Heteroonops Dalmas, 1916

Platnick, Norman I. & Dupérré, Nadine, 2009, The Goblin Spider Genus Heteroonops (Araneae, Oonopidae), With Notes on Oonops, American Museum Novitates 3649, pp. 1-72 : 17-22

Heteroonops Dalmas

Heteroonops Dalmas, 1916: 203 (type species by monotypy Oonops spinimanus Simon ).

Matyotia Saaristo, 2001: 349 (type species by original designation Matyotia tetraspinosus Saaristo ). NEW SYNONYMY.

DIAGNOSIS: Males can be recognized by the backward-pointing projection situated anteriorly on the palpal endites (figs. 272, 273); some males have a second projection situated posteriorly of the first (figs. 154, 155), and that second projection can sometimes be enlarged (figs. 85, 86). Females can be recognized by their elongated, highly spinose pedipalps, which have spines on the femora, patellae, tibiae, and tarsi (figs. 108, 109); the palpal patellae are sometimes laterally dilated (fig. 112) or elongated (fig. 320) as well. At least one other soft-bodied oonopid has a large posterior projection on the male endites ( Oonops tolucanus Gertsch and Davis, 1942 , from Mexico), but in that species the anterior projection appears to be lacking, the legs are spineless, and the female palp (assuming the sexes were correctly matched by Gertsch and Davis) is normal in size and spineless.

RELATIONSHIPS: At this point, existing knowledge of the diversity of soft-bodied oonopids is far too primitive to allow thorough testing of hypotheses about the relationships of Heteroonops , but there are at least two groups of species that may (singly or together) contain or constitute its sister group. Both groups include species from the Lesser Antilles but probably also occur in Panama and/or northern South America; unfortunately each group is currently represented by very few specimens.

One group of species occurs in Puerto Rico and through the Lesser Antilles at least to Trinidad. The male palps have a spiniform conductor, the male endites have an anterior projection that is anteriorly or ventrally (not posteriorly) directed, and females have the posterior receptacula much more extensively developed than in Heteroonops . Among the described species that may belong to this group are Oonops viridans Bryant (1942) and Oonops ebenecus Chickering (1972) from Puerto Rico, Oonops trapellus Chickering (1970) from Trinidad, and Oonops donaldi Chickering (1951) from Panama.

In the second group of species, the projections on the male endites are more hook shaped and are accompanied posteriorly by an excavation, and females again have more extensively developed posterior genitalic elements. In this group, however, the male palp lacks a conductor and the embolus is widened. The described species that may belong to this group include Oonops ronoxus Chickering (1971) from the Virgin Islands, Oonops aristelus Chickering (1972) from Antigua, and Oonops sicorius Chickering (1970) from Curaçao.

At least until the relationships of these Caribbean taxa, those currently assigned to Oonopoides Bryant (1940) , and their potential relatives in northern South America can be clarified, it seems best to restrict Heteroonops to the set of species that seem, on the basis of their posteriorly directed male endite projections and their elongated, spinose female pedipalps, to be the closest relatives of H. spinimanus .

