A new genus, Reductoonops, is established for a group of New World soft-bodied oonopine spiders, found from Mexico south to Chile, characterized by reduced size, a flattened clypeus, loss of the posterior median spinnerets, and often by loss of the four posterior eyes. Members of this group resemble those of Stenoonops Simon but lack the clump of short setae found on the dorsal surface of the palpal tarsus of both sexes in that genus, and often have four pairs of deep channels at the sides of the sternum, the most anterior pair of which demarcate a short, trapezoidal, anterior portion of the sternum. A total of 34 new species are described from Mexico (chamela, armeria, niltepec, real, nubes, jabin), Costa Rica (monte, lucha), Costa Rica and Panama (naci), Panama (almirante, escopeta, bayano), Jamaica (ferry), Curaçao (hato), Martinique (diamant), Colombia (marta, sasaima, meta, leticia), Ecuador (tandapi, pichincha, tina, domingo, otonga, palenque, napo, jatun, hedlite, molleturo, celica, yasuni), the Galapagos Islands (pinta), Peru (carpish), and Chile (elqui).
INTRODUCTION
Eye reduction is a phenomenon that has seemingly occurred multiple times within the goblin spider subfamily Oonopinae. The most obvious examples are the species that are entirely blind; they are sufficiently uncommon that, when found, they have often been placed in monotypic genera, and their relationships are therefore often enigmatic. The first such species was described as Blanioonops patellaris Simon and Fage (1922), based on females taken in a cave in Tanzania and never restudied. The second blind oonopid to be reported was described as Wanops coecus by Chamberlin and Ivie (1938), based on a male taken in a cave in Yucután, Mexico; those authors considered their species to be related to Oonops Templeton, and the male palp has a terminal embolus that does indeed resemble those found in several normal-eyed Neotropical species currently misplaced in Oonops (e.g., Oonops reticulatus Petrunkevitch and its relatives).
A third species, Dysderoides typhlos Fage (1946), was based on a single female taken in a cave in northern India; Fage also transferred into Dysderoides a six-eyed species from Venezuela originally described as Telchius micans Simon (1893). The Indian species has since been studied in detail by Grismado et al. (2014), who also described several related species from caves in northern India and Thailand; all those species seem to be completely blind, although one of the Indian species shows what appear to be remnants of the eyes, beneath the cuticle. The normal-eyed Venezuelan species transferred to Dysderoides by Fage is not at all closely related to these taxa, and belongs to a new Neotropical genus studied by Bolzern (2014).
In a series of papers, Benoit (1964, 1975a, 1975b, 1976, 1977) described no fewer than 11 blind species from central and southern Africa; one was assigned to Oonops by default, but the others were placed in new genera (Caecoonops, Termitoonops, Anophthalmoonops, Hypnoonops, and Zyngoonops). Interestingly, these species are not troglobitic, but were taken either from leaf litter or (in the case of the first three genera) from termite mounds. However, Benoits description of Zyngoonops clandestinus Benoit (1977) as lacking eyes was erroneous (see Fannes, 2013).
The past two decades have seen the addition of at least 19 additional blind species, including two species of Camptoscaphiella Caporiacco from China and New Caledonia (Deeleman-Reinhold, 1995; Baehr and Harvey, 2013), yet another new genus, Cousinea Saaristo (2001), established for a blind species from the Seychelles that has subsequently been studied in detail by Fannes (2010), and a remarkable fauna, including two cavernicolous species of Opopaea Simon and 14 subterranean species of Prethopalpus Baehr et al. from Western Australia (Harvey and Edward, 2007; Baehr et al., 2012).
Although one of the new species described below as Reductoonops jabin is blind, eye reduction is obviously not an all-or-nothing phenomenon. Most obviously, oonopids with only two eyes, rather than the normal six, have been known since O. P.-Cambridge (1908) described the aptly named Diblemma donisthorpei, based on a male taken in a greenhouse at Kew Gardens in London. It was not until Saaristo (2001) rediscovered this animal on the Seychelles that the likely source of the introduction to Great Britain was revealed. Since 1908, several additional two-eyed oonopids have been described, especially among hard-bodied (“gamasomorphine”) taxa belonging to the Zyngoonops complex, including two species of Zyngoonops (see Benoit, 1977; Fannes, 2013) and four of Coxapopha Platnick (see Platnick, 2000; Ott and Brescovit, 2004). The two-eyed, hard-bodied Argentine species described as Dysderina caeca Birabén (1954) is obviously misplaced in that genus and probably also belongs to Coxapopha or a closely related member of the Zyngoonops complex.
Most of the animals described below as members of the new genus Reductoonops are soft-bodied and have only two eyes; all four of the posterior eyes have been lost entirely (figs. 4, 58). Only one genus including two-eyed, soft-bodied forms has been described: Xyccarph Brignoli (1978), of which three species with eyes of this type are now known, all from Brazil (Höfer and Brescovit, 1996). The members of Xyccarph differ in numerous characters from the taxa described below, however; they have numerous leg spines (which are lacking in the members of Reductoonops), show bifid male endites, lack sternal grooves, and have the male palpal bulb and cymbium fully fused. In Reductoonops, there is usually an obvious seam between the bulb and cymbium, but in three species (R. real from Mexico, plus R. palenque and R. jatun from Ecuador) the seam is obvious only ventrally, and in two other species (R. domingo from Ecuador and R. pinta from the Galapagos Islands), the seam is represented at most by a series of pores.
Other taxa are known in which eye reduction is present, but less obvious, as the posterior median and lateral eyes are reduced in size but still present. Notable among these taxa are two groups from the Neotropics: the Andean genus Niarchos (Platnick and Dupérré, 2010b) and two species from southern Brazil, closely related to each other, that were misplaced in Opopaea Simon by Ott (2003). One of those misplaced species is of special interest as only two of the posterior eyes are still present, making these animals one of the very few known examples of spiders with only four eyes.
The oonopids described below have long been known in collections, where they have commonly been placed as close relatives of Stenoonops Simon, and they do seem to be members of the Stenoonops complex of genera, sharing with other genera in that complex characters such as a lightly sclerotized cephalothorax paired with an unsclerotized abdomen, lateral extensions of the sternum that protrude between the leg coxae (especially coxae II and III), a tarsal organ on which the distalmost receptor is distally bifid, with two lobes protruding from a common base (figs. 25–29, 84–88), a subdistal constriction on the dorsal surface of the leg femora that is marked by a transverse row of setae (figs. 39, 89, 90), and distal setae with fork-shaped tips on the endites (fig. 9).
One reason that these species may have been neglected in prior literature is their diminutive size. Adult males are under 1.2 mm in total length, and adult females are under 1.7 mm; the small size and pale coloration make them exceptionally difficult to study. As in Stenoonops, the larger females often have the abdominal contents significantly smaller than the cuticle, so that (at least in preserved specimens), some parts of the abdomen appear empty (as in fig. 545). In extreme cases, even one-third of the abdominal length can appear empty, and these are probably females that were preserved shortly after laying eggs.
It seems clear that most of the two-eyed species described below form a monophyletic group, as they are united by several unique and presumably synapomorphic characters. Perhaps their most obvious feature is in sternal morphology. It is common for oonopids to have three pairs of depressions extending inward from the sternal margin between legs I and II, legs II and III, and legs III and IV; in some groups, such as Stenoonops, these depressions have become narrow but distinct grooves (Platnick and Dupérré, 2010a: figs. 7, 18. 46, 55). Typical members of Reductoonops have the sternal depressions present as channels, with two steep sides, rather than as simple grooves (figs. 2, 56), and they have an additional, fourth pair of sternal channels extending inward between leg I and the endites. These anterior sternal channels demarcate a distinct, trapezoidal anterior portion of the sternum, making the animals easy to recognize (figs. 44, 270). The sternal modifications are often, but not always, accompanied by the presence of a deep, longitudinal, median incision in the labium of males (figs. 8, 242), which sometimes occurs in females as well, and the taxa lacking that deep incision in one or both sexes may thus represent the most plesiomorphic of the typical species.
The typical species of Reductoonops are found from Mexico south to Chile and are also united by a characteristic male palpal morphology. The palpal bulb is reduced in size, so much so that adult males could easily be mistaken for females if not studied in detail (as happened, for example, in the case of the genus Cousinea; see Saaristo, 2001; Fannes, 2010). The cymbium is dorsally shortened and ventrally elongated, forming a cup-shaped structure that cradles the bulb (figs. 49, 51). The terminal elements are complex, with a sinuous embolus that bears numerous spikes at its tip (fig. 36). The female genitalia are also characteristic, with a long, narrow anterior genitalic process that typically bears dorsal projections, probably of glandular nature (figs. 70, 71).
These small species have a somewhat flattened body, most obviously anteriorly, where the clypeus has become almost flat, rather then vertical (figs. 5, 59). The small size and flattened bodies suggest that these animals probably live deeper in leaf litter than do most other oonopids, and the eye reduction may well be part of a suite of adaptations for living in the smaller interstitial spaces that are available in deep litter, where little light penetrates.
In addition to the species sharing most or all of this suite of characters (reduced body size, flattened clypeus, loss of the posterior eyes, and the extra, anterior pair of sternal channels), there are some species from the northern part of the generic distribution range that lack some of these features. In R. real from Chiapas, Mexico, for example, both sexes have only two eyes, but neither has the extra pair of sternal channels. The genitalia resemble those of the other species; the male palp has a sinuous embolus, but it seems to bear only one spike at the tip. Thus it seems likely that this species may be the sister group of the more typical Reductoonops (i.e., those united by the presence of anterior sternal channels and several spikes on the embolus tip).
