Holotype ♂, ‘Gruta Apertar | da Hora | Jandaíra RN <= Rio Grande do Norte>’, ‘ISLA | 21101’, ‘15.ii.2010 | Ferreira, R.L. leg.’, ‘HOLOTYPE (male) | Cylindrogaster cavernicola | sp. n. | Det. Y. Kamimura 2017’.

Cylindrogaster cavernicola sp. n. is a median-sized species with a slender abdomen and simple forceps. This species differs from all other species of Cylindrogaster with the combination of the following characters: the well-developed tegmina; pronotum slightly longer than broad; parameres with blunt apices; and short but weakly sinuated virgae.

Male (holotype: Fig. 1). Length of body (without forceps): 11 mm. Length of forceps: 1.4 mm. Head width: 1.5 mm. Pronotum width: 1.0 mm. Pronotum length: 1.3 mm.

Figures 1-11. 

1-7 Cylindrogaster cavernicola Kamimura, sp. n. (male, holotype): habitus (1), head and thorax (2), tegmina and wings (3), penultimate sternite (4), ultimate tergite and forceps (5), and genitalia (6, 7) 8, 10 Cylindrogaster thoracicus (MM 3637; collected from Itatiaia, Brazil; det. W. D. Hincks): head and thorax (8), and genitalia (10) 9, 11 Cylindrogaster gracilis: head and thorax (9: MM 3677; collected from Itatiaia, Brazil; det. W. D. Hincks), and genitalia (11: MM 3565; collected from Minas Gerais, Brazil; det. W. D. Hincks). Scale bars 3 mm for Fig. 1; 0.5 mm for Figs 2-6, 8-11; 200 ?m for Fig. 7.

The body color of the holotype seems to have been bleached by preservation in ethanol. Body color uniformly pale amber but abdomen darker. Body, including forceps, sparsely pubescent.

Head (Fig. 2) slightly longer than broad, widest in the region of the eyes; frons tumid, occiput depressed; transverse and median suture not conspicuous but visible; hind margin strongly emarginated in middle; post-ocular carina well developed, almost straight, running from middle of the internal margin of the eyes to the hind margin of the head; lateral margins of post-ocular region bordered with strong bristles. Antennae broken, 10 (right) and 15 (left) segments remaining; first segment stout, expanded apically, length almost the same as the distance between antennal bases; second segment short, quadrate; third segment expanding apically, widest width almost the same as the length; fourth segment almost quadrate; fifth segment almost quadrate (left side) to 1.5 times longer than the width (right side); remaining segments gradually lengthening. Eyes prominent, slightly shorter than post-ocular length. Pronotum (Fig. 2) slightly longer than broad; anterior margin almost straight; sides parallel; hind margin broadly rounded; median sulcus distinct; prozona weakly raised. Tegmina (Fig. 3) well developed, 1.5 times wider than pronotum, about twice as long as pronotum; broad triangular scutellum visible. Wings (Fig. 3) well-developed. Legs long, slender; hind tarsi with first segment 2.5 times longer than third, second segment about half as long as third, claw with small arolium. Abdomen long, cylindrical; segments eight and nine slightly expanded. Penultimate (= 9^th) sternite (Fig. 4) slender, with shallow concave sides; caudal margin forming a more or less semicircular lobe. Ultimate (= 10^th) tergite (Fig. 5) moderately inflated, oval; caudal margin shallow concave between forceps. Forceps (Fig. 5) robust and short, almost straight, pubescent especially on inner margins, tapering and weakly curving inward apically, base of inner margin with small tooth. Genitalia (Figs 6, 7); virga short and sinuated; parameres (= external parameres) short and broad, rounded, with small triangular sclerotized tubercle and sparse short hairs at apex.

Female. Unknown.