DESCRIPTION: Total length of males 1.3– 1.8, of females 1.4–2.0. Cephalothorax: Carapace yellow, without any pattern, ovoid in dorsal view (figs. 98, 100, 269, 312), pars cephalica slightly elevated in lateral view (figs. 99, 101), except in H. castellus males, where strongly elevated (fig. 270), anteriorly narrowed to 0.49 times its maximum width or less, with rounded posterolateral corners, posterolateral edge without pits, posterior margin not bulging below posterior rim, anterolateral corners without extension or projections, posterolateral surface without spikes, surface of elevated portion of pars cephalica smooth (but at least in females of H. spinimanus with rows of platelets, figs. 100, 102), sides smooth, thorax without depressions, fovea absent, without radiating rows of pits, lateral margin straight, smooth, without denticles; plumose setae near posterior margin of pars thoracica absent; nonmarginal pars cephalica and pars thoracica setae dark, plumose; marginal setae dark, plumose. Clypeus margin unmodified, curved downward in front view, vertical in lateral view, high, ALE separated from edge of carapace by their radius or more, median projection absent; setae present, dark, plumose. Chilum absent. Eyes six, well developed, ALE largest, ALE oval, PME squared, PLE oval; posterior eye row recurved from above, procurved from front; ALE separated by their radius to diameter, ALE-PLE separated by less than ALE radius, PME touching throughout most of their length, PLE-PME separated by less than PME radius. Sternum longer than wide (figs. 103, 271, 313), yellowish white, uniform, not fused to carapace, median concavity absent, with radial furrows between coxae I– II, II–III, III–IV, furrow smooth, radial furrow opposite coxae III absent, surface smooth, without pits, microsculpture absent, sickle-shaped structures absent, anterior margin unmodified, posterior margin not extending posteriorly of coxae IV, anterior corner unmodified, lateral margin without infracoxal grooves, distance between coxae II and III greater than distance between coxae I and II, and coxae III and IV, extensions of precoxal triangles present, lateral margins unmodified, without posterior hump; setae abundant, dark, plumose, evenly scattered, originating from surface, without hair tufts. Chelicerae, endites, and labium yellowish white; chelicerae straight, anterior face unmodified, without teeth on promargin or retromargin, without toothlike projections, directed medially, shape normal, without prominent basal process, tip unmodified, median half of posterior surface sometimes with low denticles; setae dark, needlelike, evenly scattered; paturon inner margin with scattered setae, distal region unmodified, posterior surface unmodified, promargin unmodified, inner margin unmodified, laminate groove absent (figs. 106, 107, 275, 276, 314, 315). Labium triangular, not fused to sternum, anterior margin indented at middle, same as sternum in sclerotization, with six or more setae on anterior margin, subdistal portion with unmodified setae (figs. 104, 316). Labrum with flattened median lobe (figs. 105, 274, 317). Endites distally not excavated, same as sternum in sclerotization, anteromedian part with backward-pointing projection in males (figs. 272, 273), posteromedian part sometimes with second backward-pointing projection as well; endites scanned only in H. castellus , female H. spinimanus , and male H. vega , females usually with serrula consisting of single row of teeth (figs. 105, 318, 319), but serrula lost in males of at least H. castellus and H. vega (and possibly other males, except H. murphyorum , fig. 154). Female palp without claw (figs. 113, 323, 325); strong spines present on femora, patellae, tibiae, and tarsi (figs. 108, 109, 111, 120, 320–322); femur with retrolateral row of platelets; patella without prolateral row of ridges, often elongated or prolaterally dilated (fig. 112), sometimes with prolateral platelets (fig. 326); tibia with dorsal row of platelets (at least in H. spinimanus , fig. 110) and three trichobothria; tarsus unmodified; tarsal organ with two sensillae (figs. 121, 324). Abdomen: Ovoid, without long posterior extension, rounded posteriorly, without rows of small sclerotized platelets; dorsum white, without color pattern; book lung covers large, ovoid, without setae, anterolateral edge unmodified; posterior spiracles connected by groove (fig. 348), leading to large tracheae penetrating cephalothorax, with numerous abdominal tracheoles (fig. 132), anterior spiracles leading to booklungs with two leaves (at least in female H. spinimanus , fig. 133); pedicel tube short, unmodified, scuto-pedicel region unmodified, abdomen extending anteriad of pedicel; plumose hairs absent, matted setae on anterior ventral abdomen in pedicel area absent, cuticular outgrowths near pedicel absent; dorsum and venter with dark, plumose setae; dense patch of setae anterior to spinnerets absent; males lacking scuta; females with weak epigastric and postepigastric scuta, epigastric scutum yellow, not surrounding pedicel, not protruding, small lateral sclerites absent, without lateral joints, postepigastric scutum yellow, short, only around epigastric furrow, not fused to epigastric scutum, anterior margin unmodified, without posteriorly directed lateral apodemes. Colulus represented only by setae; spinnerets scanned only in H. castellus and female H. spinimanus , but each with more spigots than in female Oonops pulcher (figs. 126, 278, 328): ALS with major ampullate gland spigot and two or three piriform gland spigots (figs. 127, 279, 329), PMS with 2–7 aciniform gland spigots (figs. 128, 280, 330), PLS with 2–17 aciniform gland spigots (figs. 129, 281, 331). Legs: yellow, without color pattern; femur IV not thickened, same size as femora I–III, patella plus tibia I near as long as carapace, tibia I unmodified, tibiae with Emerit’s glands in at least some species (figs. 332, 343, arrow), patellae and tibiae sometimes also with differently shaped platelets without pores (figs. 333, 342), tibia IV specialized hairs on ventral apex absent, tibia IV ventral scopula absent, metatarsi I and II mesoapical comb absent, metatarsi III and IV weak ventral scopula absent; legs with spines on femora (usually), tibiae, and metatarsi, all spines longer than segment width (fig. 344); claw and sensory structures scanned only in H. castellus and female H. spinimanus , where tarsal proclaws and retroclaws have inner face striate, males apparently have teeth only on lateral surface, females have additional distal teeth on median surface (figs. 115–119, 283– 286, 334–337); inferior claws absent; tibiae each with three trichobothria, metatarsi with one distal or subdistal trichobothrium (figs. 114, 345, 346), base longitudinally narrowed, aperture internal texture gratelike, hood covered by numerous low, closely spaced ridges (figs. 282, 327); tarsal organ of legs I, II with three sensillae, of legs III and IV with two sensillae (figs. 122–125, 287–290, 338–341, 347). Genitalia: Male epigastric region with sperm pore not visible (fig. 277); furrow without V- shaped insertions, without setae; palp of normal size (figs. 291, 292), not strongly sclerotized, right and left palps symmetrical, proximal segments yellow; embolus dark, prolateral excavation absent; trochanter of normal size, unmodified; femur of 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. 296) and (at least in H. castellus ) pore-bearing platelets (fig. 297); tarsal organ with two sensillae (fig. 298); cymbium yellow, narrow in dorsal view, not fused with bulb, not extending beyond distal tip of bulb, plumose setae absent, without stout setae, without distal patch of setae; bulb white, 1 to 1.5 times as long as cymbium, stout, elongated; short conductor present (figs. 293–295). Females with longitudinal anterior receptaculum, often widened anteriorly, forming T-shaped structure, at least sometimes with associated long secretory glands (figs. 349–353); V-shaped elements of anterior portion apparently fitting into similarly shaped posterior structures when epigastric furrow is closed, moving apart when epigastric furrow is opened (figs. 96, 130, 131); posterior receptaculum well developed, at least sometimes with pore field and associated glands.

