Research Article |
Corresponding author: Antonio D. Brescovit ( antonio.brescovit@butantan.gov.br ) Academic editor: Cristina Rheims
© 2016 Antonio D. Brescovit, Alexander Sánchez-Ruiz.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Brescovit AD, Sánchez-Ruiz A (2016) Descriptions of two new genera of the spider family Caponiidae (Arachnida, Araneae) and an update of Tisentnops and Taintnops from Brazil and Chile. ZooKeys 622: 47-84. https://doi.org/10.3897/zookeys.622.8682
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New members of the spider family Caponiidae from Brazil and Chile are presented. Three new species in previously known genera are described: Taintnops paposo sp. n. from Chile, and the Brazilian Tisentnops mineiro sp. n. and Tisentnops onix sp. n., both belonging to a genus known only from its damaged type. Additionally, two new non–nopine Brazilian genera are proposed: Nasutonops gen. n. including three new species: N. chapeu sp. n., N. sincora sp. n. and N. xaxado sp. n.; and Carajas gen. n., known only from the type species C. paraua sp. n. Both new genera have entire, rather than sub-segmented tarsi. Therefore, they are not included in the caponiid subfamily Nopinae. Nasutonops gen. n. is characterized by the presence of a projected clypeal horn, unique among caponiids. Additionally, the first blind caponiids are described: Tisentnops mineiro sp. n. from the state of Minas Gerais and Carajas paraua sp. n. from the state of Pará. Both of these species are found only in caves and completely lack eyes.
Taxonomy, new species, haplogynae, new genus, Neotropical region
The family Caponiidae is currently represented by 15 genera and 98 species (
Currently nine non-nopine genera are known: Caponia Simon, Diploglena Purcell, Iraponia Kranz-Baltensperger, Platnick & Dupérré and Laoponia Platnick & Jäger from Africa and Asia, and Calponia Platnick, Caponina Simon, Notnops Platnick, Taintnops Platnick and Tisentnops Platnick from the New World. Recently, the first genera from Asia were described (
During several field trips to collect haplogynae spiders in Brazil and Chile, three new species of the genera Taintnops and Tisentnops were found (the Chilean Taintnops paposo sp. n. and the Brazilian Tisentnops mineiro sp. n. and Tisentnops onix sp. n.). The new species allowed us to gather detailed morphological information, expand the distributional range, and provide information about the natural history of these two genera. Additionally, we found two new non–nopine Brazilian genera: Nasutonops gen. n. and Carajas gen. n.; the first with three new species N. chapeu sp. n., N. sincora sp. n., N. xaxado sp. n., and the second known from only one species C. paraua sp. n. All these new genera and species are also described in this paper.
Some of the new species are remarkable among caponiids. We describe the first known blind caponiids: T. mineiro sp. n. and C. paraua sp. n.; both species are known only from caves and completely lack eyes. Another unique characteristic occurs in the new genus Nasutonops, which has a hard, distally projected clypeal horn, similar to the goblin spider genus Unicorn Platnick & Brescovit (see
Morphological observations and illustrations were made using a Leica MZ12 stereomicroscope with a camera lucida. Photographs were taken with a Leica DFC 500 digital camera mounted on a Leica MZ 16A stereomicroscope. Extended focal range images were composed with Leica Application Suite version 2.5.0. The female internal genitalia were dissected following
The specimens examined are deposited in the following collections (Abbreviation and curator in parentheses): American Museum of Natural History, New York (
Tisentnops Platnick, 1994b: 9 (Type species by original designation Caponina leopoldi Zapfe).
Tisentnops mineiro sp. n., male holotype (A–G, J), female paratype,
Members of the genus can be easily separated from all other caponiid genera by the distally widened palpal endites, as in Diploglena, but uniquely modified with a series of setae with elongated sockets in the sub-marginal and anterior margin(Fig.