The subfamily Cylindrogastrinae consists of six species belonging to the Neotropical genus Cylindrogaster (Hincks 1955; Steinmann 1986, 1989b). Their external appearance resembles those of the Diplatyinae species or Diplatyidaesensu stricto (Table 1). However, males of Cylindrogaster spp., including C. cavernicola sp. n. described here, have one gonopore on each of the paired virgae, which is a heavily sclerotized process containing the terminal part of the ejaculatory duct, whereas male diplatyids (and haplodiplatyids) have a pair of bifurcated virgae (Hincks 1955; Steinmann 1989a).

This new species is allied to C. gracilis Stal, 1855, which was recorded from Brazil (and also possibly from Peru). However, the virga is entirely straight and the parameres are much wider than the length in the latter species (Fig. 11). Another Brazilian species, C. thoracicus Dohrn, 1863, can be distinguished from C. cavernicola sp. n. by its much longer virgae (Fig. 10) and pronotum (Fig. 8).

The species epithet refers to the cave-dwelling habit of this new species, although it is presently unknown whether it is a troglobite.

Rio Grande do Norte, Brazil.

The specimen of Cylindrogaster cavernicola Kamimura, sp. n. was collected near the entrance of a cave associated with limestone rocks from the Jandaíra formation (Upper Cretaceous) in northern Rio Grande do Norte state. The caves in this region are predominantly shallow, most of which present several connections with the epigean environment (usually vertical cracks in the limestone outcrops). Accordingly, many caves in the area are strongly influenced by the external environment. Even so, given the extremely dry external environment, the caves represent a more suitable habitat for many animal species, presenting more stable temperatures and higher humidity than the epigean habitat. Furthermore, although the macro-caves are more influenced by the epigean environment, they are connected to huge systems of meso-caves, comprising small passages that are much more stable.

1 ♀, Gruta Túneis, Lagoa Santa, Minas Gerais, 10.x.2011, Ferreira, RL leg. (ISLA 43365).

The female specimen was collected near the entrance of a cave of the Sumidouro state park, located in Lagoa Santa, Minas Gerais state. Numerous nymphs presumably belonging to this species were observed throughout the years, especially on the cave walls in areas close to entrances, in the limestone caves of this region. Therefore, it is possible that this species uses the caves as a protected habitat during its development, but leaves the caves when reaching adulthood. They probably prey upon small invertebrates that are found on the walls near the entrances.

In this study, two adult females of Cylindrogaster were examined. Based on differences in body size and genital structures (see below), these two females are not conspecific. The characteristics for species diagnosis have not been established for female Cylindrogaster spp. In addition, because the collection localities of both female samples (from Minas Gerais and Pará states, respectively) are quite far from the type locality of C. cavernicola sp. n. (Rio Grande do Norte state), these female samples are tentatively treated as Cylindrogaster sp. 1 (Figs 12–17) and Cylindrogaster sp. 2 (Figs 18–21) in this study.

Figures 12–22. 

12–17 Cylindrogaster sp. 1 (female): habitus (12), spiny ridges (indicated by the blue arrowheads) on the mesothorax (13), wing base of the left hindwing (14), and genital regions (15–17) 18–21 Cylindrogaster sp. 2 (female): habitus (18), spiny ridges (indicated by the blue arrowheads) on the mesothorax (19), fustis of the left hindwing (20), and genital regions (21) 22 Cylindrogaster sp. (nymph): detail of cercus. Scale bars 3 mm for Figs 12, 18; 0.5 mm for Figs 13–15, 19–22; 200 μm for Figs 16, 17.

Wing and female genital structures have not been reported for any members of Cylindrogaster. The female genital region of Cylindrogaster sp.1 is characterized by having shorter gonapophysis VIII (gp8) than gonapophysis IX (gp9), which was slightly shorter than the finger-like gonoplac IX (gl9) (Figs 15–17).

The spermatheca of Cylindrogaster sp. 1 consisted of long thin tubes (Fig. 17) without sclerotized or pigmented spermathecal capsules. In the infraorder Protodermaptera (= Pygidicranidaesensu lato or sometimes referred to as basal dermapterans), the number of spermatheca and internal branches vary both inter- and intraspecifically (Popham 1965a; Klass 2003; Kamimura 2004). However, poor specimen quality prevented characterization of the Cylindrogaster sp. 1 spermatheca.