There are six additional species, from southern Mexico, Panama, Curaçao, and Colombia, that are even more different from typical Reductoonops members. In addition to lacking the anterior sternal channels, they still retain all four of the posterior eyes. In three species from Mexico, the male palp is much simpler, with only an embolus and a conductor, which originate from a common base (figs. 110–113), and the tip of the embolus bears no spikes; the anterior genitalic process of females is shorter and bipartite (figs. 119,120). Although these three species seem closely related to each other, it isn't likely that all the six-eyed species together form a monophyletic group. The other three species are each known from only one sex. The male of the species from Colombia has a palp that resembles the typical species in having three terminal elements (a palpal apophysis as well as the embolus and conductor), even though the embolus seems to bear no terminal spikes. Similarly, the female of the species from Curaçao has genitalia that seem closer to those of the typical species than to those of the three Mexican species or the one from Panama.
Despite retaining the normal eye number, these six species are each united with the more typical members of the genus by at least one additional character, spinneret number. Most oonopids resemble other spiders in having three pairs of spinnerets. The only exceptions that have been noted to date occur in the genus Escaphiella Platnick and Dupérré (2009b); in that genus, several species are known in which the posterior median spinnerets have fused into a single structure, which may either retain or lose functional spigots, and there is one relatively large-bodied species in which those spinnerets have been lost entirely. Although we have not been able to examine males and females of every species by scanning electron microscopy, so far as we have been able to determine, all the species assigned below to Reductoonops have only four spinnerets; the posterior median spinnerets are missing entirely, and we regard this as a synapomophy for the group, paralleled (so far as we know) only within the distantly related genus Escaphiella.
Obviously, reduction characters are not ideal as evidence of monophyly, as it is certainly possible that the loss of the posterior median spinnerets has occurred more than once within the Stenoonops complex. But we know of no potentially synapomorphic features that would ally any of the six-eyed species with any group other than the remaining species of Reductoonops. The possibility remains, of course, that even if monophyletic, Reductoonops as here delimited could be simply a highly autapomorphic lineage within some other group, for which the best candidate would presumably be Stenoonops. There is one genitalic similarity that might support that hypothesis. Although most males of Stenoonops have palps with a dorsally elongated cymbium, there are three species, namely Stenoonops petrunkevitchi Chickering (1951) from Costa Rica and Panama, plus Stenoonops tayrona Platnick and Dupérré (2010a) and Stenoonops kochalkai Platnick and Dupérré (2010a) from Colombia, in which the cymbial shape resembles somewhat that of Reductoonops. Nevertheless, those Stenoonops species all seem to retain six spinnerets, and have the distal clump of setae on the palpal tarsus of both sexes that unites all the members of Stenoonops, but is not found in Reductoonops species, which have short but widely separated setae in that position. In addition, the females of those species, where known, show no special genitalic similarities to those of Reductoonops. Thus, until additional characters are found that might ally the species treated below with just some subgroup of Stenoonops, we regard both genera as putatively monophyletic. They may be sister taxa, but several other genera belonging to the Stenoonops complex have not yet been studied in detail, and hypotheses about the number and relationships of the genera belonging to that complex would be premature.
One character that has sometimes been used in the higher classification of goblin spiders is whether the leg claws have one or two rows of teeth. In most oonopids, there are two rows of teeth on the superior claws, one on the outer margin and one on the inner margin, and those rows often have teeth that differ greatly in both size and position. However, several examples are known in which one or both sexes have only a single row of teeth. At least R. yasuni is notable in that regard, as both males and females have two rows of teeth on the anterior legs, but only a single row of teeth on the posterior legs (figs. 17–24, 76–83).
Our methods follow those of Platnick and Dupérré (2009a); only differences from the males (beyond the obvious lack of male endite modifications) are mentioned in the descriptions of females. Scans were sometimes taken from uncoated right male palps; in those cases, the images were flipped for consistency. All measurements are in mm. Species are discussed in geographic order, beginning in Mexico and continuing south. Unless otherwise indicated, all new specific names are nouns in apposition taken or shortened from the type locality. High-resolution versions of the images, a sortable version of the geocoded locality data, and a distribution map for each species will be available on the goblin spider Planetary Biodiversity Inventory (PBI) projects website ( http://research.amnh.org/oonopidae). Users should note that the relatively small published images are merely avatars for the actual image files on the website, which can each be enlarged several times before pixelating; in many cases, the website hosts significantly more images for a given species than are presented here. Similarly, the maps made available on the website, via discoverlife.org, are more useful than printed versions would be, as each dot can be associated with the actual specimen data it represents.
COLLECTIONS EXAMINED
Glossary
AMNH
American Museum of Natural History, New York, NY
CAS
California Academy of Sciences, San Francisco, CA
CDU
Darrell Ubick Collection, San Francisco, CA
FMNH
Field Museum of Natural History, Chicago, IL
INBIO
Instituto Nacional de Bio diversidad, Santo Domingo, Costa Rica
MACN
Museo Argentino de Ciencias Naturales, Buenos Aires, Argentina
MCZ
Museum of Comparative Zoology, Harvard University, Cambridge, MA
MNHN
Muséum National d'Histoire Naturelle, Paris, France
QCAZ
Museum of Invertebrates, Pontificia Universidad Católica, Quito, Ecuador
TMM
Texas Memorial Museum, Austin, TX
USNM
National Museum of Natural History, Smithsonian Institution, Washington, DC
Reductoonops , new genus
Type Species: Reductoonops yasuni, new species.
Etymology: The generic name refers to the reduced size, eyes, and spinnerets, and is masculine in gender.
Diagnosis: Members of this genus resemble those of Stenoonops and other members of the Stenoonops complex, but are reduced in size, have a flattened clypeus, have only four spinnerets, and lack the distinct clump of short setae on the dorsal surface of the palpal tarsus of both sexes that characterizes Stenoonops; they often have only two eyes, and four pairs of deep channels at the sides of the sternum, the most anterior pair of which demarcate a short anterior portion of the sternum (figs. 44, 270). They differ from those of Xyccarph in lacking leg spines and usually in having the male palpal cymbium and bulb separate, rather than fused.
Description: Total length of males 0.8–1.2, of females 0.9–1.7. Carapace, sternum, mouthparts pale orange, without pattern, ventral abdominal scuta, legs yellow, without pattern, abdomen soft portions white, without pattern. Cephalothorax: Carapace piriform in dorsal view (figs. 1, 43, 55), anteriorly narrowed to 0.49 times its maximum width or less, pars cephalica flat in lateral view, anterolateral corners with slightly sclerotized triangular projections, pars thoracica with rounded posterolateral corners, without depressions or radiating rows of pits, posterolateral edge without pits, posterior margin not bulging below posterior rim, posterolateral surface without spikes, surface of elevated portion of pars cephalica usually smooth but finely reticulate in at least some sixeyed species, at least sometimes with distinct platelets (figs. 1, 55), sides finely reticulate; fovea absent, lateral margin undulate (figs. 5, 59), rebordered, without denticles; plumose setae near posterior margin of pars thoracica absent; marginal, nonmarginal pars cephalica, pars thoracica setae dark, needlelike, scattered. Clypeus margin strongly rebordered, sinuous in front view, sloping forward in lateral view (figs. 5, 59), high, ALE separated from edge of carapace by their radius or more (figs. 4, 58), median projection absent; setae dark, needlelike. Chilum absent. Eyes usually two, sometimes six or zero; when present eyes well developed, when present, ALE largest, oval, PME squared, PLE oval; when present, posterior eye row straight from both above and front, ALE-PLE touching, PME touching throughout most of their length, PLE-PME touching. Sternum Ionger than wide, apparently fused to carapace at anterior end, surface smooth, finely reticulate, or with pits, median concavity, hair tufts absent, radial furrows between coxae I–II, II–III, III–IV smooth, additional radial furrow often present between endites and coxae I (figs. 2, 44, 56), radial furrow opposite coxae III absent, sickle-shaped structures absent, anterior margin unmodified, posterior margin extending posteriorly beyond anterior edges of coxae IV as single extension, without posterior hump, anterior corner unmodified, lateral margin without infracoxal grooves, distance between coxae II and III greater than distance between coxae I and II, or coxae