SEM images of Tisentnops mineiro sp. n., female (
Described by
SEM images of Tisentnops mineiro sp. n., female (
Tisentnops mineiro sp. n., male from Moeda, Minas Gerais (A–C), female from Santa Bárbara, Minas Gerais (
Tisentnops onix sp. n., female holotype (A–I). A habitus, dorsal view B same, ventral view C same, lateral view D carapace, dorsal view E ocular area, dorsal view F abdomen, genital area, ventral view G mouthparts, ventral view H leg I, prolateral view, showing long hairs I spinnerets, ventral view.
Known from Chile and southeastern Brazil.
Male holotype from Gruta da Serra da Moeda (20°19'58"S, 44°03'10"W), Moeda, Minas Gerais, Brazil, X.2005, R.L. Ferreira, deposited in
The specific name is the designation for the native people from the state of Minas Gerais, Brazil, where this species was found.
Tisentnops mineiro sp. n. is easily separated from other species of the genus by the absence of eyes (Figs
Male (holotype). Total length 3.6. Carapace 1.6 long, 1.2 wide. Coloration: cephalothorax orange reddish. Palps yellow. Legs yellow, except coxae I-II orange-reddish. Abdomen uniformly yellow-white. Eyes absent of setae with elongated bases forming a row on the anterior margin of the endites, infrequent on the sub-marginal part (Fig.
Female (paratype). Total length 3.8. Carapace 1.7 long, 1.2 wide. Coloration: cephalothorax and legs orange, except legs III-IV yellow and external border of endites brown. Abdomen grayish. Palpal endites as in male, but with fewer sub-marginal modified sochets (Fig.
BRAZIL, Minas Gerais: Conceição do Mato Dentro, Cave CSS-05 (18°55'02"S, 43°25'41"W), 1♀, 12–26/VIII/2013, L. Madeira (
Known only from the state of Minas Gerais, southeastern Brazil (Fig.
All 79 specimens (7♂, 24♀, 48 immature) of Tisentnops mineiro sp. n. were collected in 33 caves distributed in rock outcrops in rupestrian fields found in mountain peaks of the Atlantic Forest and lowland areas of regenerated forest or grasslands. Unlike T. onix sp. n. from limestone caves, T. mineiro sp. n. was only found in iron caves. Additionally, T. mineiro sp. n. was found in palaeoburrows “Paleotoca” (natural shelter of extinct mammals, e.g. Giant Armadillos, see
Male holotype from Maciço da Gruta da Taboa, Cave 64 (19°28'35"S, 44°55'34"W), Sete Lagoas, Minas Gerais, Brazil, 26.II.2015, F. Bondezan col., deposited in
The specific name refers to the shape of the black spot on the ocular area which resembles a lozenge-shaped onyx jewel.
Tisentnops onyx sp. n. differs from T. mineiro sp. n. by the presence of two eyes (Fig.
Male (holotype). Total length 3.7. Carapace 1.8 long, 1.4 wide. Coloration as in T. mineiro sp. n., except cephalothorax red-brown, with black spot in the ocular area, and legs olive-brown. Leg measurements: I: femur 1.2/ patella 0.6/ tibia 1.0/ metatarsus 0.5/ tarsus 0.5/ total 3.8; II: 1.05/ 0.55/ 0.6/ 0.45/ 0.45/ 3.1; III: 0.9/ 0.4/ 0.6/ 0.5/ 0.5/ 2.9; IV: 1.2/ 0.6/ 1.2/ 0.6/ 0.7/ 4.3. Legs with long trichobothria. Elongate raised sockets as in T. mineiro sp. n. Palpal cymbium twice the length of the bulb, bulb enlarged distally, next to the base of the embolus. Embolus curved, half the length of bulb and narrow medially (Fig.
Female (paratype). Total length 4.7. Carapace 1.9 long, 1.4 wide. Coloration as in male, except sternum, endites and chelicerae red-brown (Fig.
None.
Known only from the type locality in the state of Minas Gerais, southeastern Brazil (Fig.