SYNONYMY: The generic synonymy reflects our placement of the type species of Matyotia as the male of the type species of Heteroonops .

KEY TO SPECIES (except H. colombi from Cuba)

1. Females (unknown in H. iviei , H. toro , and H. macaque )....................... 2

– Males (unknown in H. singulus , H. andros , and H. spinigata ).................... 12

2. Palpal patella prolaterally dilated (as in fig. 112)........................ 3

– Palpal patella not prolaterally dilated..... 6

3. Epigastric scutum triangular, well separated from postepigastric scutum (fig. 166); anteri- or receptaculum T-shaped, with thin transverse bar (figs. 167, 168)... H. murphyorum

– Epigastric scutum rectangular, close to postepigastric scutum; anterior receptaculum otherwise....................... 4

4. Anterior receptaculum large and round, posterior receptaculum triangular (figs. 96, 97)..................... H. spinimanus

– Receptacula otherwise................ 5

5. Anterior receptaculum trumpet shaped (figs. 146, 147)............. H. singulus

– Anterior receptaculum umbrella shaped (figs. 174, 175).............. H. andros

6. Epigastric and postepigastric scuta strongly sclerotized (figs. 239, 379)............ 7

– Epigastric scutum and postepigastric scutum not strongly sclerotized............. 8

7. Epigastric scutum longer than wide (fig. 379); anterior receptaculum diamond shaped (figs. 380, 381)............... H. croix

– Epigastric scutum wider than long (fig. 239); anterior receptaculum long, gradually expanding anteriorly (figs. 240, 241) H. validus

8. Posterior receptaculum relatively small (figs. 210, 258).................... 9

– Posterior receptaculum relatively large (figs. 183, 306, 400)............... 10

9. Anterior receptaculum elongated, bearing branches (figs. 258, 259).... H. castelloides

– Anterior receptaculum T-shaped, narrow throughout its length, without branches (figs. 210, 211)............... H. vega

10. Anterior receptaculum small and rounded, posterior receptaculum heart shaped (figs. 183, 184)............ H. spinigata

– Receptacula otherwise............... 11

11. Anterior receptaculum tack shaped, posterior receptaculum rectangular (figs. 399, 400)......................... H. saba

– Anterior receptaculum terminating in knob shaped, dorsally directed expansion, posteri- or receptaculum triangular (figs. 306, 307)....................... H. castellus

12. Anteromedian part of each endite with two spines pointing backward (figs. 85, 154). 13

– Anteromedian part of each endite with one spine pointing backward............ 14

13. Anteromedian part of endites with basal spine slightly larger than apical one (fig. 154); embolus curved, only distal half darkened (figs. 156–158).......... H. murphyorum

– Anteromedian part of endites with basal spine much larger than the apical one (figs. 84, 85); embolus rather straight, proximal half darkened (figs. 87, 88)........ H. spinimanus

14. Endites with small (as wide as long), pointed spine (figs. 228, 265, 370)........... 15

– Endites with large (longer than wide), curving spine (as in figs. 191, 218).......... 17

15. Bulb constricted apically, conductor large, narrowing greatly near tip (figs. 371, 372)...................... H. croix

– Bulb not constricted apically, conductor slen- der, tapering apically.............. 16

16. Embolus curved at base, conductor short, twisted (figs. 267, 268)....... H. castellus

– Embolus strongly bent at base, conductor long, straight (figs. 230–232)........ H. validus

17. Base of embolus enlarged (as in figs. 221, 249).......................... 18

– Base of embolus not enlarged......... 19

18. Conductor narrow, spiniform (figs. 249, 250).................. H. castelloides

– Conductor large, triangular (figs. 220, 221).......................... H. iviei

19. Embolus long, conductor large, lamelliform (figs. 193–196)................ H.vega

– Embolus short, conductor narrow, spiniform.......................... 20

20. Apex of bulb rounded, not extending far beyond base of embolus (figs. 362–364)........................... H. toro

– Apex of bulb pointed, extending well beyond base of embolus................. 21

21. Apex of bulb prolonged, conductor long (figs. 390, 391)............... H. saba

– Apex of bulb not prolonged, conductor short (figs. 409, 410)............. H. macaque