III and IV, extensions of precoxal triangles absent, lateral margins with three pairs of lateral projections; setae sparse, usually densest laterally but evenly spread in R. tandapi and R. hedlite, dark, needle-like, originating from surface. Chelicerae straight, anterior face unmodified; without teeth on promargin or retromargin; fangs without toothlike projections, directed medially, shape normal, without prominent basal process, tip unmodified; setae dark, needlelike, evenly scattered; paturon inner margin with short interdigitating setae, distal region abruptly narrowed (fig. 6, 60), posterior surface unmodified, promargin with row of flattened setae (fig. 7, 61), inner margin unmodified, laminate groove absent. Labium triangular, not fused to sternum, with anterior margin indented or deeply incised at middle (fig. 8), same as sternum in sclerotization; with six or more setae on anterior margin, subdistal portion with unmodified setae. Endites same as sternum in sclerotization, distally not excavated, serrula apparently present in single row in females (figs. 63, 64), absent in males (fig. 10), anterior portion with distally fork-shaped setae (fig. 62), with projecting lobes and modified setae in males (fig. 9), posterior portion unmodified. Labrum with broad basal lobe bearing setae with basal spurs (figs. 11, 65). Female palp without claw or spines (figs. 67, 68); tibia with three trichobothria (fig. 66), patella without prolateral row of ridges, tarsus unmodified. Abdomen: Cylindrical, without long posterior extension, in females contents often shrunken (at least in preserved specimens, fig. 45), rounded posteriorly, interscutal membrane without rows of small sclerotized platelets; dorsum often iridescent, possibly because of scattered platelets (figs. 3, 45, 57). Booklung covers large, ovoid, without setae, anterolateral edge unmodified; posterior spiracles connected by groove. Pedicel tube short, unmodified, scutopedicel region unmodified, abdomen not extending anterior of pedicel; plumose hairs, matted setae on anterior ventral abdomen in pedicel area, cuticular outgrowths near pedicel all absent. Dorsal scutum absent. Epigastric scutum weakly sclerotized, not surrounding pedicel, not protruding, small lateral sclerites absent, without lateral joints in females. Postepigastric scutum weakly sclerotized, yellow, short, only around epigastric furrow (figs. 12, 69), not fused to epigastric scutum, anterior margin unmodified, without posteriorly directed lateral apodemes. Spinneret scutum, supraanal scutum both absent. Abdominal setae dark, needlelike, epigastric area setae not basally thickened (fig. 52); dense patch of setae anterior to spinnerets absent. Colulus present. Spinnerets four, posterior median pair absent (figs. 14, 73); anterior lateral spinnerets bisegmented, basal segment with oblique membranous strip (figs. 13, 72), typically with one major ampullate gland spigot and three piriform gland spigots (figs. 15, 74), posterior lateral spinnerets bisegmented, typically with two terminal spigots (figs. 16, 75). Legs: Femora with subdistal constriction on dorsal surface, accompanied by straight row of setae (figs. 39, 89, 90); femur IV not thickened, same size as femora I–III, patellae and tibiae at least sometimes with platelets of varying shape (figs. 40–42, 93–96), patella plus tibia I shorter than carapace, tibia I unmodified; tibia IV specialized hairs on ventral apex, ventral scopula, metatarsi I, II mesoapical comb, metatarsi III, IV weak ventral scopula all absent. Leg spines absent (figs. 91, 92). Tarsi without inferior claw. Superior claws scanned only in R. yasuni, those on anterior legs with three large teeth on outer margin (figs. 21, 22, 80, 81), long series of small, closely spaced teeth on inner margin of males (figs. 17, 18), those teeth longer in females (figs. 76, 77), posterior legs with three large teeth on outer margin (figs. 23, 24, 82, 83), without teeth on inner margin (figs. 19, 20, 78, 79). Trichobothrial base with numerous parallel grooves (fig. 30). Tarsal organs with three receptors on legs I, II (figs. 25, 26, 84, 85), two receptors on legs III, IV, palps (figs. 27–29, 86–88), distalmost receptor bifid, with two lobes originating on common base. Genitalia: Male epigastric region with sperm pore not visible; furrow without Ω-shaped insertions, without specialized setae. Male palp minute, not strongly sclerotized, right and left palps mirror images, proximal segments, cymbium, bulb yellow; embolus light, prolateral excavation absent; trochanter normal size, unmodified; femur normal size, two or more times as long as trochanter, without posteriorly rounded lateral dilation, attaching to patella basally; patella shorter than femur, not enlarged, without prolateral row of ridges, setae unmodified; tibia with three trichobothria (fig. 33); cymbium ovoid in dorsal view, forming cup-shaped base for palpal bulb (figs. 31, 32, 46–51), usually not fused to bulb but sometimes fused with seam obvious, sometimes fused with seam reduced (at least dorsally) to row of pores, not extending beyond distal tip of bulb, plumose setae, stout setae absent, with shortened distal setae not clumped into distinct patch (fig. 34); bulb longer than cymbium, slender, elongated; embolus typically basally sinuous, bearing numerous terminal spikes (figs. 35, 36), accompanied dorsally by conductor, ventrally by palpal apophysis (figs. 37, 38), but spikes sometime absent, palpal apophysis and/or conductor sometimes fused with embolus, palpal apophysis sometimes absent. Female genitalia with distinct anterior process, typically bearing dorsal projections (fig. 71), process set in boat-shaped structure (figs. 53, 54, 70).
Distribution: Mexico to Chile, Galapagos Islands, Jamaica, Curaçao, Martinique.
Key to Species
1. Six eyes 2
— Two or no eyes 10
2. Males (those of bayano and hato unknown) 3
— Females (those of meta unknown) 6
3. Palp with three terminal elements (fig. 350); Colombia meta
— Palp with only two terminal elements (as in fig. 111); Mexico 4
4. Common base of embolus and conductor relatively long (figs. 111, 158) 5
— Common base of embolus and conductor relatively short (fig. 133) armeria
5. Common base of embolus and conductor relatively narrow (fig. 113) chamela
— Common base of embolus and conductor relatively wide (fig. 160) niltepec
6. Anterior genitalic process long, narrow (fig. 297); Curaçao hato
— Anterior genitalic process much shorter (as in fig. 120) 7
7. Tip of anterior genitalic process greatly expanded (fig. 145) niltepec
— Tip of anterior genitalic process slightly expanded (as in fig. 120) 8
8. Abdomen with strong setae (fig. 281); Panama bayano
9. Posterior portion of anterior genitalic process relatively short (fig. 120) chamela
— Posterior portion of anterior genitalic process relatively long (fig. 140) armeria
10. No eyes (figs. 192, 193); Mexico jabin
— Two eyes 11
11. Eyes enlarged, highly reflective (figs. 370, 503); Ecuador 12
— Eyes small, not highly reflective (as in fig. 43) 13
12. Males with long extension on embolar base (fig. 498); females with tip of anterior genitalic process narrowed (fig. 505) hedlite
— Males with short extension on embolar base (fig. 365); females with tip of anterior genitalic process rounded (fig. 372) tandapi
13. Males (those of diamant, elqui, escopeta, ferry, leticia, marta, nubes, and pichincha unknown) 14
— Females 30
14. Anterior margin of labium indented at middle (as in fig. 164) 15
— Anterior margin of labium deeply incised at middle (as in fig. 8) 20
15. Conductor very narrow, paralleling embolus (figs. 390, 535) 16
16. Palpal apophysis with basal projection (fig. 390) tina
— Palpal apophysis without basal projection (fig. 535) celica
18. Palpal apophysis relatively narrow (fig. 412) domingo
— Palpal apophysis relatively wide (fig. 483) jatun
19. Conductor relatively short (fig. 173); Mexico real
— Conductor relatively long (fig. 461); Ecuador napo
20. Embolus, conductor, and palpal apophysis largely fused (figs. 555, 556); Galapagos Islands pinta
— Embolus, conductor, and palpal apophysis separate 21
21. Palpal apophysis with basal projection (figs. 251, 264) 22
— Palpal apophysis without basal projection 23
22. Basal portion of embolus directed distally (fig. 251); Costa Rica lucha
— Basal portion of embolus directed laterally (fig. 264); Panama almirante
23. Basal portion of embolus directed laterally (figs. 36, 427, 449, 523) 24
— Basal portion of embolus directed proximally (figs. 211, 234, 336, 569) 27
24. Palpal apophysis very narrow (fig. 449) palenque
25. Palpal bulb with projection near base of terminal elements (fig. 523) molleturo
26. Base of embolus with elevated ridge (fig. 427) otonga
— Base of embolus without elevated ridge (fig. 36) yasuni
— Palpal apophysis flattened, leaflike (fig. 234) naci
28. Embolar base enlarged, with lateral projection (fig. 336) sasaima
29. Palpal apophysis relatively narrow (fig. 211); Costa Rica monte
— Palpal apophysis wider (fig. 569); Peru carpish
30. Anterior margin of labium deeply incised at middle (as in fig. 8) 31
— Anterior margin of labium indented at middle (as in fig. 164) 36
31. Tip of anterior genitalic process expanded (figs. 278, 323, 380) 32
— Tip of anterior genitalic process narrowed (figs. 54, 218, 577) 34
32. Tip of anterior genitalic process short (figs. 323, 380) 33
— Tip of anterior genitalic process longer (fig. 278); Panama escopeta
33. Anterior genitalic process basally wide (fig. 323); Colombia leticia
— Anterior genitalic process basally narrow (fig. 380); Ecuador pichincha
34. Anterior genitalic process relatively wide (fig. 577); Peru carpish
— Anterior genitalic process relatively narrow (figs. 54, 218) 35
35. Anterior genitalic process abruptly narrowed at tip (fig. 218); Costa Rica monte
