..............................................................................................Psyllipsocus ramburii
134 C. Lienhard & R. L. Ferreira
Note: This is the rare (or potential?) apterous form of P. ramburii, corresponding to P. dubius Badonnel which
is here considered as a junior synonym of the latter (see Discussion of P. ramburii, above). For micropterous,
brachypterous and macropterous forms, see couplets 5, 10 and 15.
5 Forewing at or slightly vaulted (elytriform), reaching posterior half of abdomen and bearing some clearly
differentiated veins ................................................................................................................................................... 6
Forewing strongly reduced, just reaching basal abdominal segments and lacking clearly differentiated veins
(Fig. 2E). Paraproct with a long anal spine .............................................................................Psyllipsocus ramburii
Note: This is the micropterous form of P. ramburii; for apterous, brachypterous and macropterous forms, see
couplets 4, 10 and 15.
6 Forewing margin with long setae (their length several times width of marginal vein) (Fig. 7A). Paraproct lacking
a conspicuous anal spine (Figs 7FG, 8C) ......................................................................................... Psocathropos 7
Forewing margin bare (except for some microscopic hairs which are much shorter than width of marginal vein).
Paraproct with a long anal spine (Lienhard & Ferreira, 2014: g. 1F) .............................................. Psyllipsocus 8
7 Forewing Rs simple (Fig. 7A); hindwing strongly reduced, bare and veinless (Fig. 7B). Hypandrium shorter than
its basal width (Fig. 7H). Male paraproct ventrally pointed, with a posteroventral sclerotized ridge (Fig. 7G).
Female paraproct and epiproct with some conspicuous cylinder setae (Fig. 7F) .................. Psocathropos lachlani
Note: This is the brachypterous form of P. lachlani; for the rare macropterous form, see couplet 14.
Forewing Rs bifurcate (Fig. 8A); hindwing much shortened but with some clearly differentiated veins and some
long marginal setae (Fig. 8B). Hypandrium longer than its basal width (Fig. 8F). Male paraproct as in female
(Fig. 8C), ventrally rounded, lacking sclerotized ridge. Female paraproct and epiproct lacking cylinder setae .......
............................................................................................................................................. Psocathropos pilipennis
Note: Macropterous form unknown in this species.
8 Forewing with a brown transversal band about in middle (Lienhard & Ferreira, 2014: g. 1C) ...............................
...................................................................................................................................................Psyllipsocus spinifer
Note: For the much rarer macropterous form of this species, see couplet 22.
– Forewing unmarked ..................................................................................................................................................9
9 P2 with a stout sensillum in basal half (Lienhard & Ferreira, 2013b: g. 4F) ....................Psyllipsocus clunjunctus
Note: Macropterous form of this species not known.
Stout P2-sensillum not differentiated (Fig. 2D) .....................................................................................................10
10 Forewing slightly vaulted (elytriform), hindwing a tiny membranous ap only reaching hind margin of rst
abdominal tergite ...................................................................................................................Psyllipsocus delamarei
Note: This species is only known from one female from Argentina (see Badonnel, 1962); it seems to be very close
to the brachypterous form of P. ramburii.
Forewing not elytriform, hindwing with some veins, not much shorter than half length of forewing ......................
.................................................................................................................................................Psyllipsocus ramburii
Note: This is the brachypterous form of P. ramburii; for apterous, micropterous and macropterous forms, see
couplets 4, 5 and 15.
11 Female: distal part of gynosome strongly curved (Lienhard & Ferreira, 2013a: g. 1d) ..............Neotrogla curvata
Female: distal part of gynosome straight or only slightly curved (Lienhard et al., 2010b: gs 2df, 8c) ...............12
12 Female: membranous part of gynosome not lobate (Lienhard et al., 2010b: g. 8c) .................. Neotrogla truncata
Female: membranous part of gynosome with a spiny dorsal lobe and a pair of spiny lateral lobes (Lienhard et al.,
2010b: g. 2f) .........................................................................................................................................................13
13 Female: distal lobe of subgenital plate (egg-guide) apically with a large bare median lobe and a pair of minute
pilose lateral lobes (Lienhard et al., 2010b: g. 1c) ............................................................... Neotrogla brasiliensis
Female: pilose lateral lobes of egg-guide not much smaller than bare median lobe (Lienhard et al., 2010b: g. 5)
........................................................................................................................................................Neotrogla aurora
14 Forewing and hindwing with numerous long marginal setae (their length several times width of marginal vein)
(Mockford, 1993: gs 462, 463) ............................................................................................. Psocathropos lachlani
Note: This is the rare macropterous form of P. lachlani; for the brachypterous form, see couplet 7.
Wing margins bare (except for some microscopic hairs which are shorter than width of marginal vein) ............. 15
15 Distal closed cell in forewing absent (i.e. no crossvein between R1 and Rs) (Fig. 2A) .........Psyllipsocus ramburii
Note: This is the relatively rare macropterous form of P. ramburii; for apterous, micropterous and brachypterous
forms, see couplets 4, 5 and 10.