Tisentnops onix sp. n. is a troglobite spider. The female was collected from a cave, 15–20 meters from the entrance, under a rock in the aphotic area. The soil is very humid, composed of large and small rocks (Fig.
Taintnops Platnick, 1994b: 9 (type species by original designation Taintnops goloboffi Platnick).
Members of Taintnops can be separated from other caponiids as follows: from Nopinae genera by having entire, rather than subsegmented tarsi, from Caponia, Calponia, Carajas gen n., Nasutonops gen. n. and Notnops by having only two eyes (Fig.
Taintnops paposo sp. n. from Taltal, Antofagasta, Chile, female paratype (A–J) A habitus, dorsal view B carapace, ocular area, dorsal view C carapace, ventral view D leg I, metatarsus and tarsus distal, retrolateral view E pedipalp, prolateral view F same, retrolateral view G leg I, prolateral view H genital area, ventral view I spinnerets, posterior view J same, ventral view.
SEM images of Taintnops paposo sp. n., female paratype (A–I) A chelicerae, ventral view B left endite and labium, ventral view C left serrula, ventral view D left pedipalp, prolateral view E same, distal area, prolateral view F paired claws on right leg I, prolateral view G trichobotria base on leg II, dorsal view H sensilla on leg II, dorsal view I paired claws on right leg IV, prolateral view.
Described by
Holotype and paratype females from Reserva Nacional Paposo (24°57,82'S, 70°27,961'W), 52m asl, Taltal, Antofagasta Province, Chile, 16/VII/2012, A.D. Brescovit, A.J. Santos & A. Taucare-Rios col., deposited in
The specific name is a noun in apposition taken from the type locality.
Females differ from those of Taintnops goloboffi Platnick (see
Male. Unknown.
Female (holotype). Total length 3.9. Carapace 1.3 long, 0.8 wide. Carapace, mouthparts and sternum reddish. Border of eyes black. Legs and pedipalp orange. Abdomen uniformly creamy white, with epyginal area orange (Fig.
None.
Known only from Taltal area, in the Antofagasta Region.
Both specimens were collected under rocks during the day. Silk retreats were not observed in the area.
Nasutonops xaxado sp. n.
The generic name comes from the Latin words nasutus (with a large nose) combined with nops (less eyes), and is masculine in gender.
Males and females can be easily distinguished from all other known caponiids by the presence of a clypeal horn, projected distally (Fig.
Nasutonops xaxado sp. n., male from Serra Talhada, Pernambuco (A–F) female, same locality (G–L) A habitus, dorsal view B carapace, lateral view C ocular area, lateral view D left pedipalp, prolateral view E same, retrolateral view F same, ventral view G habitus, dorsal view H same, ventral view I carapace, ventral view J ocular area, dorsal view K genital area, ventral view L spinnerets, ventral view.
Moderate-sized caponiids with six eyes (Fig.
SEM images of Nasutonops xaxado sp. n., male from Serra Talhada, Pernambuco (A–L) A carapace, lateral oblique view B ocular area, lateral oblique view C striated area between posterior eyes, detail, dorsal view D clypeal projection, lateral view E same with chelicerae, anterior view F same, anterior view G chelicerae, stridulatory area, lateral view H serrula, lateral view I left pedipalp, retrolateral view J embolus, retrolateral view K tip of embolus, distal area L bulb, ventral view.
SEM images of Nasutonops xaxado sp. n., male (A–D) and female (E–L) from Serra Talhada, Pernambuco. A paired claws on right leg II, prolateral view B unpaired claw on right leg II, prolateral view C trichobothria base on right leg II, dorsal view D tarsal organ on right leg II, dorsal view E ocular area, lateral oblique view F same, anterior view G chelicerae, stridulatory area, lateral view H clypeal projection, lateral view I same, dorsal view J striated area between posterior eyes, detail, dorsal view K stridulatory pick on left pedipalp, prolateral view L distal area of left pedipalp, anterior view.