— Anterior genitalic process gradually narrowed at tip (fig. 54); Ecuador yasuni
36. Anterior genitalic process basally wide (fig. 340); Colombia sasaima
— Anterior genitalic process basally narrower 37
37. Anterior genitalic process relatively short, wide (figs. 183, 313, 473, 547, 582) 38
— Anterior genitalic process relatively long, narrow (as in figs. 198, 223, 273, 468) 42
38. Anterior genitalic process with a distal cap (figs. 313, 473, 547, 582) 39
— Anterior genitalic process without a distal cap (fig. 183); Mexico real
39. Cap of anterior genitalic process much wider than rest of process (fig. 547); Galapagos Islands pinta
— Cap of anterior genitalic process not much wider than rest of process (figs. 313, 473, 582) 40
40. Cap of anterior genitalic process relatively long (fig. 313); Martinique diamant
— Cap of anterior genitalic process shorter (figs. 473, 582) 41
41. Anterior genitalic process narrowed behind cap (fig. 473); Ecuador jatun
— Anterior genitalic process not narrowed (fig. 582); Chile elqui
42. Anterior genitalic process distinctly narrowed at tip (figs. 188, 198, 273, 439) 43
— Anterior genitalic process at most slightly narrowed at tip (figs. 223, 297, 318, 397, 402, 434, 468, 510, 542) 46
43. Middle portion of anterior genitalic process greatly widened (fig. 188); Mexico nubes
— Middle portion of anterior genitalic process not widened (figs. 198, 273, 439) 44
44. Anterior genitalic process bulging on one side (figs. 198, 273) 45
— Anterior genitalic process not bulging (fig. 439); Ecuador palenque
45. Anterior genitalic process set in relatively long boat-shaped structure (fig. 198); Costa Rica lucha
— Anterior genitalic process set in relatively short boat-shaped structure (fig. 273); Panama almirante
46. Anterior genitalic process with widened tip (figs. 397, 402, 468) 47
— Tip of anterior genitalic process not widened (figs. 223, 297, 318, 434, 510, 542) 49
47. Tip of anterior genitalic process with pair of lateral extensions (fig. 468) napo
— Tip of anterior genitalic process without lateral extensions (figs. 397, 402) 48
48. Anterior genitalic process sinuous (fig. 397) tina
— Anterior genitalic process straight (fig. 402) domingo
49. Anterior genitalic process set in boat-shaped structure with steep sides (figs. 318, 434) 50
— Anterior genitalic process set in boat-shaped structure with low sides (figs. 223, 297, 510, 542) 51
50. Tip of anterior genitalic process rounded (fig. 318); Colombia marta
— Tip of anterior genitalic process squared (fig. 434); Ecuador otonga
51. Tip of anterior genitalic process rounded (fig. 542) celica
— Tip of anterior genitalic process not rounded (figs. 223, 297, 510) 52
52. Anterior genitalic process relatively short, wide (fig. 297); Jamaica ferry
— Anterior genitalic process relatively long, narrow (figs. 223, 510) 53
53. Anterior genitalic process set in relatively long boat-shaped structure (fig. 510); Ecuador molleturo
— Anterior genitalic process set in relatively short boat-shaped structure (fig. 223); Costa Rica and Panama naci
Reductoonops chamela , new species (figs. 97–120)
Types: Male holotype, female allotype, two male paratypes, and two female paratypes taken from a Berlese sample of forest litter taken at an elevation of 150–300 ft in the Estación de Biología Chamela, at km 59 on Route 200, north of La Barra de Navidad, Jalisco, Mexico (Apr. 4–7, 1985; R. Schuh, B. Massie), deposited in AMNH (PBI_OON 1463).
Diagnosis: Members of this species resemble those of R. armeria and R. niltepec in retaining the four posterior eyes. Males differ from those of R. armeria in having a much longer base subtending both the embolus and conductor, and from those of R. niltepec in having that base narrower (figs. 110–113). Females differ from those of R. armeria in having the posterior portion of the anterior genitalic process much shorter, and from those of R. niltepec in having that portion of the process wider distally than proximally (figs. 119, 120).
Male (PBI_OON 1463, figs. 97–113): Total length 0.95. Surface of elevated portion of pars cephalica finely reticulate. Eyes six; ALE touching. Sternum without anterior channels, surface finely reticulate, without pits, microsculpture only at sides; setae densest laterally. Labium anterior margin indented at middle. Anterolateral edge of endites with sharply pointed, laterally directed projection. Embolus and conductor originating from single long, wide base; cymbium not fused with bulb.
Female (PBI_OON 1463, figs. 114–120): Total length 1.31. Anterior genitalic process bipartite, posterior portion about twice as long as anterior portion, widest anteriorly.
Other Material Examined: MEXICO: Jalisco: La Manganilla, 12 mi NW La Barra de Navidad, Sept. 23, 1973, Berlese, litter near logs with fungus (A. Newton, FMNH 56114, PBI_OON 10724), 4♂.
Distribution: Mexico (Jalisco).
Reductoonops armeria , new species (figs. 121–140)
Types: Male holotype and male paratype from Las Humedades, Armería, Colima, Mexico (Jan. 19, 1943; F. Bonet), deposited in AMNH (PBI_OON 31161).
Diagnosis: Members of this species resemble those of R. chamela and R. niltepec in retaining the four posterior eyes. Males differ from those of R. chamela and R. niltepec in having a much shorter base subtending both the embolus and conductor (figs. 132–135). Females differ from those of R. chamela in having the posterior portion of the anterior genitalic process much longer, and from those of R. niltepec in having that portion of the process wider distally than proximally (figs. 139, 140).
Male (PBI_OON 31161, figs. 121–136): Total length 1.02. Surface of elevated portion of pars cephalica finely reticulate. Eyes six; ALE touching. Sternum without anterior channels, surface finely reticulate, without pits, microsculpture only at sides; setae densest laterally. Labium anterior margin indented at middle. Endite tip with rounded, laterally directed lobe. Combined base of embolus and conductor short, conductor narrow, embolus sharply bent at three-fourths its length; cymbium not fused with bulb.
Female (PBI_OON 37320, figs. 137–140): Total length 1.19. Anterior portion of anterior genitalic process subdistally expanded, planaria shaped.
Other Material Examined: MEXICO: Colima: 12 mi E Manzanillo, 19°01′N, 104°10′W, May 11, 1963 (W. Gertsch, W. Ivie, AMNH PBI_OON 37320), 2♂, 2♀.
Distribution: Mexico (Colima).
Reductoonops niltepec , new species (figs. 141–160)
Types: Male holotype and male paratype taken 2 miles southeast of Niltepec, 16°32′N, 94°33′W, Oaxaca, Mexico (Aug. 16, 1966; J., W. Ivie), deposited in AMNH (PBI_OON 37321).
Diagnosis: Members of this species resemble those of R. chamela and R. armeria in retaining the four posterior eyes. Males differ from those of R. armeria in having a much longer base subtending both the embolus and conductor, and from those of R. chamela in having that base wider (figs. 157–160). Females differ from those of R. chamela and R. armeria in having the posterior portion of the anterior genitalic process wider proximally than distally (figs. 144, 145).
Male (PBI_OON 37321, figs. 141, 146–160): Total length 1.03. Surface of elevated portion of pars cephalica finely reticulate. Eyes six; ALE touching. Sternum without anterior channels, surface finely reticulate, without pits, microsculpture only at sides; setae densest laterally. Labium anterior margin indented at middle. Endite tip with squared, anteromedially directed lobe. Embolus and conductor originating from long, wide common base; cymbium not fused with bulb.
Female (PBI_OON 21105, figs. 142–145): Total length 1.15. Anterior genitalic process with long posterior portion wider posteriorly than anteriorly, short anterior portion greatly widened.
Other Material Examined: MEXICO: Oaxaca: 20 mi S Juchetenango, May 29, 1971, Berlese, oak litter, elev. 6000 ft (S. Peck, AMNH PBI_OON 21105), 2♀; 11.6 km N junction Routes 190 and 135, on 135, July 21, 1987, Berlese, wet oak forest (R. Anderson, FMNH 61606, PBI_OON 10892), 1 ♂; 3 mi SE Tlacolula, 16°56′N, 96°25′W, Aug. 30, 1966 (J., W. Ivie, AMNH PBI_OON 37322), 1♂.
Distribution: Mexico (Oaxaca).
Reductoonops real , new species (figs. 161–183)
Type: Male holotype from El Real, Chiapas, Mexico (July 3, 1950; C., M. Goodnight), deposited in AMNH (PBI_OON 1432).
Diagnosis: Members of this species resemble those of the majority of species in having only two eyes, but show no trace of the anterior sternal channels found in most of those species (figs. 163, 176, 177). The male palp shows the typically sinuous embolus, but the embolus tip bears only a single projection (best seen in dorsal view, fig. 175), rather than the group of projections found in other species. The female anterior genitalic process is of the typically unipartite form, but is shorter and more ovoid than in other species (figs. 182, 183).
Male (PBI_OON 1432, figs. 161–176): Total length 1.22. Surface of elevated portion of pars cephalica smooth. Eyes two; ALE touching. Sternum without anterior channels, surface smooth, without pits, micro sculpture absent; setae densest laterally. Labium anterior margin indented at middle. Endite tip with slight lobe, laterally protruding, anteromedially directed macro seta. Embolus sinuous, tip with single spur; cymbium fused with bulb but with clearly defined seam between.
Female (PBI_OON 1467, figs. 177–183): Total length 0.98. Anterior genitalic projection relatively short, unipartite, torpedo shaped.
Other Material Examined: MEXICO: Chiapas: 5 mi NE Chiapa, 16°45′N, 92°58′W, Aug. 26, 1966, hillside (J., W. Ivie, AMNH PBI_OON 1467), 2 ♀; Ruinas de Palenque, Apr. 3, 1974 (C. Alteri, AMNH PBI_OON 1466), 1 ♀; Plateau 6 mi S Tuxtla Gutiérrez, 16°42′N, 93°07′W, Aug. 21, 1966 (J., W. Ivie, AMNH PBI_OON 1468), 3 ♀.
Distribution: Mexico (Chiapas).
