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 and micropterous; the existence of an apterous form is somewhat doubtful and has yet to be confirmed (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: fig. 37e and Badonnel, 1986: fig. 22); spermapore region with a characteristic pair of more or less sclerotized longitudinal straps (Fig. 2G; see also Mockford, 1993: fig. 62 and Badonnel, 1986: fig. 22); macropterous form occasionally with slightly shortened wings (not as short as in the typical brachypterous form, see Lienhard, 1998: figs 37c, 38c) and rather variable venation (Fig. 3, see also Badonnel, 1986: figs 14-18); in micropterous form length of minute wing rudiments variable, relatively long and bearing some vein rudiments (Lienhard, 1998: fig. 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 confirmed 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 figured 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 justified. The terminalia 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 definitively confirmed 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: 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. – Teófilo 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 macropterous. 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 typical 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 sensillum (Fig. 4D). Epiproct characteristically pigmented (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 articulated seta (not much longer than long seta of setal organ) (Fig. 4BF); female paraproct simple, male paraproct 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 fig. 5 in Badonnel, 1946; Fig. 6B similar to fig. 107 in Mockford, 2011). Sclerotization of ovipositor v1 and v2 variable; sclerotized median axis of v1 weakly developed or absent; median axis of v2 usually well-sclerotized (Fig. 5A), but weakly developed or completely absent in pale individuals (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); spermatophores not sclerotized, somewhat variable in shape, but always similar to that figured in Fig. 5C; the spermatheca of a female may contain up to about a dozen spermatophores (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 artificial 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.

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 significant difference between the African and the Brazilian specimens could be observed. The structure figured by Badonnel (1946: fig. 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.

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. – Teófilo 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 first 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 flap (Fig. 7B). P2 lacking stout sensillum. Pedicel with well-developed microspades organ (Fig. 7E). Paraproctal trichobothria with weakly differentiated basal florets; 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 fine membranes (Fig. 7G); depending on slide-mounting this structure may appear like a free-ending appendix (see Badonnel, 1967: fig. 58; Mockford, 1993: fig. 467; Lienhard, 1998: fig. 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: fig. 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 figured (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 figures given by Townsend (1912) and Brèthes (1923) for P. termitorum and P. astizi respectively fit 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 figures 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 figure 58 is erroneously assigned to P. pilipennis.

Gambrella pilipennis Enderlein, 1931: 221.

Psocathropos pilipennis (Enderlein); Menon, 1942: 35.

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 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 figures 25-26) which we have never observed in P. pilipennis. In our opinion, his figures 25 and 26 illustrate forewings of P. lachlani, while his figures 23, 24 and 27 illustrate wings of P. pilipennis, and his figure 28 the male genitalia of the latter. Badonnel (1967) gives excellent figures of the wings and the male genitalia, but it has to be noted that his figure 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).