According to Hass and Kukalová-Peck (2001), Diplatyidae (Diplatys) and Haplodiplatyidae (Haplodiplatys) are separated from other members of the infraorder Protodermaptera by the presence of (1) a narrow, slender, and elongated fustis head, (2) a long and slender costal area, (3) a concave and strongly three-dimensional anojugal arm (FAJ: anojugal fulcalare), (4) the distal end of CuA3+4 (cubitus anterior 3 and 4) lying between the 8th and 9th branches of AP1+2 (anal posterior 1 and 2), and (5) two proximal branches of AP1+2 diverging close together. The characteristics unique to Diplatyidae and Haplodiplatyidae (1–3) were observed in Cylindrogaster sp. 1 (Fig. 14). The latter two traits were not observed as the poor quality of the specimen prevented opening of the wings.

The genus Haplodiplatys (Haplodiplatyidae) is characterized by multiple plesiomorphic features, including laterally symmetrical tegmina and absence of a spiny ridge (a component of the tegmina-locking device) on the dorsal side of the mesothorax (Hass and Kukalová-Peck 2001). The female sample of Cylindrogaster sp. 1, however, possessed well-developed spiny ridges (Fig. 13), similar to those in Diplatys spp. (Diplatyidae).

1 ♀, Cave N5SM1-017, Parauapebas, Pará, 17.ii.2011, CARSTE leg. (ISLA 15558).

The specimen was found in the cave N5SM1-017 (also known as GEM-1190 cave). This cave consists of a shallow iron ore cave (14 m horizontal projection), located in Carajás region (Pará state, Brazil). The surrounding vegetation is composed of Amazon forest with a dense canopy. The cave has only one wide and shaded entrance, with the presence of vegetation (lichens, moss, and ferns). Furthermore, it does not present an aphotic zone, being lightened throughout its whole extension. The litter is abundant near the entrance zone and sparse in the rest of the cavity. Several caves were sampled in this region, and as a single specimen was found, it is likely that this species is accidental or seeks only temporarily shelter in caves.

The genital structures of this species are essentially similar to those of Cylindrogaster sp. 1, but the spermatheca was missing likely due to incorrect dissection. Cylindrogaster sp. 2 differs from Cylindrogaster sp. 1 in having triangular shaped gl9 (Fig. 21) vs. finger-like in Cylindrogaster sp. 1 (Figs 15, 16), as well as a much smaller body size (Fig. 18 vs. 12). The wing and mesothoracic structures of Cylindrogaster sp. 2 were basically the same as those of Cylindrogaster sp. 1. Refer to Figures 19 and 20 for images of the spiny ridges and fustis, respectively.