Distal closed cell present, distally delimited by a crossvein between R1 and Rs (Fig. 4G) ..................................16
16 In forewing Rs and M joined by a crossvein (Lienhard & Ferreira, 2014: g. 3A) ...............................................17
In forewing Rs and M fused for a length (Fig. 4G) ................................................................................................ 19
17 Forewing unmarked ................................................................................................................... Psyllipsocus falcifer
Brazilian cave psocids 135
Forewing with some pigmented areas ....................................................................................................................18
18 AP at, its marginal length more than twice its height, patches of wing pattern associated to veins (Lienhard &
Ferreira, 2014: g. 5A) ............................................................................................................Psyllipsocus marconii
AP tall, its marginal length not much exceeding its height, an isolated longitudinal patch present in middle of cells
r1, r3 and r5 (Lienhard & Ferreira, 2014: g. 7A) ..................................................................... Psyllipsocus thaidis
19 First portion of pterostigmal R1 not longer than half length of R1-Rs crossvein (Fig. 4G) ..................................20
First portion of pterostigmal R1 clearly longer than half length of R1-Rs crossvein, often longer than this
crossvein .................................................................................................................................................................22
20 Distal closed cell almost twice as long as marginal length of pterostigma; paraproct with a long, basally non-
articulated anal spine (Lienhard & Ferreira, 2013b: gs 4A, 6B) ..........................................................................21
Distal closed cell only about half as long as marginal length of pterostigma (Fig. 4G); anal spine of paraproct
replaced by a stout, relatively short and basally articulated seta (Fig. 4BF) ............................ Psyllipsocus yucatan
21 Female: spermatheca with a conspicuous longitudinal saw-like sclerite (Lienhard & Ferreira, 2013b: g. 7J). Male:
abdominal apex with a complex clunial bridge, hypandrium with four apical setae (Lienhard & Ferreira, 2013b:
g. 5) .......................................................................................................................................... Psyllipsocus serrifer
Female: spermatheca with a slender le-like sclerite (Lienhard & Ferreira, 2013b: g. 8F). Male: abdominal apex
with a simple clunial bridge, hypandrium with two apical setae (Lienhard & Ferreira, 2013b: g. 9) .....................
.................................................................................................................................................... Psyllipsocus similis
22 Forewing and hindwing with a brown transversal band in basal half (Lienhard & Ferreira, 2014: g. 1AB) ..........
...................................................................................................................................................Psyllipsocus spinifer
Note: This is the relatively rare macropterous form of this species; for the brachypterous form, see couplet 8.
Wings unmarked or wing pattern different ............................................................................................................. 23
23 Forewing particularly narrow, FW/FWw > 3 ......................................................................................................... 24
Forewing of normal shape, FW/FWw 3 .............................................................................................................. 25
24 Forewing with narrow pigment bands bordering most of veins (Lienhard & Ferreira, 2014: g. 16A). Male:
postero-median V-shaped incision of phallosome not deeper than length of antero-median part of phallosome
sclerite (Lienhard & Ferreira, 2014: g. 17AB) ..............................................................Psyllipsocus angustipennis
Pigment bands broader in apical half of forewing, covering more than half of the surface of cells m1 and m2
(Lienhard & Ferreira, 2014: g. 17C). Male: postero-median V-shaped incision of phallosome deeper than length
of antero-median part of phallosome sclerite (Lienhard & Ferreira, 2014: g. 17F) ............. Psyllipsocus proximus
25 Forewing anal vein with a brown spot at the base of each hair; exceptionally forewings completely unmarked (see
pale specimens of P. subtilis, couplet 28) ............................................................................................................... 26
Forewing anal vein lacking such spots; distal end of forewing veins bordered by brown pigment (Lienhard &
Ferreira, 2014: gs 14A, 15A) ..................................... Psyllipsocus punctulatus (and female spec. cf. punctulatus)
26 Height of AP exceeding its marginal length; forewing with a conspicuous U-shaped or V-shaped brown patch
covering most of cell r1 (Lienhard & Ferreira, 2014: g. 13A) ..........................................Psyllipsocus radiopictus
AP lower than its marginal length; pigmentation of cell r1 different .....................................................................27
27 Forewing with a broad brown band bordering apical portion of R1 and R1-Rs crossvein, forming a conspicuous
angulate patch which borders cell r1 basally (Lienhard & Ferreira, 2014: g. 12A) .................................................
.............................................................................................................................................Psyllipsocus fuscistigma
At most narrow pigment bands bordering apical portion of R1 and R1-Rs crossvein ........................................... 28
28 Forewing with four narrow pigment stripes parallel to veins R2+3 and R4+5, situated in cells r1 (one stripe), r3
(two stripes) and r5 (one stripe); these stripes clearly separated from the narrow pigmented zones bordering these
veins. In very pale specimens forewings unmarked. Male: clunium simple, lacking clunial rods. Female: v3 with
a marginal row of thick setae, clearly thicker than other v3-setae of similar length (Lienhard & Ferreira, 2014: g.
11AE) ......................................................................................................................................... Psyllipsocus subtilis
Forewing with a dark brown drop-shaped spot or stripe or a light brown diffuse patch in middle of cells r1 and r3.