Reductoonops nubes , new species (figs. 184–188)
Type: Female holotype from Berlese sample of litter taken in a coffee grove at an elevation of 5800 ft at Las Nubes, Volcán de Tacana, Chiapas, Mexico (Aug. 6, 1950; C., M. Goodnight), deposited in AMNH (PBI_OON 1447).
Diagnosis: These females have two eyes (separating them from R. chamela, R. armeria, and R. niltepec, which have six eyes, and from R. jabin, which lack eyes) and anterior sternal channels (separating them from R. real). The anterior genitalic process is long and greatly narrowed anteriorly (figs. 187, 188).
Male: Unknown.
Female (PBI_OON 1469, figs. 184–188): Total length 1.46. Surface of elevated portion of pars cephalica smooth. Eyes two; ALE touching. Sternum with anterior channels, surface smooth, micro sculpture absent, setae densest laterally. Labium anterior margin indented at middle. Anterior genitalic projection unipartite, basally ovoid, distally narrowed.
Other Material Examined: MEXICO: Chiapas: Cacahuatán, Las Nubes, Aug. 6, 1950, Berlese, coffee area, elev. 5800 ft (C., M. Goodnight, AMNH PBI_OON 1675), 1 ♀ (without abdomen, so genitalia not examined); Unión Juárez, Aug. 11, 1950 (C., M. Goodnight, AMNH PBI_OON 1469), 1 ♀.
Distribution: Mexico (Chiapas).
Reductoonops jabin , new species (figs. 189–196)
Type: Female holotype from Cenote (sinkhole) Jabin, 1.5 km W Kaua, Yucatán, Mexico (Jan. 5, 2003; J. Reddell, M. Reyes), deposited in TMM (34502, PBI_OON 21100).
Diagnosis: Members of this species differ from those of all other Reductoonops in being completely blind (figs. 192, 193). The anterior genitalic process is relatively short and wide (figs. 195, 196). One other completely blind oonopid has been described from Yucatán, Wanops coecus, but that species is very different, lacking sternal channels and having highly spinose legs.
Male: Unknown.
Female (PBI_OON 21100, figs. 189–196): Total length 1.66. Surface of elevated portion of pars cephalica smooth. Eyes absent. Sternum with anterior channels, surface smooth, microsculpture absent; setae densest laterally. Labium anterior margin indented at middle. Anterior genitalic projection relatively short, wide.
Other Material Examined: None.
Distribution: Mexico (Yucatán).
Reductoonops monte , new species (figs. 199–218)
Types: Male holotype and female allotype taken by sifting leaf litter at an elevation of 1500 m at Monteverde, Puntarenas, Costa Rica (Feb. 23–27, 1991; H., A. Howden), deposited in AMNH (PBI_OON 1435).
Diagnosis: Males can be recognized by the hook-shaped process on the palpal apophysis, best seen in prolateral view (fig. 210), females by the very wide arms at the base of the anterior genitalic process (figs. 217, 218). The species occurs sympatrically with R. naci, but differs from that and all other known species in these characters.
Male (PBI_OON 1435, figs. 199–213): Total length 0.95. Surface of elevated portion of pars cephalica smooth. Eyes two; ALE touching. Sternum with anterior channels, surface smooth, without pits, microsculpture absent; setae densest laterally. Labium anterior margin deeply incised. Endites with bipartite distal lobe situated beside short, modified seta situated at anterolateral corner. Palpal apophysis with hook-shaped projection; cymbium not fused with bulb.
Female (PBI_OON 1435, figs. 214–218): Total length 1.34. Anterior genitalic process long, narrow, base with wide arms bearing relatively straight anterior margin.
Other Material Examined: None
Distribution: Costa Rica (northern Puntarenas).
Reductoonops naci , new species (figs. 219–238)
Types: Male holotype, male paratype, female allotype, and two female paratypes taken from humus at an elevation of 950 m in the Nacimiento Parque Municipal, 9°58′57″N, 84°10′28″W, Heredia, Costa Rica (Jan. 27–28, 2010; C. Víquez, B. Hernandez, A. Solis), deposited in INBIO (PBI_OON 1618).
Diagnosis: Males can be recognized by the distally pointed apophysis on the palp (figs. 233– 238), females by the bowl-shaped arms at the base of the anterior genitalic process (figs. 222, 223).
Male (PBI_OON 1618, figs. 224–238): Total length 0.97. Surface of elevated portion of pars cephalica smooth. Eyes two; ALE touching. Sternum with anterior channels, surface smooth, without pits, microsculpture absent; setae densest laterally. Labium anterior margin deeply incised. Endites with bipartite distal lobe situated beside short, modified seta situated at anterolateral corner. Palpal apophysis leaf shaped, with distal point; cymbium not fused with bulb.
Female (PBI_OON 1618, figs. 219–223): Total length 0.99. Labium anterior margin indented at middle. Anterior genitalic process very long, narrow, set in bowl-shaped base.
Other Material Examined: COSTA RICA: Cartago: Estación Biológica Barbilla, Parque Nacional Barbilla, Jan. 19, 2001, humus, elev. 600 m (W. Arana, INBIO PBI_OON 30982), 1 ♂, 1 ♀; Parque Nacional Tapanti, June 4, 1997, litter, tropical montane evergreen forest, elev. 1500 m (R. Anderson, INBIO PBI_OON 30983), 1 ♀; Reserva Forestal Río Macho, km 70, Inter-American Highway, 9°39′N, 83°51′W, Mar. 22–26, 1999, sifting forest litter, elev. 2850 m (J. Miller, USNM PBI_OON 27798), 2 ♀. Limón: Finca Alberto Moore, Hone Creek, July 8–11, 2005, humus, cacao/cordia/banana (C. Víquez, INBIO PBI_OON 29683), 1 ♀; Valle del Silencio, Mar. 23, 2003 (C. Víquez, INBIO 73778, PBI_OON 49622), 1 ♀. Puntarenas: Estación Biológica Las Cruces, 8°46′N, 82°58′W, Mar. 14, 1973, Berlese, sawdust on and under slabs, old sawmill site, elev. 4000 ft (J. Wagner, J. Kethley, FMNH 33528, PBI_OON 10035), 2 ♂, 4 ♀, Mar. 14–15, 1973, Berlese, leaf litter, elev. 4000 ft (J. Wagner, J. Kethley, FMNH 33523, 33530, 71271, PBI_OON 10030, 10037, 10630, 10649, 38427, 43489), 5 ♂, 17 ♀, Mar. 16, 1973, forest floor litter above stream, elev. 4000 ft (J. Wagner, J. Kethley, FMNH 33521, ex 56586, PBI_OON 10028, 37035, 37976), 3 ♂, 10 ♀, Mar. 18, 1973, Berlese, virgin forest floor litter, moderate slope, elev. 4000 ft (J. Wagner, J. Kethley, FMNH 33514, 56584, PBI_OON 10021, 10799), 11 ♀, Mar. 19, 1973, Berlese, leaf litter in stream bed, elev. 4000 ft (J. Wagner, J. Kethley, FMNH 33529, PBI_OON 10036), 1 ♀, Berlese, under bark, rotten log galleries, elev. 4000 ft (J. Wagner, J. Kethley, FMNH 33515, PBI_OON 10022), 1 ♀, Mar. 20, 1973, Berlese, sawdust under plank, elev. 4000 ft (J. Wagner, J. Kethley, FMNH 33517, PBI_OON 10024), 2 ♂, 6 ♀, Mar. 21, 1973, Berlese, mixed floor litter, elev. 4700 ft (J. Wagner, J. Kethley, FMNH 33519, PBI_OON 10026), 4 ♂, 8 ♀; Estación Sirena, Península de Osa, May 18–30, 2001, humus (A. Azofeifa, INBIO PBI_OON 30978), 1 ♀; Fundación Neotrópica, 10 km W Rincón, Península de Osa, 8°42′30″N, 83°31′30″W, June 23, 1997, ridge forest litter, elev. 180 m (R. Anderson, INBIO PBI_OON 51308), 1 ♂, 1 ♀; Isla del Coco, Apr. 19, 2013 (C. Víquez, INBIO 106123, PBI_OON 49621), 1 ♂; Monteverde, Aug. 24, 1983, roadside scrub, elev. 1500 m (J., F. Murphy, AMNH PBI_OON 36826), 1 ♂, Feb. 23–27, 1991, sifting leaf litter, elev. 1500 m (H., A. Howden, AMNH PBI_OON 49605), 1 ♂; Parque Nacional Manuel Antonio, June 19, 1976 (C., M. Goodnight, AMNH PBI_OON 1430), 2 ♀; Quepos, Cerro Nara, 9°29′03″N, 84°09′12″W, Aug. 2002, under rock, elev. 900 m (C. Víquez, INBIO PBI_OON 30981), 1 ♀ ; Rancho Quemado, Península de Osa, Mar. 1992 (F. Quesada, INBIO 45746, PBI_OON 30979), 1 ♀. San José: Parque Nacional Braulio Carrillo, Apr. 28–30, 1983, nature trail along a cloud forest/rainforest transect, elev. 1100 m (D. Ubick, CDU PBI_OON 3635), 1 ♀. PANAMA: Chiriquí: Cerro Punta, Mar. 6, 1959, Berlese, old mossy log, elev. 6250 ft (H. Dybas, FMNH 56888, PBI_OON 10831), 1 ♀ ; on road to Cerro Punta, Mar. 8, 1959, Berlese, damp litter near spring, elev. 5600 ft (H. Dybas, FMNH 33660, PBI_OON 10162), 1 ♂; 2 km E Cerro Punta, June 2, 1977, moss and lichens on trees, elev. 2400 m (H., A. Howden, AMNH PBI_OON 37062), 1 ♂; 5 mi W El Hato del Volcán, Aug. 10, 1983, rainforest edge, elev. 1000 m (J., F. Murphy, AMNH PBI_OON 36827), 1 ♀ ; 12 km W El Hato del Volcán, June 27, 1976, Berlese, cloud forest litter, elev. 4500 ft (A. Newton, MCZ 72911, PBI_OON 28118), 2 ♀; El Volcán, Feb. 19, 1936 (W. Gertsch, AMNH PBI_OON 1464), 1 ♀, Aug. 9–14, 1950 (A. Chickering, MCZ PBI_OON 51313), 2 ♀ ; Finca Hartmann, 15 km NW Santa Clara, May 20, 1977, elev. 1200 m (S., J. Peck, FMNH 33645, PBI_OON 10147), 2 ♂, 1 ♀; Finca Lerida, near Boquete, Mar. 17, 1959, Berlese, concentrated forest floor litter, elev. 7800 ft (H. Dybas, FMNH 56508, PBI_OON 10750), 1 ♂, 3 ♀, Berlese, forest floor litter under palms, elev. 7750 ft (H. Dybas, FMNH PBI_OON 51314), 1 ♂, Mar. 18, 1959, Berlese, concentrated floor debris in damp ravine, elev. 6900 ft (H. Dybas, FMNH 56559, PBI_OON 10778), 1 ♀; Finca Palo Santo, W Nueva California, Mar. 5, 1959, Berlese, chips, lead mold at base of cut stump, elev 4750 ft (H. Dybas, FMNH PBI_OON 10182), 1 ♀, Mar. 9, 1959, Berlese, forest floor litter in deep ravine, elev. 4900 ft (H. Dybas, FMNH PBI_OON 51312), 1 ♂, Mar. 10, 1959, Berlese, forest floor litter at base of log and cut stump, elev. 5000 ft (H. Dybas, FMNH PBI_OON 51311), 2 ♂, 5 ♀ ; Parque Nacional Volcán Barú, 5.9 km E Cerro Punta, June 14, 1995, oak ridge bamboo forest litter, elev. 2400 m (R. Anderson, AMNH PBI_OON 51310), 1 ♀. Darién: Chucanti, Nov. 11–14, 2009 (R. Miranda, S. Fernandez, INBIO PBI_OON 51316), 3 ♂, 1 ♀.