1 nymph, Cavidade RF 86, Barão de Cocais, Minas Gerais, 9.ii.2015, ATIVO AMBIENTAL leg. (ISLA 15507) – 1 nymph, Cavidade CBT_09, Barão de Cocais, Minas Gerais, 19.i.2015, ATIVO AMBIENTAL leg. (ISLA 15508) – 1 nymph, S11D, S11D-0003, Serra / Sul, Canaã dos Carajás, Pará, 16.xii.2014, CARSTE leg. (ISLA 17201) – 1 nymph, Gruta da Lapinha, Lagoa Santa, Minas Gerais, 7.vii.2011, Ferreira, RL leg. (ISLA 21083) – 3 nymphs, Cave GEM-1194, Parauapebas, Pará, 23.ii.2011, CARSTE leg. (ISLA 21087) – 1 nymph, CAPA 03, Itabirito, Minas Gerais, 11.xi.2013, SPELAYON leg. (ISLA 21088) – 1 nymph, Cave Mll GEM-1705, Parauapebas, Pará, 13.iv.2011, CARSTE leg. (ISLA 21091) – 2 nymphs, Cave Mll GEM-1738, Parauapebas, Pará, 3.xi.2011, CARSTE leg. (ISLA 21092) – 1 nymph, Cave Mll GEM-1694, Parauapebas, Pará, 20.iii.2011, CARSTE leg. (ISLA 21095) – 4 nymphs, Didi Vieira cave, Afonso Cláudio, Espírito Santo, 23.iii.2005, Souza MS et al. leg. (ISLA 21098) – 1 nymph, Sitio Paraíso cave, Ecoporanga, Espírito Santo, 22.vii.2004, Souza MS et al. leg. (ISLA 21099) – 2 nymphs, Gruta do Roxo, Novo Oriente de Minas, Minas Gerais, 20.vii.2002, Souza MS et al. leg. (ISLA 21100) – 1 nymph, SEP-0407 (Geraldo Gusso), Felipe Guerra, Rio Grande do Norte, 2.x.2010, Ferreira, RL leg. (ISLA 21102) – 1 nymph, Lapa de Urtiga, Vazante, Minas Gerais, 16.ix.2010, Ferreira, RL leg. (ISLA 2735).

Remarks. A total of 21 nymphs of Cylindrogaster were examined. Nymphal cerci were frequently lost in these specimens, but when present, they were always segmented instead of unsegmented forceps of adults (Fig. 22).

Species diagnosis has not been established for nymphal samples of Cylindrogaster spp.

3 ♂♂, 6 ♀♀, 7 nymphs, Gruta dos Farias cave, Barbalhas, Ceará, 30.iv.2007, Ferreira, RL leg. (ISLA 15565) – 1 nymph, Cave GEM-1623, Parauapebas, Pará, 16.iii.2011, CARSTE leg. (ISLA 21085) – 1 nymph, Gruta Ecos cave, Cocalzinho de Goiás, Goiás, 4.iv.2006, CECAV leg. (ISLA 21096).

While most earwigs found in Brazilian caves seem to be accidental, this species was present as a large population within Gruta dos Farias cave, a sandstone cave located in Barbalhas municipality (Ceará state, Brazil). Many adults and nymphs were observed only in guano piles in deeper areas of the cave, which has a stream trespassing its entire conduit, strongly suggesting that the population is troglophilic. They are likely feeding on bat guano or preying upon small invertebrates.

All adult specimens examined in this study had fully developed tegmina, but lacked hind wings. Four species of Euborellia from the Neotropical region, E. boliviana Brindle, 1971, E. ambigua (Borelli, 1906), E. caraibea Hebard, 1921, and E. janeirensis, also have such characteristics. Among these species, E. janeirensis is distinguished from the others by the presence of well-developed lateral longitudinal ridges on the male abdominal tergites VI and IX, and one or more white/yellow distal antennal segments (Steinmann 1989a). The external morphologies and male genitalia of the specimens examined in this study agreed well with those described previously for E. janeirensis, including brown markings on the femora (Figs 23 and 27), the shapes of the forceps (Figs 23 and 27), and the shapes of the parameres and denticulated pads in the penis lobe (Fig. 26).

Figures 23–35. 

23–26 Euborellia janeirensis (male): habitus (23), penultimate sternite + manubrium (24), and genitalia (25, 26) 27–28 Euborellia janeirensis (female): habitus (27), and spermatheca (28) 29–32 Euborellia brasiliensis (male): habitus (29), penultimate sternite + manubrium (30), and genitalia (31, 32) 33Anisolabididae gen. sp. 1 (female): habitus 34Anisolabididae gen. sp. 2 (nymph): habitus 35Anisolabididae gen. sp. 3 (nymph) habitus. Scale bars 3 mm for Figs 23–25, 27, 29, 30, 33–35; 0.5 mm for Figs 26, 28, 31; 100 μm for Fig. 32.