Male: clunial rods present. Female: v3-pilosity uniform .......................................................................................29
29 Forewing with a dark brown spot or stripe in middle of cells r1 and r3. Male: phallic cradle of hypandrium
anteriorly broadly rounded (Lienhard & Ferreira, 2014: gs 8, 9) ................................Psyllipsocus clunioventralis
These cells only with a light brown diffuse patch covering most of their surface. Male: phallic cradle of hypandrium
anteriorly truncate (Lienhard & Ferreira, 2014: g. 10) ...........................................................Psyllipsocus didymus
136 C. Lienhard & R. L. Ferreira
GENERAL DISCUSSION
Analysis of distribution
This analysis is based on the data presented in Appendix
2. Detailed considerations concerning each of the 19
endemics (15 Psyllipsocus spp., 4 Neotrogla spp.) have
already been published by Lienhard et al. (2010b) and
Lienhard & Ferreira (2013a, b, 2014) when describing
these previously unknown species. During this
project, about 400 adult individuals of 23 species of
Psyllipsocidae and Prionoglarididae from 124 Brazilian
caves were identi ed (Table 1). The total number
of caves wherein each species was recorded and the
number of caves wherein it was associated with another
species of these families are presented in Table 2. The
proportion of Brazilian caves occupied by these families
is unknown; because of uneven sampling effort their
absence cannot be assumed for caves where they are not
currently recorded. This discussion only concerns caves
with identi ed adults; caves with nymphs that cannot be
identi ed are not included (see also Introduction).
As previously observed by Lienhard & Ferreira (2014) and
Yoshizawa et al. (2014), these insects live predominantly
in dry resource-poor caves. Two closely related species
of Psyllipsocus or two species of Neotrogla have never
been found in the same cave. In most of the caves only
one species was present, rarely two species (23 caves),
exceptionally three species (viz. Gruta de Ubajara, CE:
Psoc. lachlani, P. spinifer, P. yucatan; Gruta Janelão, MG:
N. brasiliensis, P. serrifer, P. spinifer; Gruta do Lagedo
Grande, RN: P. clunjunctus, P. spinifer, P. subtilis). In
most caves with two or three species, only one of them is
a Brazilian endemic; in seven caves two endemics were
found and in two caves three endemics live together
(Gruta Janelão and Gruta do Lagedo Grande, see above).
In three caves (cave PEA 343, BA; Gruta Caboclo, MG;
Gruta Janelão, MG) specimens of both genera, Neotrogla
and Psyllipsocus, were present simultaneously. Among
the 15 endemics of Psyllipsocus, 8 species were recorded
from a single cave each (Table 2). Thus, very strict local
endemism appears to be high in this genus.
The cosmopolitan and often domicole species
P. ramburii is the most common cave Psyllipsocus in
Brazil (present in 30 caves, in 23 of them exclusively).
In about 40% of the caves at least one of the non-
endemics P. ramburii, P. yucatan, Psoc. lachlani or Psoc.
pilipennis is present; 32% of the caves are exclusively
inhabited by non-endemics. The high frequency of the
opportunistic, parthenogenetic (thelytokous) and often
domicole P. ramburii indicates that this species is a
particularly good disperser; but it may sometimes have
been introduced by human activities.
In all 9 caves with two or three Brazilian endemics at
least one of them is P. spinifer or P. clunjunctus. Together
with P. serrifer, these are the most widely distributed
endemics. P. spinifer is known from 20 caves (19
municipalities, 8 states), P. clunjunctus from 16 caves (9
municipalities, 5 states) and P. serrifer from 13 caves (9
municipalities, 2 states) (for the distribution of the latter
two species, see also map in Lienhard & Ferreira, 2013b).
P. spinifer is very often associated with other species,
being found alone in only ve caves. It is interesting to
see that the non-domicole, non-parthenogenetic endemic
P. spinifer, the second most common Brazilian cave
species, is much more often associated with a probably
more specialized local endemic than the most common,
cosmopolitan, domicole and parthenogenetic P. ramburii
(Table 2). It cannot be excluded that the presence of this
opportunistic generalist contributes to the elimination
of locally endemic specialists. Only in two of 30 caves
P. ramburii has been recorded together with a Brazilian
endemic (P. falcifer, P. serrifer).
Some evolutionary considerations
This rather speculative discussion follows the short
remarks that we made in the recent paper wherein most of
the endemic cave species of Psyllipsocus were described
(Lienhard & Ferreira, 2014). Rather than as a discussion
of concrete results we consider this part as an outlook
on possible future research perspectives concerning the
fascinating cave psocids of the families Psyllipsocidae
and Prionoglarididae.
The only clearly troglobitic representative of these
families from the New World is the prionoglaridid
Speleopsocus chimanta known from a humid cave in
south eastern Venezuela (see Key and Appendix 1).
All other species are winged, have well developed
compound eyes and are relatively well-pigmented,
except for the cosmopolitan and often domicole pale and
Table 1. Species richness, rate of endemism and distribution for Brazilian genera of Psyllipsocidae and Prionoglarididae based on adult
specimens collected in caves.
species endemics caves municipalities states
Psocathropos 213128
Psyllipsocus 17 15 (88%) 102 51 13
Neotrogla 4 4 (100%) 19 9 3
Total 23 19 (83%) 124 59 13
Brazilian cave psocids 137
Table 2. Brazilian cave species of Psyllipsocidae (Psocathropos, Psyllipsocus) and Prionoglarididae (Neotrogla) with total number of caves wherein adults of each species were collected
and, on corresponding line or column, number of caves where the species was associated with another one.