Distribution: Widespread in Costa Rica (including Cocos Island) and Panama.
Reductoonops lucha , new species (figs. 197, 198, 239–253)
Types: Male holotype, three male paratypes, female allotype, and female paratype taken at an elevation of 1500 m at the Project Darwin base camp, La Lucha, Cerro Amuo, 9.114388°N, 83.093421°W, Puntarenas, Costa Rica (Feb. 19–27, 2008; C. Víquez), deposited in INBIO (PBI_ OON 31133).
Diagnosis: Males resemble those of the Panamanian species R. almirante in having a triangular palpal apophysis, but have a much less sinuous embolus (figs. 250–253); females also resemble those of R. almirante in having an asymmetrical anterior genitalic process, but that process is longer and is situated on a much longer base (figs. 197, 198).
Male (PBI_OON 31133, figs. 239, 242–253): Total length 1.06. Surface of elevated portion of pars cephalica smooth. Eyes two; ALE touching. Sternum with anterior channels, surface smooth, without pits, micro sculpture absent; setae densest laterally. Labium anterior margin deeply incised. Endite tip with narrow, posteromedially directed lobe situated distal to short, modified seta on enlarged base. Palpal apophysis triangular, embolar base only slightly sinuous; cymbium not fused with bulb.
Female (PBI_OON 30980, figs. 197, 198, 240, 241): Total length 1.45. Labium anterior margin indented at middle. Anterior genitalic process long, asymmetrical.
Other Material Examined: COSTA RICA: Puntarenas: Cerro Frantzius, Feb. 26–28, 2005, elev. 2200 m (R. Anderson, INBIO PBI_OON 30980), 1 ♀.
Distribution: Costa Rica (southeastern Puntarenas).
Reductoonops almirante , new species (figs. 254–266, 269–273)
Types: Male holotype, male paratype, female allotype, and female paratype from Berlese sample of overflow of debris on ground from split tree hole 6 ft up on the trail to the dam on Nigua Creek, Almirante, Bocas del Toro, Panama (Mar. 25, 1959; H. Dybas), deposited in FMNH (PBI_OON10395).
Diagnosis: Males resemble those of the Costa Rican species R. lucha in having a triangular palpal apophysis, but have a much more sinuous embolus (figs. 263–266); females also resemble those of R. lucha in having an asymmetrical anterior genitalic process, but that process is shorter and is situated on a much shorter base (figs. 272, 273).
Male (PBI_OON 10395, figs. 254–266, 269): Total length 1.00. Surface of elevated portion of pars cephalica smooth. Eyes two; ALE touching. Sternum with anterior channels, surface smooth, without pits, microsculpture; setae densest laterally. Labium anterior margin deeply incised. Endites with bipartite terminal lobe, proximal portion with two modified setae. Palpal apophysis triangular, basal portion of embolus highly sinuous; cymbium not fused with bulb.
Female (PBI_OON 10395, figs. 270–273): Total length 1.17. Labium anterior margin indented at middle. Anterior genitalic process asymmetrical, situated on short base.
Other Material Examined: PANAMA: Bocas del Toro: Almirante, Apr. 1, 1959, Berlese, thatch on nest of snapping ant (H. Dybas, FMNH 33890, PBI_OON 10392), 1♂; Almirante, trail to left of trail to dam on Nigua Creek, Mar. 30, 1959, Berlese, concentrated floor litter on hill (H. Dybas, FMNH PBI_OON 10402), 1♀.
Distribution: Panama (Bocas del Toro).
FIGURES 1–12.
Reductoonops yasuni, new species, male. 1. Carapace, dorsal view. 2. Sternum, ventral view. 3. Abdomen, dorsal view. 4. Carapace, anterior view. 5. Same, lateral view. 6. Chelicerae, anterior view. 7. Same, posterior view. 8. Labium and endites, ventral view. 9. Tip of endite, ventral view. 10. Labrum and endites, dorsal view. 11. Labrum, dorsal view. 12. Epigastric region, ventral view.
FIGURES 13–27.
Reductoonops yasuni, new species, male. 13. Spinnerets, ventral view. 14. Same, posterior view. 15. Anterior lateral spinneret, posterior view. 16. Posterior lateral spinneret, posterior view. 17. Claws of leg I, apical view. 18. Same, leg II. 19. Same, leg III. 20. Same, leg IV. 21. Claws of leg I, lateral view. 22. Same, leg II. 23. Same, leg III. 24. Same, leg IV. 25. Tarsal organ from leg I, dorsal view. 26. Same, leg II. 27. Same, leg III.
FIGURES 28–42.
Reductoonops yasuni, new species, male. 28. Tarsal organ from leg IV, dorsal view. 29. Same, palp. 30. Trichobothrial base from metatarsus II, dorsal view. 31. Palp, prolateral view. 32. Same, retrolateral view. 33. Palpal tibia, dorsal view. 34. Palpal tarsus, dorsal view. 35. Embolus, prolateral view. 36. Same, ventral view. 37. Same, retrolateral view. 38. Same, dorsal view. 39. Femur IV, dorsal view. 40. Patella I, same. 41. Tibia I, same. 42. Tibia IV, same.
FIGURES 43–54.
Reductoonops yasuni, new species, male (43, 44, 46–51) and female (45, 52–54). 43. Carapace, dorsal view. 44. Sternum, ventral view. 45. Abdomen, ventral view. 46, 49. Palp, prolateral view. 47, 50. Same, ventral view. 48, 51. Same, retrolateral view. 52. Epigastric area, ventral view. 53. Genitalia, ventral view. 54. Same, dorsal view.
FIGURES 55–66.
Reductoonops yasuni, new species, female. 55. Carapace, dorsal view. 56. Sternum, ventral view. 57. Abdomen, dorsal view. 58. Carapace, anterior view. 59. Same, lateral view. 60. Chelicerae, anterior view. 61. Same, posterior view. 62. Labium and endites, ventral view. 63. Labrum and endites, dorsal view. 64. Serrula dorsal view. 65. Labrum, dorsal view. 66. Palpal tibia, dorsal view.
FIGURES 67–81.
Reductoonops yasuni, new species, female. 67. Palp, prolateral view. 68. Same, retrolateral view. 69. Epigastric region, ventral view. 70. Genitalia, dorsal view. 71. Anterior genitalic process, dorsal view. 72. Spinnerets, ventral view. 73. Same, posterior view. 74. Anterior lateral spinneret, posterior view. 75. Posterior lateral spinneret, posterior view. 76. Claws of leg I, apical view. 77. Same, leg II. 78. Same, leg III. 79. Same, leg IV. 80. Claws of leg I, lateral view. 81. Same, leg II.
FIGURES 82–96.
Reductoonops yasuni, new species, female. 82. Claws of leg III, lateral view. 83. Same, leg IV. 84. Tarsal organ from leg I, dorsal view. 85. Same, leg II. 86. Same, leg III. 87. Same, leg IV. 88. Same, palp. 89. Femur I, dorsal view. 90. Same, femur IV. 91. Tibia and metatarsus I, ventral view. 92. Same, leg IV. 93. Patella I, dorsal view. 94. Same, patella IV. 95. Tibia I, dorsal view. 96. Same, tibia IV.