Taxonomists generally examine only the terminal region of the male genital organs. The male genitalia of E. janeirensis were approximately 22 mm in length, and more than twice the body length with forceps (Fig. 25). The manubrium, which is an extension in the basal, inner margin of the penultimate sternite (Burr 1915b; Ramamurthi 1958), was approximately 6 mm in length (Fig. 24). In male earwigs, the retractor muscles of the genitalia originate from this structure (Popham 1965a). As reported by Mariani (1994), the spermatheca of this species was a long and thin blind duct lacking a capsule at the distal end (Fig. 28).

Males of Euborellia spp. directly insert the elongated virga into the female spermatheca during copulation (Kamimura 2000; Lieshout and Elgar 2011). Elongation of the virga (and the genitalia as a whole, which functions as the virgal sheath when in repose) is considered an adaptation for removal of rival sperm from the female spermatheca, which is usually longer than the virga (Kamimura 2000, 2005, 2013, 2015; Lieshout and Elgar 2011). Thus, genital elongation in E. janeirensis suggests intensive sperm competition in this species.

Brazil, Argentina, Paraguay, and Venezuela.

2 ♂♂, 2 nymphs, Gruta dos Coelhos cave, Lima Duarte, Minas Gerais, 11.vii.2005, Ferreira, RL leg. (ISLA 15564).

This species was found in deeper areas (aphotic zones) of Gruta dos Coelhos and Gruta do Pião caves, both associated with quartzite rocks and located in the Ibitipoca state park (Lima Duarte municipality, Minas Gerais state, Brazil). In the latter cave, several individuals were found walking near root masses, probably searching for prey. There are several caves in the area, and at least three distinct inventories of cave fauna were performed over the last 10 years. However, this species was only found in two caves, suggesting that, although probably not accidental, their association with subterranean habitats is uncommon.

This apterous anisolabidid species from Brazil (Fig. 29) was originally described as Heterolabis brasiliensis by Borelli (1912), as the type species of the monotypic genus Heterolabis. Popham and Brindle (1966) recognized the type species as a member of the genus Euborellia Burr, 1910 and proposed the combination Euborellia brasiliensis (Borelli, 1912) Popham & Brindle, 1966. Therefore, they considered the generic name Heterolabis as a synonym of Euborellia. Their decision was followed by Reichardt (1968b), Sakai (1970), and Steinmann (1977, 1978).

Srivastava (1978) described a second species belonging to Heterolabis, i.e. Heterolabis punctata Srivastava, 1978. Its transfer to the genus Euborellia was proposed by Sakai (1982, 1987) and Srivastava (1999). Since the binomen Euborellia punctata was preoccupied by Euborellia punctata Borelli, 1927, they proposed the replacement names Euborellia srivastavai Sakai, 1987 and E. mindanoensis Srivastava, 1999, respectively. An alternative generic position for H. punctata was proposed by Steinmann (1989b); who recognized it as a member of the genus Epilabis Burr, 1915. Since the relevant binomen was preoccupied by Epilabis punctata Srivastava, 1976, he proposed the replacement name Epilabis harlequin Steinmann, 1989.

Currently Heterolabis Borelli, 1912 is considered a synonym of Euborellia, and the species previously considered as members of Heterolabis is as follows:

1. Euborellia brasiliensis (Borelli, 1912): Popham and Brindle (1966)

Synonym: Heterolabis brasiliensis Borelli, 1912

2. Euborellia srivastavai Sakai, 1987

Synonyms: Heterolabis punctata Srivastava, 1978; Epilabis harlequin Steinmann, 1989 (junior objective synonym); Euborellia mindanoensis Srivastava, 1999 (junior objective synonym).

The generic name Heterolabis Borelli, 1912 is an invalid homonym of Heterolabis Kriechbaumer, 1889 (Ichneumonidae).

Brazil.