species
total
P. lachlani
P. pilipennis
P. ramburii
P. yucatan
P. angustipennis
P. clunioventralis
P. clunjunctus
P. didymus
P. falcifer
P. fuscistigma
P. marconii
P. proximus
P. punctulatus
P. radiopictus
P. serrifer
P. similis
P. spinifer
P. subtilis
P. thaidis
N. aurora
N. brasiliensis
N. curvata
non-endemics Psocathropos lachlani 922 1 2
Psocathropos pilipennis 41
Psyllipsocus ramburii 30 22 1 12
Psyllipsocus yucatan 13 22 1 2 1
endemics Psyllipsocus angustipennis 3
Psyllipsocus clunioventralis 11
Psyllipsocus clunjunctus 16 32
Psyllipsocus didymus 1
Psyllipsocus falcifer 811
Psyllipsocus fuscistigma 11
Psyllipsocus marconii 11
Psyllipsocus proximus 1
Psyllipsocus punctulatus 11
Psyllipsocus radiopictus 11
Psyllipsocus serrifer 13 111
Psyllipsocus similis 3
Psyllipsocus spinifer 20 2122 13 11 1 1 1 2
Psyllipsocus subtilis 621
Psyllipsocus thaidis 1
Neotrogla aurora 6
Neotrogla brasiliensis 312
Neotrogla curvata 71
Neotrogla truncata 3
138 C. Lienhard & R. L. Ferreira
wingless forms of Psyllipsocus ramburii, with reduced
but still multifacetted eyes (see Key). The only character
interpreted as a cave adaptation (Lienhard, 2000) is
the presence of leg trichobothria in the prionoglaridid
subfamiliy Speleketorinae, to which the Brazilian genus
Neotrogla belongs (Lienhard, 2004a; Lienhard et al.,
2010b). This genus is only known from caves. However,
most of the Psyllipsocus species known from North and
Middle America live on the bark of trees, in soil litter
or on lichen-covered rock outcrops (Mockford, 2011).
Unfortunately, all available data on Brazilian Psyllipsocus
concern material collected in caves, though some species
of this genus might also live in other microhabitats in
Brazil, outside of caves, as in the northern parts of the
New World. It would be of particular interest to discover
non-cavernicolous populations of the endemic species or
of species closely related to them.
Most of the local endemics are only known from one
cave or from some caves situated close together (often in
the same municipality or neighbouring municipalities).
Some isolated relict populations of formerly more
widely distributed species might be at the origin of this
high diversity of cave endemics. These ancestors were
probably not strictly cavernicolous. The relatively wide
distribution of P. spinifer, P. clunjunctus and P. serrifer
may be the result of secondary dispersal of these
endemics out of their region of origin.
The endemic cave Psyllipsocus form a heterogeneous
assemblage of isolated species or small groups of
species. However, except for the most common endemic
P. spinifer, they all show some kind of reduction of the
phallosome, apparently due to homoplasy (Lienhard
& Ferreira, 2014) (Note: males of P. thaidis and
P. radiopictus are not known). All known Brazilian
prionoglaridids (i.e. the four species of the endemic cave
genus Neotrogla) show also a strong reduction of the
phallosome (Lienhard et al., 2010b; Lienhard & Ferreira,
2013a; Yoshizawa et al., 2014). Therefore we proposed
the speculative hypothesis that the locally endemic
cave species of these genera might have evolved under
a particular selection pressure favouring reduction of
male primary external genitalia (Lienhard & Ferreira,
2014). This selection pressure could be similar to that
favouring reduction of the phallosome in the African
cave prionoglaridid Afrotrogla, known from caves
in semidesertic regions of Namibia and South Africa
(Lienhard, 2007). All these psocids live in dry resource-
poor caves and are the only known representatives of the
suborder Trogiomorpha with reduced phallosomes. For
the genus Neotrogla, Yoshizawa et al. (2014) showed
that this reduction of the male intromittent organ is
very probably related to reversed sexual selection. The
only psocid species where reversed sexual selection has
clearly been documented at present, Lepinotus patruelis,
also belongs to the suborder Trogiomorpha but is not
cavernicolous and has normal male genitalia (Wearing-
Wilde, 1995, 1996; Yoshizawa et al., 2014). It would be
interesting to elucidate the in uence of a subterranean
mode of life in dry caves on sexual selection. Miller &
Svensson (2014: p. 427) “suggest that deeper ecological
perspectives on sexual selection may alter some of the
fundamental assumptions of sexual selection theory and
rapidly lead to new discoveries”. Prionoglarididae and
Psyllipsocidae could play a key role in future research
in this eld. We suggest that adaptations to life in dry,
resource-poor cave environments could affect sexual
selection and thus promote sexual role reversal in these
insects.
However, the absence of data on non-cavernicolous
representatives of these families from Brazil makes it
impossible to estimate the pertinence of the supposed
correlation between a subterranean mode of life and the
presence of reduced male genitalia. The most widely
distributed endemic cave species is P. spinifer, the only
endemic with a normal male phallosome, like that of
all non-cavernicolous species of the genus. Thus, this
species may also live outside of caves in Brazil. For
the local endemics with reduced phallosome it may
be speculated that they evolved as cave refugees from
populations which were not competitive under out-of-
cave conditions.