FIGURES 97–109.
Reductoonops chamela, new species, male. 97. Carapace, dorsal view. 98. Same, anterior view. 99. Eye group, anterior view. 100. Sternum, ventral view. 101. Labium and endites, ventral view. 102. Tip of endite, ventral view. 103. Spinnerets, posterior view. 104. Palp, prolateral view. 105. Same, ventral view. 106. Same, retrolateral view. 107. Same, dorsal view. 108. Palpal bulb, prolateral view. 109. Same, retrolateral view.
FIGURES 110–120.
Reductoonops chamela, new species, male (110–113) and female (114–120). 110. Embolus, prolateral view. 111. Same, ventral view. 112. Same, retrolateral view. 113. Same, dorsal view. 114. Eye group, anterior view. 115. Spinnerets, posterior view. 116, 117. Sternum, ventral view. 118. Abdomen, ventral view. 119. Genitalia, ventral view. 120. Same, dorsal view.
FIGURES 121–135.
Reductoonops armeria, new species, male. 121. Carapace, anterior view. 122. Eye group, anterior view. 123. Spinnerets, posterior view. 124. Labium and endites, ventral view. 125. Tip of endite, ventral view. 126. Palp, prolateral view. 127. Same, ventral view. 128. Same, retrolateral view. 129. Same, dorsal view. 130. Palpal bulb, prolateral view. 131. Same, retrolateral view. 132. Embolus, prolateral view. 133. Same, ventral view. 134. Same, retrolateral view. 135. Same, dorsal view.
FIGURES 136–145.
136–140. Reductoonops armeria, new species, male (136) and female (137–140). 141–145. R. niltepec, new species, male (141) and female (142–145). 136, 137, 141, 142. Sternum, ventral view. 138, 143. Abdomen, ventral view. 139, 144. Genitalia, ventral view. 140, 145. Same, dorsal view.
FIGURES 146–160.
Reductoonops niltepec, new species, male. 146. Carapace, anterior view. 147. Eye group, anterior view. 148. Spinnerets, posterior view. 149. Labium and endites, ventral view. 150. Tip of endite, ventral view. 151. Palp, prolateral view. 152. Same, ventral view. 153. Same, retrolateral view. 154. Same, dorsal view. 155. Palpal bulb, prolateral view. 156. Same, retrolateral view. 157. Embolus, prolateral view. 158. Same, ventral view. 159. Same, retrolateral view. 160. Same, dorsal view.
FIGURES 161–175.
Reductoonops real, new species, male. 161. Carapace, dorsal view. 162. Eye group, anterior view. 163. Anterior portion of sternum, ventral view. 164. Labium and endites, ventral view. 165. Tip of endite, ventral view. 166. Palp, prolateral view. 167. Same, ventral view. 168. Same, retrolateral view. 169. Same, dorsal view. 170. Palpal bulb, prolateral view. 171. Same, retrolateral view. 172. Embolus, prolateral view. 173. Same, ventral view. 174. Same, retrolateral view. 175. Same, dorsal view.
FIGURES 176–188.
176–183. Reductoonops real, new species, male (176) and female (177–183). 184–188. R. nubes, new species, female. 176, 177, 184. Sternum, ventral view. 178, 185. Abdomen, ventral view. 179. Carapace, anterior view. 180. Eye group, anterior view. 181. Spinnerets, posterior view. 182, 187. Genitalia, ventral view. 183, 188. Same, dorsal view. 186. Epigastric area, ventral view.
FIGURES 189–198.
189–196. Reductoonops jabin, new species, female. 197, 198. R. lucha, new species, female. 189, 190. Sternum, ventral view. 191. Abdomen, ventral view. 192. Carapace, anterior view. 193. Same, closeup, showing absence of eyes. 194. Spinnerets, posterior view. 195, 197. Genitalia, ventral view. 196, 198. Same, dorsal view.
FIGURES 199–213.
Reductoonops monte, new species, male. 199, 200. Sternum, ventral view. 201. Spinnerets, posterior view. 202. Labium and endites, ventral view. 203. Tip of endite, ventral view. 204. Palp, prolateral view. 205. Same, ventral view. 206. Same, retrolateral view. 207. Same, dorsal view. 208. Palpal bulb, prolateral view. 209. Same, retrolateral view. 210. Embolus, prolateral view. 211. Same, ventral view. 212. Same, retrolateral view. 213. Same, dorsal view.
FIGURES 214–223.
214–218. Reductoonops monte, new species, female. 219–223. R. naci, new species, male (219) and female (220–223). 214, 219, 220. Sternum, ventral view. 215, 221. Abdomen, ventral view. 216. Epigastric area, ventral view. 217, 222. Genitalia, ventral view. 218, 223. Same, dorsal view.
FIGURES 224–238.
Reductoonops naci, new species, males (227–236, specimen from Heredia, Costa Rica; 237, specimen from Cartago, Costa Rica; 238, specimen from Darién, Panama). 224. Spinnerets, posterior view. 225. Labium and endites, ventral view. 226. Tip of endite, ventral view. 227. Palp, prolateral view. 228. Same, ventral view. 229. Same, retrolateral view. 230. Same, dorsal view. 231. Palpal bulb, prolateral view. 232. Same, retrolateral view. 233. Embolus, prolateral view. 234, 237, 238. Same, ventral view. 235. Same, retrolateral view. 236. Same, dorsal view.
FIGURES 239–253.
Reductoonops lucha, new species, male (239, 242–253) and female (240, 241). 239, 240. Sternum ventral view. 241. Abdomen, ventral view. 242. Labium and endites, ventral view. 243. Tip of endite, ventral view. 244. Palp, prolateral view. 245. Same, ventral view. 246. Same, retrolateral view. 247. Same, dorsal view. 248. Palpal bulb, prolateral view. 249. Same, retrolateral view. 250. Embolus, prolateral view. 251. Same, ventral view. 252. Same, retrolateral view. 253. Same, dorsal view.
FIGURES 254–268.
254–266. Reductoonops almirante, new species, male. 261, 268. R. escopeta, new species, female. 254, 268. Spinnerets, posterior view. 255. Labium and endites, ventral view. 256. Tip of endite, ventral view. 257. Palp, prolateral view. 258. Same, ventral view. 259. Same, retrolateral view. 260. Same, dorsal view. 261. Palpal bulb, prolateral view. 262. Same, retrolateral view. 263. Embolus, prolateral view. 264. Same, ventral view. 265. Same, retrolateral view. 266. Same, dorsal view. 267. Eye group, anterior view.
FIGURES 269–278.
269–273. Reductoonops almirante, new species, male (269) and female (270–273). 274– 278. R. escopeta, new species, female. 269, 270, 274, 275. Sternum, ventral view. 271, 276. Abdomen, ventral view. 272, 277. Genitalia, ventral view. 273, 278. Same, dorsal view.
FIGURES 279–289.
Reductoonops bayano, new species, female. 279, 280. Sternum, ventral view. 281. Abdomen, ventral view. 282. Carapace, dorsal view. 283. Same, lateral view. 284. Habitus, anterior view. 285. Same, lateral view. 286. Eye group, anterior view. 287. Spinnerets, posterior view. 288. Genitalia, ventral view. 289. Same, dorsal view.
FIGURES 290–300.
290–297. Reductoonops ferry, new species, female. 298–300. R. hato, new species, female. 290, 291, 298, 299. Sternum, ventral view. 292, 300. Abdomen, ventral view. 293. Labium and endites, ventral view. 294. Spinnerets, posterior view. 295. Epigastric region, ventral view. 296. Genitalia, ventral view. 297. Same, dorsal view.
FIGURES 301–313.
301–305. Reductoonops hato, new species, female. 306–313. R. diamant, new species, female. 301, 309. Labium and endites, ventral view. 302, 310. Spinnerets, posterior view. 303, 311. Epigastric region, ventral view. 304, 312. Genitalia, ventral view. 305, 313. Same, dorsal view. 306, 307. Sternum, ventral view. 308. Abdomen, ventral view.
FIGURES 314–323.
314–318. Reductoonops marta, new species, female. 319–323. R. leticia, new species, female. 314, 319. Sternum, ventral view. 315, 320. Abdomen, ventral view. 316, 321. Epigastric region, ventral view. 317, 322. Genitalia, ventral view. 318, 323. Same, dorsal view.
FIGURES 324–338.
324, 327–338. Reductoonops sasaima, new species, male. 325. Same, female. 326. R. meta, new species, male. 324–326. Sternum, ventral view. 327. Spinnerets, posterior view. 328. Labium and endites, ventral view. 329. Tip of endite, ventral view. 330. Palp, prolateral view. 331. Same, ventral view. 332. Same, retrolateral view. 333. Palpal bulb, prolateral view. 334. Same, retrolateral view. 335. Embolus, prolateral view. 336. Same, ventral view. 337. Same, retrolateral view. 338. Same, dorsal view.
FIGURES 339–352.
339, 340. Reductoonops sasaima, new species, female. 341–352. R. meta, new species, male. 339. Genitalia, ventral view. 340. Same, dorsal view. 341. Labium and endites, ventral view. 342. Tip of endite, ventral view. 343. Palp, prolateral view. 344. Same, ventral view. 345. Same, retrolateral view. 346. Same, dorsal view. 347. Palpal bulb, prolateral view. 348. Same, retrolateral view. 349. Embolus, prolateral view. 350. Same, ventral view. 351. Same, retrolateral view. 352. Same, dorsal view.
