Opopaea Simon, 1891

Daniela, Andriamalala, D. C., Hormiga, Gustavo & D. C., 2013, Systematics Of The Goblin Spider Genus Opopaea (Araneae, Oonopidae) In Madagascar, Bulletin of the American Museum of Natural History 2013 (380), pp. 1-156 : 17-34

Opopaea Simon, 1891

Opopaea Simon, 1891: 560 (type species by monotypy Opopaea deserticola Simon ). DIAGNOSIS: Members of the genus Opo-

depression; Saaristo, 2001; Saaristo and Marusik, 2008) (figs. 4, 47, 48, 807, 808); and lack of an elongated distal tooth on the claws of leg IV (figs. 74, 75, 76).

(3) Both sexes have: lateral striations on the carapace (figs. 9, 10, 12, 13); smooth sternum surface (figs. 11, 14, 15, 16); abdomen completely covered dorsally and ventrally with scuta; long and thin lateral apodemes that extend past the posterior spiracles (figs. 457, 460); pedicel tube with a pair of small dorsolateral, triangular extensions; and a pair of curved scutal ridges on the scutopedicel region (figs. 42, 45, 146, 211).

DESCRIPTION: Male: Total length: 1.02– 2.14. Cephalothorax: Carapace without any pattern, posterolateral edge without pits, posterior margin not bulging below posterior rim, anterolateral corners without extension or projections, posterolateral surface without spikes, thorax without depressions, fovea absent, without radiating rows of pits; lateral margin straight; nonmarginal pars thoracica setae needlelike. Clypeus: median projection absent. Eyes: six, well developed. Sternum longer than wide, uniform, median concavity absent, smooth with radial furrows between coxae I–II, II–III, III–IV, anterior margin unmodified, posterior margin not extending posteriorly of coxae IV, anterior corner unmodified, extensions of precoxal triangles absent, lateral margins between coxae III unmodified. Mouthparts: chelicerae anterior face unmodified; without teeth on both promargin and retromargin; fangs without toothlike projections, shape normal, without prominent basal process, tip unmodified; setae needlelike; paturon distal region unmodified, posterior surface unmodified, promargin unmodified, inner margin unmodified. Labium sclerotization as in sternum. Endites distally not excavated, serrula present in a single row, posteromedian part unmodified, same as sternum in sclerotization. Abdomen: Without long posterior extension, rounded posteriorly, interscutal membrane rows of small sclerotized platelets absent posteriorly; dorsum soft portions without color pattern. Booklung covers without setae. Posterior spiracles connected by groove. Pedicel tube short with a pair of dorsolateral, triangular extensions, scutum not extending far dorsal of pedicel, matted setae on anterior ventral abdomen in pedicel