In the trogiomorphan species with the most striking
reduction of the phallosome (i.e. all species of Neotro-
gla and Afrotrogla and of the Psyllipsocus clunjunctus
species group), females have evolved novel differentia-
tions of the spermapore region. Yoshizawa et al. (2014)
showed that, in Neotrogla, the novel female organ func-
tions as an intromittent organ during copulation. Such a
function was also suggested hypothetically for the novel
female organs in Afrotrogla (Lienhard, 2007) and the
Psyllipsocus clunjunctus group (Lienhard & Ferreira,
2013b). Among the Brazilian endemics with a reduced
phallosome there are two particularly successful spe-
cies, in terms of their distribution range, P. clunjunctus
and P. serrifer, both belonging to the clunjunctus spe-
cies group. They are not only characterized by the almost
complete absence of phallosome sclerites and the pre-
sence of a novel female “micropenis”, but they also have
evolved a novel male accessory genital organ, the clunial
bridge (Lienhard & Ferreira, 2013b). Might these novel
structures be the reason of their success?
It is dif cult to ascribe to pure coincidence these varied
phenomena relating to similar modi cations of genital
morphology and reproductive behaviour in species that
are not particularly closely related but which share a
cavernicole life history. Perhaps we are here confronted
with a new category of adaptations to a life in caves
different to the classical troglomorphic adaptations
(reduction of wings, pigmentation and eyes; see Dethier
& Hubart, 2005), and as yet, not directly explained.
Brazilian cave psocids 139
ACKNOWLEDGEMENTS
We thank Edward L. Mockford (Illinois State University,
Normal, Illinois, USA) and John Hollier (MHNG) for
critical reading of the manuscript and useful comments.
CL thanks John Hollier for interesting discussions,
Eliane De Coninck (Tervuren) for the loan of the type-
series of Psyllipsocus collarti and Manuela Lienhard
for composing the photo plate of Fig. 1. RLF is very
grateful to all members of the team from the Centro
de Estudos em Biologia Subterrânea (UFLA) for their
help in collecting material. Funding was provided by
the Conselho Nacional de Pesquisa (CNPq - process nº
477712/2006–1, and CNPq grant nr. 301061/2011–4)
and the Fundação de Amparo à Pesquisa do Estado de
Minas Gerais (FAPEMIG - processes CRA - APQ 01826-
08 and CRA - PPM-00433-11).
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APPENDIX 1: Checklist of South American Psyllipsocidae
and Prionoglarididae, with indication of general distribution
(only known from caves, except for Psocathropos spp.,
Psyllipsocus delamarei, P. ramburii and P. yucatan).
Psyllipsocidae
Psocathropos Ribaga, 1899: 156; type species: P. lachlani
Ribaga.
Psocathropos lachlani Ribaga, 1899: 157. Widely distributed in
the tropics (often domicole).
Psocathropos pilipennis (Enderlein, 1931: 221). Widely
distributed in the tropics (sometimes domicole).
Psyllipsocus Selys-Longchamps, 1872: 145; type species:
P. ramburii Selys-Longchamps.
Psyllipsocus angustipennis Lienhard, 2014; in: Lienhard &
Ferreira, 2014: 239. Brazil.
Psyllipsocus clunioventralis Lienhard, 2014; in: Lienhard &
Ferreira, 2014: 225. Brazil.
Psyllipsocus clunjunctus Lienhard, 2013; in: Lienhard &
Ferreira, 2013b: 424. Brazil.
Psyllipsocus delamarei Badonnel, 1962: 187. Argentina.
Psyllipsocus didymus Lienhard, 2014; in: Lienhard & Ferreira,
2014: 229. Brazil.
Psyllipsocus falcifer Lienhard, 2014; in: Lienhard & Ferreira,
2014: 216. Brazil.
Psyllipsocus fuscistigma Lienhard, 2014; in: Lienhard &
Ferreira, 2014: 233. Brazil.
Psyllipsocus marconii Lienhard, 2014; in: Lienhard & Ferreira,
2014: 220. Brazil.
Psyllipsocus proximus Lienhard, 2014; in: Lienhard & Ferreira,
2014: 242. Brazil.
Psyllipsocus punctulatus Lienhard, 2014; in: Lienhard &
Ferreira, 2014: 236. Brazil.
Psyllipsocus radiopictus Lienhard, 2014; in: Lienhard &
Ferreira, 2014: 234. Brazil.
Psyllipsocus ramburii Selys-Longchamps, 1872: 146.
Cosmopolitan (often domicole).
Psyllipsocus serrifer Lienhard, 2013; in: Lienhard & Ferreira,
2013b: 429. Brazil.
Psyllipsocus similis Lienhard, 2013; in: Lienhard & Ferreira,
2013b: 433. Brazil.
Psyllipsocus spinifer Lienhard, 2014; in: Lienhard & Ferreira,
2014: 212. Brazil.
Psyllipsocus subtilis Lienhard, 2014; in: Lienhard & Ferreira,
2014: 229. Brazil.
Psyllipsocus thaidis Lienhard, 2014; in: Lienhard & Ferreira,
2014: 223. Brazil.
Psyllipsocus yucatan Gurney, 1943: 212. Widely distributed in
the tropics.
Prionoglarididae
Neotrogla Lienhard, 2010; in: Lienhard et al., 2010b: 612; type
species: N. brasiliensis Lienhard.
Neotrogla aurora Lienhard, 2010; in: Lienhard et al., 2010b:
619. Brazil.
Neotrogla brasiliensis Lienhard, 2010; in: Lienhard et al.,
2010b: 614. Brazil.