2 ♂♂, 9 ♀♀, 8 nymphs, Cave N5SM2-099 (= GEM-1799 cave), Parauapebas, Pará, 5.v.2011, CARSTE leg. (ISLA 15563) – 1 ♀, Cave N5SM2-019 (= GEM-1739 cave), Parauapebas, Pará, 31.x.2010, CARSTE leg. (ISLA 15562) – 3♀, Cave N5SM2-099 (= GEM-1799 cave), Parauapebas, Pará, 31.x.2010, CARSTE leg. (ISLA 15560) – 1 ♂, 2 ♀♀, 8 nymphs, Cave N5SM2-019 (= GEM-1739 cave), Parauapebas, Pará, 5.v.2011, CARSTE leg. (ISLA 15561) – 1 nymph, Cave N5SM2-019 (= GEM-1739 cave), Parauapebas, Pará, 31.x.2010, CARSTE leg. (ISLA 21084) – 1 nymph, Cave N5SM2-019 (= GEM-1739 cave), Parauapebas, Pará, 5.v.2011, CARSTE leg. (ISLA 21089) – 1 nymph, Cave N5SM2-019 (= GEM-1739 cave), Parauapebas, Pará, 5.v.2011, CARSTE leg. (ISLA 21090) – 3 nymphs, Cave N5SM2-019 (= GEM-1739 cave), Parauapebas, Pará, 31.x.2010, CARSTE leg. (ISLA 21094).

Many specimens of P. dorsalis were found in two iron ore caves (N5SM2-019 cave – synonym of GEM-1739 cave, and N5SM2-099 cave – synonym of GEM-1799 cave), both located in Carajás region (Pará state). These caves occur in an iron ore plateau surrounded by the Amazon forest. However, they are in an area of metallophilic savannah, a vegetation type usually found at the top of plateaus. The caves are considerably large (> 100 m in horizontal projection) compared to other caves in the area. Although many caves were sampled in the plateau (at least 100 caves), this species was found in only these two caves, which contain huge colonies of the insectivorous bat genus Pteronotus (Pteronotus gymnonotus Wagner, 1843, in the N5SM2-019 cave and Pteronotus parnellii (Gray, 1843) in the N5SM2-099 cave). These colonies produce large guano piles, where several individuals of P. dorsalis were observed. The populations of P. dorsalis, which include both adults and nymphs, were observed in both dry and rainy seasons in the two caves, strongly suggesting that they are troglophilic.

The external morphologies and genital structures of male and female specimens were examined (Figs 36–41) and matched those of Paralabellula dorsalis (Burmeister 1838; Brunner 1906; Brindle 1971a,b,c; Steinmann 1989a; Briceño 1997; Eberhard et al. 1998). Winged and wingless morphs have been reported for a Costa Rican population of this species, where wingless individuals always predominated in the wild (Briceño and Eberhard 1987). Significantly more winged adults emerged when nymphs were subjected to low nutritional conditions (Briceño and Eberhard 1987). All adults (3 males and 13 females) examined in this study were winged morphs with fully developed tegmina and hind wings (Figs 36, 39). These findings may have been due to the poor nutritional conditions in caves. Although previous authors noted that each branch of the male forceps has a small inner tooth at the base (see Brindle 1971a, b, c; Sakai 1993), all males (n = 3) examined in this study lacked this tooth (Fig. 37). However, one male specimen from Moniquira, Colombia, determined by A. Brindle, also lacked a tooth at the base of the forceps (Fig. 42).

Figures 36–45. 

36–38 Paralabellula dorsalis (male): habitus (36), forceps (dorsolateral view) (37), and genitalia (38) 39–41 Paralabellula dorsalis (female): habitus (39), spermatheca and spiny area in the genitalia (40, 41). Repaired wound patches are indicated by the red arrowheads in Figs 40 and 4142 Paralabellula dorsalis (male; a specimen from Colombia, det. A. Brindle; OMHH S. Sakai Collection, 31.iii.2002 [01-25]): abdomen and forceps (dorsolateral view) 43–44 Doru luteipes (female): habitus (43), and spermatheca (44) 45 Doru lineare (female; coll. Y. Kamimura, at Varzelândia, Minas Gerais, Brazil, outside of caves on 13.iii.2016): spermatheca. Scale bars 3 mm for Figs 36, 39, 42, 43; 0.5 mm for 37; 200 μm for Figs 38, 40, 41, 44, 45.