137. Epigynum, dorsal view. Scale bars: 0.1 mm.

area absent, cuticular outgrowths near pedicel absent. Epigastric scutum strongly sclerotized, not protruding, small lateral sclerite absent. Postepigastric scutum strongly sclerotized, covering nearly full length of abdomen, anterior margin unmodified. Spinneret scutum present, incomplete ring, with fringe of needlelike setae, supraanal scutum absent, dense patch of setae anterior to spinnerets absent. Anterior lateral spinnerets (ALS) with single major ampullate gland spigot and one to two piriform gland spigots, posterior medians (PMS) with one to two minor ampullate gland spigots, posterior laterals (PLS) with one minor ampullate and two to three aciniform gland spigots. Epigastric area setae uniform, needlelike. Legs: Without spines, femur IV not thickened, same size as femora I–III, patella plus tibia I shorter than carapace, tibia I unmodified, tibia I Emerit’s glands absent, tibia IV ventral scopula absent, metatarsi I and II mesoapical comb absent, metatarsi III and IV weak ventral scopula absent, tarsal proclaws and retroclaws inner face striate; tarsus I superior claws with one tooth on lateral surface of proclaw, five teeth on median surface of proclaw, one tooth on lateral surface of retroclaw, five teeth on median surface of retroclaw; tarsus II superior claws with one tooth on lateral surface of proclaw, five teeth on median surface of proclaw, one tooth on lateral surface of retroclaw, five teeth on median surface of retroclaw; tarsus III superior claws with one tooth on lateral surface of proclaw, four teeth on median surface of proclaw, one tooth on lateral surface of retroclaw, four teeth on median surface of retroclaw; tarsus IV inner claws with four large prolateral teeth and six small distal teeth, outer claws with four large teeth (figs. 75, 76). Tibia with two to three trichobothria, metatarsi with one trichobothria (figs. 71, 73). Trichobothria base round- ed, aperture internal texture gratelike, hood smooth (figs. 89, 90, 91, 92). Tarsal organ semiexposed and with longitudinal ridges, with four raised receptors on legs I–II and two on legs III–IV (figs. 81–84). Genitalia: Epigastric region with sperm pore visible; furrow without setae. Palp of normal size, right and left palps symmetrical; embolus light, prolateral excavation absent; trochanter of normal size, unmodified; femur not enlarged, of normal size, without posteriorly rounded lateral dilation; patella without prolateral row of ridges, setae unmodified; cymbium narrow in dorsal view, completely fused with bulb, seam visible only in some undescribed Opopaea species from Australia (B. Baehr, personal commun.), not extending beyond distal tip of bulb, plumose setae absent, without stout setae, without distal patch of setae; bulb 1 to 1.5 times as long as cymbium, slender, elongated.

Female: Total length: 1.25–2.43. Cephalothorax: Carapace with angular posterolateral corners. Plumose setae near posterior margin of pars thoracica absent. Clypeus: setae present, needlelike. Sternum fused to carapace, radial furrow opposite coxae III absent, surface smooth, without pits, without lumps. Female palp: claws absent; spines absent; tarsus unmodified, patella without prolateral row of ridges. Abdomen: Book lung covers anterolateral edge unmodified. Dorsal scutum strongly sclerotized, middle surface punctate, sides punctate, anterior half without projecting denticles. Epigastric scutum without lateral joints. Legs: Leg spines absent. Tarsus I superior claws with five teeth on lateral surface of proclaw, nine or more teeth on median surface of proclaw, five teeth on lateral surface of retroclaw, nine or more teeth on median surface of retroclaw; tarsus II superior claws with five teeth on lateral surface of proclaw, nine or more teeth on median surface of proclaw, five teeth on lateral surface of retroclaw, nine or more teeth on median surface of retroclaw; tarsus III superior claws with five teeth on lateral surface of proclaw, five teeth on lateral surface of retroclaw; tarsus IV inner claws with four large prolateral teeth and six small distal teeth, outer claws with four large teeth (figs. 75, 76). Trichobothria: tibia, each with three; metatarsus, each with one. The tracheal system of all 26 Malagasy Opopaea consists of two large trunks arising just below the epigastric furrow from each of the two tracheal spiracles. Each tracheal trunk goes anteriorly through the pedicel into the prosoma and posteriorly each trunk branches into numerous tracheoles that spread in the

abdomen (figs. 63–67). Epigynum internally with a T-shaped anterior sclerites (Asc) which ends posteriorly with a small median projection (Na, sensu Burger, 2009). The receptaculum is connected anteriorly to a short and straight tube (Gap) and bear at its anterior surface different numbers (three to four) and pattern of small holes that seems to have glands enclosed inside of them. Receptaculum inverted-arc shaped, of different size, always shorter than lateral apodemes and with a slit posteriorly (figs. 51, 60).

PHYLOGENETICS: The monophyly of Opopaea has not been tested yet in a phylogenetic framework. Putative synapomorphies for the genus could be the ‘‘postgynum depression’’ (figs. 47–50) and a series of holes on the anterior part of the receptaculum (fig. 54).