Neotrogla curvata Lienhard & Ferreira, 2013a: 4. Brazil.
Neotrogla truncata Lienhard, 2010; in: Lienhard et al., 2010b:
622. Brazil.
Brazilian cave psocids 141
Speleopsocus Lienhard, 2010; in: Lienhard et al., 2010a: 186;
type species: S. chimanta Lienhard.
Speleopsocus chimanta Lienhard, 2010; in: Lienhard et al.,
2010a: 187. Venezuela.
APPENDIX 2: Comprehensive list of Brazilian caves wherein
adults of Psyllipsocidae and Prionoglarididae were collected
(leg. R. L. Ferreira unless other collector mentioned)
P. = Psyllipsocus; Psoc. = Psocathropos; in bold type =
non-endemic species (widely distributed in the tropics or
cosmopolitan)
Alagoas (AL)
Delmiro Gouveia (AL), Gruta Morcego, 23.iv.2006: Psoc.
lachlani
Murici (AL), Toca da Raposa 1 (granite), 13.i.2007: P.
radiopictus, P. yucatan
Murici (AL), Toca da Raposa 2 (granite), 13.i.2007: P. yucatan
Amazonas (AM)
Apuí (AM), Gruta Apiacá 1, Parque Nacional do Juruena,
13.ix.2011: P. proximus
Bahia (BA)
Campo Formoso (BA), Toca do Angico, 9.i.2008: P. spinifer
Campo Formoso (BA), Toca do Morrinho, i.1997: P. spinifer
Campo Formoso (BA), Toca do Pitu, 10.vii.2008: Neotrogla
truncata
Campo Formoso (BA), Toca da Tiquara, 8.i.2008, i.2009,
xii.2010: P. yucatan
Curaçá (Patamuté) (BA), Toca d’agua de Patamuté, 6.i.2008:
Psoc. lachlani, P. spinifer
Ourolândia (BA), Toca dos Ossos, 31.vii.2007, 10.vi.2012,
i.2013: Neotrogla truncata
Palmeiras (BA), Gruta Ioiô, 1.i.2008: Neotrogla truncata
Santa Maria da Vitória (BA), cave PEA 341, x.2012: Neotrogla
curvata
Santa Maria da Vitória (BA), cave PEA 342, x.2012: Neotrogla
curvata
Santa Maria da Vitória (BA), cave PEA 343, 15.v.2011, leg. S.
S. Salgado: Neotrogla curvata, P. yucatan
São Desidério (BA), Gruta do Catitu, 24.vii.200: P. clunjunctus
São Desidério (BA), Gruta do Sumidouro do João Baio,
29.vii.2006: P. clunjunctus, P. spinifer
São Félix do Coribe (BA), cave PEA 377, 10.v.2011, leg. S. S.
Salgado: P. serrifer
São Félix do Coribe (BA), cave PEA 378, x.2012: Neotrogla
curvata
São Félix do Coribe (BA), cave PEA 380, 21.vii.2011, leg. S. S.
Salgado: Neotrogla curvata
São Félix do Coribe (BA), cave PEA 381, 18.vii.2011, leg. S. S.
Salgado: Neotrogla curvata
São Félix do Coribe (BA), cave PEA 383, 18.vii.2011, leg. S. S.
Salgado: Neotrogla curvata
Várzea Nova (BA), Gruta Jurema, 20.vii.2008: P. clunjunctus
Ceará (CE)
Araripe (CE), Gruta do Brejinho, 1.v.2007: P. spinifer
Tejuçuoca (CE), Gruta do Veado Campeiro, 16.ix.2008: P.
fuscistigma, P. spinifer
Ubajara (CE), Gruta do Araticum, 1.i.2007: Psoc. pilipennis,
P. spinifer
Ubajara (CE), Gruta dos Mocós, 3.i.2007: P. yucatan
Ubajara (CE), Gruta do Morcego Branco, 3.i.2007: P. spinifer,
P. yucatan
Ubajara (CE), Gruta de Ubajara, 30.xii.2006: Psoc. lachlani, P.
spinifer, P. yucatan
Espírito Santo (ES)
Castelo (ES), Gruta do Limoeiro, 7.i.2005: P. ramburii, P.
yucatan
Santa Teresa (ES), Gruta do André Huscki, 4.i.2005: Psoc.
lachlani
Goiás (GO)
Damianópolis (GO), Lapa do Ribeirão dos Porcos, 29.vii.2001,
5.x.2001, 27.vi.2002: P. clunjunctus, P. spinifer
Minas Gerais (MG)
Arcos (MG), Caverna do Alinhamento, 1.vi.2002: P. ramburii
Arcos (MG), Gruta da Bocaininha, 3.xii.2008: P. serrifer
Arcos (MG), Gruta Labirinto, 28.i.2006: P. serrifer
Arinos (MG), Lapa do Salobo, 18.vii.2010: P. yucatan
Cordisburgo (MG), Gruta de Maquiné, 10.vii.2000, 18.v.2010:
Psoc. lachlani, P. ramburii
Cordisburgo (MG), Gruta do Salitre, 22.iv.2011: P. serrifer
Cordisburgo (MG), Gruta Santo Amaro 1, 28.ix.2010: P.