Based on samples collected from Costa Rica, Briceño (1997) reported the detailed genital structures of this species, which included a horn-like structure and several heavily sclerotized toothed plates in the penis lobe. These structures were also found in the three male samples examined in this study (ho and tp, respectively, in Fig. 38). The spermatheca of this species is a long, thin, blind duct lacking a capsule at the distal end (Mariani 1994; Briceño 1997). Briceño (1997) also reported the presence of multiple fine spines around the spermathecal opening. All such features were present in the specimens examined in this study (Fig. 40, 41).

Using specimens fixed during copulation, Briceño (1997) also examined coupling of the male and female genitalia. Toothed plates were exposed by eversion of the inflated penis lobe, and contacted the inner walls of the vagina, including the spiny area. The horn-like structure functioned as a guiding sheath for the virga, potentially to facilitate insertion into the female spermatheca. There is accumulating evidence that many male animals inflict wounds on the female during mating through use of their genital structures (Lange et al. 2013; Tatarnic et al. 2014; Reinhardt et al. 2014). However, this mode of mating, termed traumatic mating, has been reported in only two species of earwigs, Echinosoma denticulatum Hincks, 1959 (Pygidicranidae: Echinosomatinae; Kamimura and Lee 2014a) and Marava arachidis (Yersin, 1860) (Spongiphoridae: Spongiphorinae; Kamimura et al. 2016b). While Briceño (1997) failed to mention the occurrence of copulatory wounding in P. dorsalis, melanized patches on the membranous region at the spermathecal opening were observed in this study (n = 2; Figs 40, 41). This finding suggests that the male genitalia cause wounding during copulation.

West Indies, Mexico, Costa Rica, Panama and northern South America (including Brazil).

1 ♀, Gruta do Vento cave, Pains, Minas Gerais, 12.x.2000, R. L. Ferreira leg. (ISLA 495) – 1 ♀, Gruta Zé Geraldão cave, Pains, Minas Gerais, 10.iii.2009, R. A. Zampaulo leg. (ISLA 534).

Doru luteipes is an extremely common species in Brazil (Reis et al. 1988), being frequently associated with crops, especially corn. The few observed specimens were found from the Pains region, which is considered the speleological area with the highest concentration of caves in Brazil (and probably in South America). Although more than 300 caves have been examined in this area (Zampaulo 2010), this species was found near the entrances of only two caves, suggesting that they are definitely accidental.

Only female samples of Doru, which are difficult to identify to the species level, were collected in this study. The external morphologies (Fig. 43) matched those of D. luteipes (Scudder 1876; Brindle 1971d; Steinmann 1979; 1993). The females of D. luteipes are similar in appearance to Doru lineare (Eschscholtz, 1822) [Brindle 1971d; Steinmann 1979, 1993: see also Sakai 1993, 1995 for proposal of a synonymy of this species with Doru taeniatum (Dohrn, 1862)]; however, the spermathecal morphologies, including the shape of the spermathecal capsule with seven constrictions (Fig. 44: the spermathecal morphology was examined for the sample from Gruta do Vento cave), were the same as those described by Mariani (1994) for D. luteipes. According to Mariani (1994), the spermathecal capsule of D. lineare is shorter with fewer and weaker constrictions. This was confirmed in a female sample collected at Varzelândia, Minas Gerais, Brazil on 13.iii.2016 by Y.K., with conspecific males (Fig. 45). However, parthenogenesis has been reported for some Brazilian populations of D. lineare (Cocco et al. 2013), suggesting possible polymorphisms in the spermathecal morphology. Thus, the identification of the samples is tentative.

Colombia, Surinam, Brazil, Peru, Bolivia, and Argentina.