COMPOSITION: There are currently 50 described species of Opopaea worldwide. Only the 27 species known from Madagascar are treated here: Opopaea andranomay , n. sp., O. ankarafantsika , n. sp., O. ankarana , n. sp., O. antsalova , n. sp., O. andringitra , n. sp., O. antsiranana , n. sp., O. bemarivo , n. sp., O. bemaraha , n. sp., O. berenty , n. sp., O. betioky , n. sp., O. concolor (Blackwall, 1859) , O. itampolo , n. sp., O. kirindy , n. sp., O. manderano , n. sp., O. mahafaly , n. sp., O. manongarivo , n. sp., O. maroantsetra , n. sp., O. namoroka , n. sp., O. sandranantitra , n. sp., O. torotorofotsy , n. sp., O. tsimaloto , n. sp., O. tsimbazaza , n. sp., O. tsimembo , n. sp., O. tsinjoriaky , n. sp., O. tsingy , n. sp., O. vohibazaha , n. sp., and O. foulpointe , n. sp.

NATURAL HISTORY AND DISTRIBUTION: In Madagascar, Opopaea species were found all over the island, in different types of habitat, such as montane rainforest, tropical dry forest, spiny/thicket forest, gallery forests, tropical deciduous forest, secondary forests near the beach, and mango and palm plantations from 10–1990 m elevations. Opopaea have also been collected in the litter, in dead wood, from trees, and even in the canopy. Given their distribution, one would think of Opopaea as a generalist, however, the results from the MAXENT model of their distribution suggests that Opopaea species are very sensitive to precipitation. Most of the predicted suitable areas for the genus are found in the eastern, humid part of the island, especially in the northeast and the southeast of Madagascar (map 6) and the MAXENT model’s internal jackknife test of variable importance showed that precipitation variables such as ‘‘Precipitation of driest month (bio14)’’ and ‘‘Precipitation of wettest quarter (bio16)’’ were the two most important predictors of Opopaea ’s habitat distribution in Madagascar ( table 2 ).

KEY TO THE OPOPAEA SPECIES OF MADAGASCAR

1. Males.......................... 2

– Females........................ 25

2. Cymbiobulbus globular (figs. 668–669, 675– 676).................. O. aroantsetra

– Cymbiobulbus elongated (all other species, figs. 156, 188, 221)................ 3

3. Cymbiobulbus without protuberance (figs. 224, 226)................. O. ankarafantsika

– Cymbiobulbus with protuberance (all other species, fig. 187–188, 252–253, 805–806).... 4

181. Dorsal view. 182. Ventral view. Scale bars: 0.2 mm.

4. Cymbiobulbus with one distal protuberance (figs. 731, 732)................... 5

– Cymbiobulbus with different pattern of protuberance (figs. 252, 474)......... 6

5. Cymbiobulbus tapering anteriorly with dorsal margin curved in an arc shape. Cymbiobulbus tip with sclerotized truncated leaflike extensions (figs. 966, 967)... O. vohibazaha

– Cymbiobulbus not tapering anteriorly but with a truncated edge. Cymbiobulbus tip without extensions (figs. 731, 732, 735, 736)................ O. sandranantitra

6. Cymbiobulbus with one posterior protuberance (fig. 766).................... 7

– Cymbiobulbus otherwise (fig. 474)..... 10

7. Cymbiobulbus tip with a long, fat, leafshaped sclerotized extension directed to the prolateral side.......... O. torotorofots y

– Cymbiobulbus tip without a long, fat, leafshaped sclerotized extension directed to the prolateral side.................... 8

8. Cymbiobulbus with the area before fenestra (abf) very thick and sclerotized all the way toward the embolar region. Palp tip short, beaklike, and sclerotized (figs. 864, 865, 868, 869)................... O. tsimembo

– Cymbiobulbus with the area before fenestra (abf) neither thick nor sclerotized...... 9