ramburii
Cordisburgo (MG), Gruta Tão Lucas, 14.xi.2010: P. falcifer
Cordisburgo (MG), Lapinha do Atamis, 13.xi.2010: P. falcifer
Coromandel (MG), Gruta João do Pó, 5.x.2000: P. ramburii
Coromandel (MG), Gruta Ronan, 3.x.2000: P. ramburii
Coromandel (MG), Gruta Ronan II, 4.x.2000: P. ramburii
Diamantina (MG), Gruta do Salitre, 13.xii.2007: P. ramburii
Doresópolis (MG), Gruta P43, 9.xi.2003: P. serrifer
Itabirito (MG), Gruta MP1, 29.viii.2005: P. serrifer
Itabirito (MG), Gruta MP8, 8.ix.2005: P. similis
Itacarambi (MG), Gruta Bonita, 19.iii.2003: P. angustipennis
Itambé do Mato Dentro (MG), Baixada dos Crioulos 2,
29.vii.2004: P. similis
Itumirim (MG), Gruta Santo Antônio, 25.x.2002: P. ramburii
Januária (MG), Gruta Caboclo, 27.vii.2003: Neotrogla
brasiliensis, P. spinifer
Januária (MG), Gruta Ossos, vii.2003, iii.2013: Neotrogla
brasiliensis
Januária / Itacarambi (MG), Gruta Brejal, 25.vii.2003: female
cf. P. punctulatus
Januária / Itacarambi (MG), Gruta Janelão, 28.vii.2003:
Neotrogla brasiliensis, P. serrifer, P. spinifer
Januária / Itacarambi (MG), Gruta Preguiça, 26.vii.2003: P.
angustipennis
João Pinheiro (MG), Gruta do Sapecado, 15.x.2010: P. ramburii
Lagoa da Prata (MG), Gruta Salão de Festas, 4.v.2003: P.
serrifer
Lagoa Santa (MG), Gruta da Lapinha, 12.vi.2002: P. ramburii
Matozinhos (MG), Gruta dos Irmãos Piriás, 28.vii.2000: P.
ramburii
Matozinhos (MG), Gruta Lavoura, 28.+ 29.v.1997, 26.vi.1997,
19.ii.2000: P. ramburii
Matozinhos (MG), Gruta Pequenas III, 31.vii.2002: P. serrifer
Matozinhos (MG), Meandro Abismante, 6.vii.2002: P. ramburii
Matutina (MG), Gruta 9, 10.x.2010: P. ramburii
Moeda (MG), SMS 19, 3.xii.2005: P. similis
142 C. Lienhard & R. L. Ferreira
Moeda (MG), SMS 29, 11.xii.2005: P. ramburii
Montalvânia (MG), Gruta Nossa Senhora do Perpétuo Socorro,
14.vii.2007: Psoc. lachlani, P. marconii
Nacip Raydan (MG), Gruta Manga de Pedra (granito),
21.vii.2002: Psoc. lachlani
Pains (MG), Buraco do Nando, 12.x.2003: P. serrifer
Pains (MG), Gruta Brasical, 28.ix.2003: P. serrifer
Pains (MG), Gruta do Capão, 5.v.2001: P. ramburii
Pains (MG), Gruta dos Estromatólitos, 7.xi.2000: P. falcifer
Pains (MG), Gruta Paiol de Milho, 13.x.2003: P. falcifer, P.
spinifer
Pains (MG), Gruta Paranoá, 15.i.2008: P. ramburii, P. serrifer
Pains (MG), Gruta Retiro, 27.xi.1999: P. ramburii
Pains (MG), Gruta Ronco, 28.xi.1999: P. falcifer
Pains (MG), Gruta do Sobradinho, 5.v.2001: P. serrifer
Paracatu (MG), Gruta da Fazenda Tamanduá II, 14.x.2010: P.
ramburii
Paracatu (MG), Lapa do Brocotó, 16.ix.2010: P. ramburii
Paracatu (MG), Lapa de Santo Antônio, 13.vii.2010: P.
ramburii
Presidente Olegário (MG), Lapa Vereda da Palha, 13.x.2010:
P. clunjunctus
Santa Maria do Suaçuí (MG), Gruta do Rio Suaçuí, 19.vii.2002:
P. ramburii, P. yucatan
Sete Lagoas (MG), Gruta Rei do Mato, 3.+4.xi.2011: P. falcifer,
P. ramburii
Teó lo Otoni (MG), Lapa da Vaca Parida, 26.i.2005: Psoc.
lachlani, P. yucatan
Vazante (MG), Gruta da Escarpa, xi.2008: P. falcifer
Vazante (MG), Lapa da Delza, 12.vii.2010: P. ramburii
Vazante (MG), Lapa das Urtigas, 16.ix.2010: P. falcifer
Mato Grosso (MT)
Apiacás (MT), Casa de pedra do Navalha, Parque Nacional do
Juruena, 9.ix.2011: P. angustipennis
Chapada dos Guimarães (MT), Gruta Kiogo Brado, 27.x.2006:
P. clunioventralis, P. spinifer
Paranaíta (MT), Gruta da Pedra Preta, 19.ix.2011: P. didymus
Piauí (PI)
Coronel José Dias (PI), Coroa de Frade, ix.2008: P. thaidis
Coronel José Dias (PI), Toca do Garrincho, 11.ix.2008: Psoc.
pilipennis
Coronel José Dias (PI), Toca do Inferno, 12.ix.2008: P.
punctulatus, P. spinifer
Coronel José Dias (PI), Toca das Moendas, 10.ix.2008: P.
spinifer
Rio Grande do Norte (RN)
Baraúna (RN), Caverna Britador, 11.vi.2010, leg. D. M. Bento:
P. clunjunctus
Baraúna (RN), Caverna Cipós, 11.vi.2010, leg. D. M. Bento:
P. clunjunctus
Baraúna (RN), Caverna Escada, 27.i.2010, leg. D. M. Bento:
P. clunjunctus
Baraúna (RN), Caverna Esquecida, 17.vi.2010, leg. D. M.