Nasutonops xaxado sp. n., female from Serra Talhada, Pernambuco (A–B) Nasutonops chapeu sp. n., male and female from Irecê, Bahia (C–G), Nasutonops sincora sp. n., female from Contendas do Sincorá, Bahia (H–I) (A–B, F–G, H–I) female internal genitalia A, F, H dorsal view B, G, I anterior view (C–E) male palp, A prolateral view B ventral view D prolateral view (if = internal transversal fold, psb = pair of sclerotized bars). Scale bars: A–G 0.5 mm.
Nasutonops sincora sp. n., female from Contendas do Sincorá, Bahia (A–G). A habitus, dorsal view B same, ventral view C carapace, ventral view D ocular area, lateral view E same, dorsal view F genital area, ventral view G spinnerets, ventral view. Nasutonops xaxado sp. n. (H–J) HSEM images of internal genitalia, dorsal view I same, posterior view J same, anterior view (if = internal transversal fold; psb = pair of sclerotized bars). Scale bars: A–G 0.5 mm.
Known only from Brazilian Caatinga in the states of Bahia and Pernambuco, Brazil.
Male holotype and female paratype from Serra Talhada (7°59'9"S, 38°17'45"W), Pernambuco, Brazil, 2008-2009, H. Amorin col., deposited in
The specific name is a nickname of the type locality, known as the ‘’Capital do Xaxado’’. The Xaxado is a popular dance in this region.
Females of Nasutonops xaxado sp. n. resemble those of N. chapeu sp. n. in lacking the connection of a pair of sclerotized bars in the female internal genitalia (Fig.
Male (Holotype). Total length 3.80, with clypeal horn. Carapace 1.7 long, 1.2 wide. Coloration: carapace orange-reddish, except apex of endites and labium white, legs and palps orange. Eye median diameters 0.3, interdistances 0.15. Leg measurements: I: femur .1.1/ patella 0.5/ tibia 0.8/ metatarsus 0.7/ tarsus 0.4/ total 3.5; II: 1.1/ 0.5/ 0.7/ 0.7/ 0.4/ 3.4; III: 0.9/ 0.5/ 0.6/ 0.6/ 0.5/ 3.1; IV: 1.1/ 0.6/ 1.0/ 0.9/ 0.6/ 4.2. Abdomen gray with epiandric plate and spinnerets orange. Palpal cymbium not covered with thick layer of distal setae (Fig.
Female (allotype). Total length 4.5, with clypeal horn. Carapace length 1.7 width 1.2. Coloration as in male. Eye median diameters 0.3, interdistances 0.2. Leg measurements: I: femur .1.0/ patella 0.5/ tibia 0.8/ metatarsus 0.6/ tarsus 0.5/ total 3.4; II: 1.0/ 0.5/ 0.7/ 0.7/ 0.5/ 3.4; III: 0.9/ 0.4/ 0.6/ 0.6/ 0.4/ 2.9; IV: 1.0/ 0.5/ 0.8/ 0.9/ 0.5/ 3.7. Internal genitalia with enlarged anterior ends and short posterior ends of the pair of sclerotized bars (Figs
Left metatarsus and tarsus IV of male absent.
BRAZIL: Pernambuco, Serra Talhada (7°59'9"S, 38°17'45"W), 1♂ 1♀, 2008-2009, H. Amorin col. (
Known only from the type locality in the Brazilian Caatinga.
The specimens were collected in soil with pitfall traps.
Male holotype from Parque Estadual Morro do Chapéu (11°29'19,2"S, 41°15'27,6"W), Morro do Chapéu, Bahia, 21.I.2012, 1097 m asl., I.L.F. Magalhaes et al. col, and female allotype from Área da Mineradora Galvani, Irecê (11°18'14"S, 41°51'21"W), Bahia, 27.VI.-27.IX.2011, M.C. Nunes col., deposited in
Although the male and female were not collected from the exact same locality, both specimens were collected near one another in the state of Bahia, and they have the same body coloration.
The specific name is an apposition to the type locality.