9. Tip of the cymbiobulbus on prolateral side with a very thin and compact extension directed downward and bearing above it a round hole............. O. tsimbazaza

– Tip of the cymbiobulbus on prolateral side without a thin and compact extension direct- ed downward and not bearing above it a round hole............. O. antsiranana

10. Cymbiobulbus with one anterior and one posterior protuberance (fig. 475)....... 11

– Cymbiobulbus with more than two posterior protuberances (fig. 156)............. 24

11. Sternum posterior part with protrusions or lumps (figs. 245, 247)............... 12

– Sternum posterior part without protrusion or lumps (fig. 142)................... 16

12. Sternum posterior part constricted then enlarged posteriorly, with two lateral Lshaped longitudinal lumps. Cymbiobulbus resembling a parrot beak. Palpal femur inserted at the end of patella (figs. 466, 469, 474, 475)............... O. foulpointe

– Sternum posterior part not constricted nor enlarged posteriorly, without lateral L-shaped longitudinal lumps. Cymbiobulbus not resembling a parrot beak. Palpal femur not inserted at the end of patella......... 13

13. Posterior part of sternum with one longitudinal median protrusion. Carapace with two lateral denticles forming two sharp, spikelike lateral denticules. Sternum posterior part more or less rectangular (figs. 245, 247)...... 14

– Sternum posterior part with a U-shaped median protrusion (fig. 525).......... 15

14. Tip of the cymbiobulbus large, round, pointing downward (fig. 249)...... O. ankarana

– Tip of cymbiobulbus also rounded, but bearing a more or less long sclerotized embolic extension that folds laterally on the retrolateral side......... O. manongarivo

15. Dark-brown coloration. Carapace pars cephalica strongly elevated medially with numerous rows of thick and stiff hairs. Sternum surface punctuate. Tip of cymbiobulbus tubular, sclerotized, and curved downward (figs. 513, 523–525, 531)....... O. kirindy

– Orange coloration. Carapace pars cephalica not elevated medially without rows of thick and stiff hairs. Sternum surface smooth. Tip of cymbiobulbus not tubular, but thick and sclerotized (figs. 276–278, 283–284).......................... O. antsalova

16. Distal part of cymbiobulbus truncated and rectangular (figs. 187–188)... O. andringitra

– Distal part of cymbiobulbus neither truncat- ed nor rectangular................. 17

17. Opercula small (figs. 498, 694)........ 18

– Opercula medium to large (figs. 374, 430)... 22

18. Paired scutal ridges not joined or consisting of two separate lobes (figs. 694, 796).... 19

– Paired scutal ridges medially joined or consisting of one large lobe (figs. 498, 923)..... 20

19. Palpal patella obviously larger and longer than cymbiobulbus and connected medially to femur (figs. 699, 700)..... O. namoroka

– Palpal patella not obviously larger nor longer than cymbiobulbus and connected subbasally to femur (figs. 801, 802)...... O. tsimaloto

20. Palpal fenestra top (FT) raised dorsally (figs. 932, 939), palp tip (pt) with two short, liplike extensions (figs. 932, 933, 941)....................... O. tsinjoriaky

– Palpal fenestra top (FT) not raised dorsally, palp tip (pt) without a short, liplike extensions (figs. 507, 508)............... 21

21. Cymbiobulbus tip (pt) terminated with a sclerotized, long, and straight extension (figs. 600, 601, 609, 611)... O. manderano

– Cymbiobulbus tip (pt) not terminated with a sclerotized, long, and straight extension (figs. 507, 508)............ O. itampolo

22. Color yellowish, carapace flat and elongated, hexagonal (figs. 375–378); abdomen elongat- ed, rectangular (fig. 370)..... O. bemarivo

– Color orange to dark orange, carapace neither flat nor hexagonal; abdomen ovoid..... 23