Bento: P. clunjunctus
Felipe Guerra (RN), Caverna Arapuá, 3.viii.2010, leg. D. M.
Bento: P. subtilis
Felipe Guerra (RN), Caverna Beira-Rio, 19.iii.2010, leg. D. M.
Bento: P. subtilis
Felipe Guerra (RN), Caverna Rumana, 19.i.2010, 5.viii.2010,
leg. D. M. Bento: P. clunjunctus, P. subtilis
Felipe Guerra (RN), Caverna Trapiá, 6.i.2010, 4.viii.2010, leg.
D. M. Bento: P. yucatan
Felipe Guerra (RN), Gruta Carrapateira, 24.iv.2007: P.
clunjunctus
Felipe Guerra (RN), Gruta da Catedral, 14.ix.2008: P.
clunjunctus
Felipe Guerra (RN), Gruta Roncador, 1.vi.2006: Psoc.
pilipennis
Felipe Guerra (RN), Lapa do Engano, 5.viii.2010, leg. D. M.
Bento: P. clunjunctus
Governador Dix-Sept Rosado (RN), Caverna Capoeira do João
Carlos, 3.vi.2010, leg. D. M. Bento: P. subtilis
Governador Dix-Sept Rosado (RN), Gruta do Lagedo Grande,
21.vii.2010, leg. D. M. Bento: P. clunjunctus, P. spinifer.
P. subtilis
Governador Dix-Sept Rosado (RN), Gruta do Marimbondo
Caboclo, 20.vii.2010, leg. D. M. Bento: P. subtilis
Jandaíra (RN), Gruta Aroeira, 11.i.2006: Psoc. pilipennis
Mossoró (RN), Caverna Trinta, 10.vi.2010, leg. D. M. Bento:
P. clunjunctus
Rio Grande do Sul (RS)
Torres (RS), Furna da Lagoa de Itapeva, 20.v.2008: P. ramburii
São Paulo (SP)
Altinópolis (SP), Gruta Edgar 1, 28.ii.2006: P. spinifer
Altinópolis (SP), Gruta do Itambé, 27.ii.2006: Psoc. lachlani,
P. ramburii
Altinópolis (SP), Gruta Olho de Cabra, 2.iii.2006: P. ramburii,
P. spinifer
Altinópolis (SP), Gruta do Paraná, 1.iii.2006: P. ramburii, P.
spinifer
Ilha Bela (SP), Gruta da Serraria, 22.iv.2006: P. yucatan
Itirapina (SP), Gruta da Toca, 22.x.2004: P. ramburii
Tocantins (TO)
Arraias (TO), cave Ponto 008, 4.-6.x.2010, leg F. Pellegatti et
al.: Neotrogla aurora
Arraias (TO), cave Ponto 014, 4.-6.x.2010, leg F. Pellegatti et
al.: Neotrogla aurora
Aurora do Tocantins (TO), Gruta Asa Branca 1, 7.i.2009, leg.
R. A. Zampaulo: Neotrogla aurora
Aurora do Tocantins (TO), Gruta Biritite, 5.i.2009, leg. R. A.
Zampaulo: Neotrogla aurora
Aurora do Tocantins (TO), Gruta Couve-Flor, 7.i.2009, leg. R.
A. Zampaulo: Neotrogla aurora
Aurora do Tocantins (TO), Gruta das Rãs, 8.i.2009, leg. R. A.
Zampaulo: P. clunjunctus
Dianópolis (TO), Gruta Imperial, iii.2013: Neotrogla aurora
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Psocoptera captured in emergence traps, light traps and a Malaise trap at the Reserva Ducke in 1977-78 are enumerated and discussed. Eighty four species were trapped, mainly in small numbers, and the few common species were present throughout the trapping period. Fewer species were caught in emergence traps than in other trap types. Distribution of Brazilian psocids is briefly discussed.
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A new genus of Sensitibillini from Brazilian caves (Psocodea: 'Psocoptera': Prionoglarididae). - The genus Neotrogla Lienhard gen. n. is described for three new cave-dwelling species from Brazil: Neotrogla brasiliensis Lienhard sp. n. (from Minas Gerais State), N. aurora Lienhard sp. n. (from Tocantins State) and N. truncata Lienhard sp. n. (from Bahia State). These species are the first Neotropical representatives of the sub family Speleketorinae and the first New World representatives of the tribe Sensitibillini, previously known only from southern Africa. This distributional pattern of Sensitibillini is tentatively interpreted as due to Western Gondwanan vicariance. In the females of Neotrogla a complex of accessory structures to the spermathecal duct is described and denoted by the new term "gynosome". A hypothesis of functional complementarity, during copulation, between the "penis-like" gynosome and the strongly reduced male phallosome of Neotrogla is presented.