Females of Nasutonops chapeu sp. n. resemble those of N. xaxado sp. n. in lacking the connection of the pair of elongate sclerotized bars in the female internal genitalia (Fig.
Male (holotype). Total length 3.2, with clypeal horn. Carapace 1.3 long, 1.05 wide. Coloration of body as in N. xaxado sp. n., except abdomen grayish. Eye median diameters 1.0, inter distances 0.6. Leg measurements: I: femur 1.0/ patella 0.4/ tibia 0.75/ metatarsus 0.55/ tarsus 0.3/ total 3.0; II: 0.9/ 0.4/ 0.65/ 0.55/ 0.3/ 2.8; III: 0.75/ 0.35/ 0.5/ 0.5/ 0.25/ 2.35; IV: 0.95/ 0.4/ 0.8/ 0.7/ 0.45/ 3.3.
Female (allotype). Total length 4.3, with clypeal horn. Carapace 1.55 long, 1.1 wide. Coloration as in male. Eye median diameters 1.0, inter distances 0.8. Leg measurements: I: femur 1.1/ patella 0.45/ tibia 0.75/ metatarsus 0.65/ tarsus 0.4/ total 3.35; II: 1.0/ 0.4/ 0.75/ 0.6/ 0.45/ 3.2; III: 0.8/ 0.35/ 0.6/ 0.6/ 0.35/ 2.7; IV: 1.1/ 0.55/ 0.9/ 0.9/ 0.5/ 3.95. Internal genitalia with long pair of elongate sclerotized bars with narrow anterior ends and slender posterior ends, and with an enlarged area on the anterior third (Fig.
None.
Known only from the type locality in the Brazilian Caatinga from the state of Bahia.
Female holotype from Floresta Nacional Contendas do Sincorá (13°46'–14°00'S, 41°03'–41°10'W), Contendas do Sincorá, Bahia, Brazil, X.2007–X.2008, Y.G. Santos col., deposited in
The specific name is an apposition to the type locality.
The female of Nasutonops sincora sp. n. differs from others females of the genus by the elevated area of the pair of transverse elongate sclerotized bars that are connected anteriorly, forming a rounded, small plate (11H–I) in the female genitalia.
Male. Unknown.
Female (holotype). Total length 4.5, with shortest clypeal horn. Carapace 1.7 long, 1.2 wide. Coloration: cephalothorax and legs orange, ocular area black, abdomen cream, covered with gray hairs, genital area orange and spinnerets yellow (12 A–B). Clypeal horn truncate at tip (Fig.
None.
Known only from the type locality in the Brazilian Caatinga from the state of Bahia.
Carajas paraua sp. n.
The generic name is an apposition to the type locality where all known specimens were collected and is masculine in gender.
Members of Carajas gen. n. can be easily separated from other caponiid genera by having anteriorly and posteriorly strongly projected endites (Figs
Carajas paraua sp. n., male (A–F) and female (G–K) from Parauapebas, Pará. A habitus dorsal view B same, ventral view C left leg I, prolateral view D left leg IV, prolateral view E left pedipalp, prolateral view F same, retrolateral view G left pedipalp prolateral view H same, retrolateral view I habitus, dorsal view J carapace, ventral view K genital area, ventral view.
SEM images of Carajas paraua sp. n., male from type locality. A carapace, ocular area, dorsal view B carapace, ventral view C left chelicerae, prolateral view D mouthparts, ventral view E labium, distal area, ventral view F labium, distal area, prolateral view G paired claws on right leg IV, prolateral view H right tibiae and metatarsus I showing long trichobotria, prolateral view I tricobothria base on right leg I, dorsal view J pedipalp, prolateral view (inset: detail of embolus) K bulb, dorsal view L same,ventral view (inset: detail of embolus).
SEM images of Carajas paraua sp. n., three females (
Moderate-sized caponiids (Fig.
Carajas paraua sp. n., male from
Known only from caves in Carajás region, in the state of Pará, Brazil.