FIGURES 353–367.
Reductoonops tandapi, new species, male. 353. Carapace, anterior view. 354. Eye group, anterior view. 355. Spinnerets, posterior view. 356. Labium and endites, ventral view. 357. Tip of endite, ventral view. 358. Palp, prolateral view. 359. Same, ventral view. 360. Same, retrolateral view. 361. Same, dorsal view. 362. Palpal bulb, prolateral view. 363. Same, retrolateral view. 364. Embolus, prolateral view. 365. Same, ventral view. 366. Same, retrolateral view. 367. Same, dorsal view.
FIGURES 368–380.
368–372. Reductoonops tandapi, new species, male (368) and female (369–372). 373–380. R. pichincha, new species, female. 368, 369, 373, 374. Sternum, ventral view. 370, 375, 377. Carapace, anterior view. 371, 379. Genitalia, ventral view. 372, 380. Same, dorsal view. 376. Labium and endites, ventral view. 378. Spinnerets, posterior view.
FIGURES 381–392.
Reductoonops tina, new species, male. 381. Spinnerets, posterior view. 382. Labium and endites, ventral view. 383. Tip of endite, ventral view. 384. Palp, prolateral view. 385. Same, ventral view. 386. Same, retrolateral view. 387. Palpal bulb, prolateral view. 388. Same, retrolateral view. 389. Embolus, prolateral view. 390. Same, ventral view. 391. Same, retrolateral view. 392. Same, dorsal view.
FIGURES 393–402.
393–397. Reductoonops tina, new species, male (393) and female (394–397). 398–402. R. domingo, new species, female. 393, 394, 398. Sternum, ventral view. 395, 399. Abdomen, ventral view. 396, 401. Genitalia, ventral view. 397, 402. Same, dorsal view. 400. Epigastric region, ventral view.
FIGURES 403–414.
Reductoonops domingo, new species, male. 403. Sternum, ventral view. 404. Spinnerets, posterior view. 405. Labium and endites, ventral view. 406. Tip of endite, ventral view. 407. Palp, prolateral view. 408. Same, ventral view. 409. Same, retrolateral view. 410. Same, dorsal view. 411. Embolus, prolateral view. 412. Same, ventral view. 413. Same, retrolateral view. 414. Same, dorsal view.
FIGURES 415–429.
Reductoonops otonga, new species, male. 415, 416. Sternum, ventral view. 417. Spinnerets, posterior view. 418. Labium and endites, ventral view. 419. Tip of endite, ventral view. 420. Palp, prolateral view. 421. Same, ventral view. 422. Same, retrolateral view. 423. Same, dorsal view. 424. Palpal bulb, prolateral view. 425. Same, retrolateral view. 426. Embolus, prolateral view. 427. Same, ventral view. 428. Same, retrolateral view. 429. Same, dorsal view.
FIGURES 430–439.
430–434. Reductoonops otonga, new species, female. 435–439. R. palenque, new species, male (435) and female (436–439). 430, 435, 436. Sternum, ventral view. 431, 437. Abdomen, ventral view. 432. Epigastric region, ventral view. 433, 438. Genitalia, ventral view. 434, 439. Same, dorsal view.
FIGURES 440–451.
Reductoonops palenque, new species, male. 440. Labium and endites, ventral view. 441. Tip of endite, ventral view. 442. Palp, prolateral view. 443. Same, ventral view. 444. Same, retrolateral view. 445. Same, dorsal view. 446. Palpal bulb, prolateral view. 447. Same, retrolateral view. 448. Embolus, prolateral view. 449. Same, ventral view. 450. Same, retrolateral view. 451. Same, dorsal view.
FIGURES 452–463.
452. Reductoonops jatun, new species, male. 453–463. R. napo, new species, male. 452, 453. Spinnerets, posterior view. 454. Labium and endites, ventral view. 455. Tip of endite, ventral view. 456. Palp, prolateral view. 457. Same, ventral view. 458. Same, dorsal view. 459. Palpal bulb, prolateral view. 460. Embolus, prolateral view. 461. Same, ventral view. 462. Same, retrolateral view. 463. Same, dorsal view.
FIGURES 464–473.
464–468. Reductoonops napo, new species, male (464) and female (465–468). 469–473. R. jatun, new species, male (469) and female (470–473). 464, 465, 469, 470. Sternum, ventral view. 466, 471. Abdomen, ventral view. 467, 472. Genitalia, ventral view. 468, 473. Same, dorsal view.
FIGURES 474–485.
Reductoonops jatun, new species, male. 474. Labium and endites, ventral view. 475. Tip of endite, ventral view. 476. Palp, prolateral view. 477. Same, ventral view. 478. Same, retrolateral view. 479. Same, dorsal view. 480. Palpal bulb, prolateral view. 481. Same, retrolateral view. 482. Embolus, prolateral view. 483. Same, ventral view. 484. Same, retrolateral view. 485. Same, dorsal view.
FIGURES 486–500.
Reductoonops hedlite, new species, male. 486. Carapace, anterior view. 487. Eye group, anterior view. 488. Spinnerets, posterior view. 489. Labium and endites, ventral view. 490. Tip of endite, ventral view. 491. Palp, prolateral view. 492. Same, ventral view. 493. Same, retrolateral view. 494. Same, dorsal view. 495. Palpal bulb, prolateral view. 496. Same, retrolateral view. 497. Embolus, prolateral view. 498. Same, ventral view. 499. Same, retrolateral view. 500. Same, dorsal view.
FIGURES 501–510.
501–505. Reductoonops hedlite, new species, male (501) and female (502–505). 506–510. R. molleturo, new species, male (506) and female (507–510). 501, 502, 506, 507. Sternum, ventral view. 503, 508. Carapace, anterior view. 504, 509. Genitalia, ventral view. 505, 510. Same, dorsal view.
FIGURES 511–525.
Reductoonops molleturo, new species, male. 511. Carapace, anterior view. 512. Eye group, anterior view. 513. Spinnerets, posterior view. 514. Labium and endites, ventral view. 515. Tip of endite, ventral view. 516. Palp, prolateral view. 517. Same, ventral view. 518. Same, retrolateral view. 519. Same, dorsal view. 520. Palpal bulb, prolateral view. 521. Same, retrolateral view. 522. Embolus, prolateral view. 523. Same, ventral view. 524. Same, retrolateral view. 525. Same, dorsal view.
FIGURES 526–537.
Reductoonops celica, new species, male. 526. Spinnerets, posterior view. 527. Labium and endites, ventral view. 528. Tip of endite, ventral view. 529. Palp, prolateral view. 530. Same, ventral view. 531. Same, retrolateral view. 532. Palpal bulb, prolateral view. 533. Same, retrolateral view. 534. Embolus, prolateral view. 535. Same, ventral view. 536. Same, retrolateral view. 537. Same, dorsal view.
FIGURES 538–547.
538–542. Reductoonops celica, new species, male (538) and female (539–542). 543–547. R. pinta, new species, male (543) and female (544–547). 538, 539, 543, 544. Sternum, ventral view. 540, 545. Abdomen, ventral view. 541, 546. Genitalia, ventral view. 542, 547. Same, dorsal view.
FIGURES 548–559.
Reductoonops pinta, new species, male. 548. Spinnerets, posterior view. 549. Labium and endites, ventral view. 550. Tip of endite, ventral view. 551. Palp, prolateral view. 552. Same, ventral view. 553. Same, retrolateral view. 554. Palpal bulb, prolateral view. 555. Same, retrolateral view. 556. Embolus, prolateral view. 557. Same, ventral view. 558. Same, retrolateral view. 559. Same, dorsal view.
FIGURES 560–571.
Reductoonops carpish, new species, male. 560. Spinnerets, posterior view. 561. Labium and endites, ventral view. 562. Tip of endite, ventral view. 563. Palp, prolateral view. 564. Same, ventral view. 565. Same, retrolateral view. 566. Palpal bulb, prolateral view. 567. Same, retrolateral view. 568. Embolus, prolateral view. 569. Same, ventral view. 570. Same, retrolateral view. 571. Same, dorsal view.
FIGURES 572–582.
572–577. Reductoonops carpish, new species, male (572) and female (573–577). 578–582. R. elqui, new species, female. 572, 573, 578. Sternum, ventral view. 574, 579. Spinnerets, posterior view. 575, 580. Labium and endites, ventral view. 576, 581. Genitalia, ventral view. 577, 582. Same, dorsal view.
ACKNOWLEDGMENTS
This study is part of the oonopid PBI project supported by the U.S. National Science Foundation (grant DEB-0613754) and organizations in several other countries. The assistance of the many participants in that project is immensely appreciated. As always, we thank the many curators of collections that have supplied specimens: Jonathan Coddington (USNM), Gonzalo Giribet and Laura Leibensperger (MCZ), Charles Griswold and Darrell Ubick (CAS, CDU), Clifford Keil and Mauricio Vega (QCAZ), Martín Ramírez (MACN), James Reddell (TMM), Christine Rollard (MNHN), Petra Sierwald (FMNH), and Carlos Víquez (INBIO). We also thank Steve Thurston for composing the plates, and Cristian Grismado and an anonymous reviewer for helpful comments on a draft of the manuscript.