Male holotype and female allotype from Gruta N5S8 (06°06'29"S, 50°07'57"W), Flona de Carajás, Parauapebas, Pará, Brazil, 7-12.X.2008, R. Andrade, deposited in
The specific name is a noun in apposition taken from the Brazilian Tupi Indian language that means parrot (‘’Papagaio’’ in Portuguese) and refers to this common bird in the region of Parauapebas.
With the characters of the genus and genitalia as above described.
Male (holotype). Total length 4.2. Carapace 2.1 long, 1.5 wide. Coloration: cephalothorax uniformly orange-reddish, except border of sternum brown. Legs and palps yellow. Abdomen uniformly grayish. Leg measurements: I: femur 1.8/ patella 1.0/ tibia 1.4/ metatarsus 1.1/ tarsus 1.0/ total 6.3; II: 1.6/ 1.0/ 1.4/ 1.0/ 0.9/ 5.9; III: 1.5/ 0.7/ 1.1/ 1.0/ 0.8/ 4.1; IV: 2.0/ 1.0/ 2.1/ 1.3/ 1.0/ 7.4. Palp as in figures 13D–E, 14J–L and 16A–C.
Female (allotype). Total length 4.3. Carapace 2.1 long, 1.5 wide. Coloration as in male. Leg measurements: I: femur .1.3/ patella 0.7/ tibia 1.1/ metatarsus 0.7/ tarsus 0.6/ total 4.4; II: 1.1/ 0.6/ 0.9/ 0.7/ 0.6/ 3.9; III: 1.0/ 0.6/ 0.8/ 0.7/ 0.5/ 3.6; IV: 1.4/ 0.8/ 1.4/ 0.9/ 0.7/ 5.2. Internal genitalia as described for the genus (Fig.
Tisentnops mineiro sp. n. (A), Tisentnops onix sp. n. (B–D), Carajas paraua sp. n. (E–I). A adult female on the soil B Gruta da Taboa, Minas Gerais, Brazil, showing the rocks on the soil where specimens of T. onix sp. n. were collected C adult female on the soil D same E adult female on rock F adult male on rock G adult female on rock H immature on rock I same.
BRASIL. Pará: Parauapebas, Flona de Carajás, Cave N1-15 (06°02'03"S, 50°16'17"W), Magangá, 2♀ 1imm., 28/IX-30/X/2007, R. Andrade col. (
Known only from caves in the state of Pará, northern Brazil (Fig.
All 767 specimens (34♂, 179♀, 554 immatures) of Carajas paraua sp. n. were collected in 104 caves distributed in rock outcrops covered by canga vegetation (details in
The phylogenetic relationships of Caponiidae are unknown, although some studies have inferred the relationship of certain genera of the family (
Among the genera described here, the most remarkable is Nasutonops gen. n. All species have a distally projected clypeal horn, not found in other Caponiidae. Despite this interesting modification, the ocular area and genital structures link this species to Caponina (see
The first blind caponiids described here are also remarkable (T. mineiro sp. n. and C. paraua sp. n.). They were found only in caves and are totally eyeless. Furthermore, they have other troglobitic adaptations, such as very long trichobothria (Figs
The description of these two new Tisentnops species allowed us to increase the knowledge of this genus. Specifically, the configuration of the female internal genitalia was studied for the first time. The female genitalia of Tisentnops resembles that found in Calponia, Carajas gen. n., Diploglena, Notnops, Laoponia and Iraponia, with an elongate membranous anteromedian receptaculum formed by an anteriorly directed median tubular duct leading to a globose sac (see Figs
In summary, the female internal genitalia of non-nopine genera may indicate a monophyletic group formed by those genera with a median duct and a globose sac on the membranous anteromedian receptaculum. In this case, Diploglena and Tisentnops belong to this group and would be closely related as suggested by
We thank Igor Cizauskas for the help in collecting haplogynae spiders and for the distribution maps and natural history of Brazilian species. Alexandre B. Bonaldo for his assistance using the Scanning Electron Microscope from