23. Opercula with sclerotized ridges (fig. 146), carapace strongly elevated with chelicera with bosses and detached from clypeus edge (figs. 147–149).......... O. andranomay

202. Epigynum, dorsal view. Scale bars: 0.1 mm.

– Opercula without sclerotized ridges, carapace slightly elevated with chelicera without bosses and not detached from clypeus edge (figs. 557– 560, 892–895)..................... 24

24. Eyes large (ratio of diameter of ALE to female body size more than 0.040 mm, figs. 557–560)............. O. mahafaly

– Eyes small (ratioof diameter of ALE to female body size less than 0.040 mm, figs. 557–560)............... O. tsingy

25. Postgynum depression U-shaped (figs. 385, 441, 602)....................... 26

– Postgynum depression not U-shaped (figs. 158, 190)............................. 28

26. Carapace flattened, abdomen elongated, rectangular (figs. 368, 375, 376)... O. bemarivo

– Carapace not flattened, abdomen ovoid (figs. 424, 432)................... 27

27. Postgynum depression ridge laterally sclerotized (fig. 603)........... O. manderano

– Postgynum depression ridge not laterally sclerotized (figs. 441, 442)..... O. betioky

28. Postgynum depression V-shaped (figs. 285, 509, 838)......................... 29

– Postgynum not depression V-shaped (figs. 476, 706)............................. 32

29. Paired scutal ridges medially joined.......................... O. Itampolo

– Paired scutal ridges not medially joined (figs. 234, 827)................... 30

30. Postgynum depression very narrow, V- shaped (figs. 838, 839)..... O. tsimbazaza

– Postgynum depression a very wide V-shape (figs. 285, 286)................... 31

31. Postgynum depression ridge thick, parmula above epyginum scutal ridge... O. antsalova

– Postgynum depression ridge not thick, par- mula detached from epyginum scutal ridge..................... O. concolor

32. Postgynum depression slitlike (figs. 318, 319, 476, 477)....................... 33

– Postgynum depression otherwise (fig. 772) 38

33. Abdomen apodemes extend beyond K of abdomen length.......... O. foulpointe

– Abdomen apodemes short (not reaching K of abdomen length).................. 34

34. Abdomen laterally flattened (figs. 306, 555)........................... 35

214. Dorsal view. 215. Ventral view. Scale bars: 0.2 mm.

– Abdomen not laterally flattened (fig. 657).. ......................... 36

35. Abdomen round, opercula diameter less than K of pedicel diameter (figs. 301, 307)...................... O. antsiranana

– Abdomen not round, opercula diameter K or more than K of pedicel diameter, epigynum scutal ridge dropping down medially (figs. 550, 556, 567).................. O. mahafaly

36. Parmula above epigynal scutal ridge, scutal ridge divided (figs. 737, 738)............................. O. sandranantitra

– Parmula and scutal ridge otherwise..... 37

37. Parmula within scutal ridge (fig. 968)...................... O. vohobazaha

– Parmula below scutal ridge (fig. 671)....................... O. maroantsetra

38. Postgynum depression inverted-arc shaped (figs. 773, 870, 934)................ 39

39. Postgynum depression short and narrow (figs. 157, 189, 222, 255, 352)......... 40

– Postgynum depression otherwise....... 43

40. Color dark orange, scutal ridge medially dropping into a reverse triangle (figs. 170, 189).................. O. andringitra

– Color orange, scutal ridge not medially dropping into a reverse triangle....... 41

41. Paired extensions on pedicel area small and widely separated from each other, scutal ridge straight (fig. 341).......... O. bemaraha

– Paired extensions on pedicel area large and close to each other, scutal ridges curved....... 42

42. Opercula outer edge thick and sclerotized (fig. 211)............. O. ankarafantsika

– Opercula outer edge not thick and not sclerotized (fig. 211)........ O. ankarana

43. Postgynum ridge thin and not sclerotized (figs. 409, 532, 636, 902)............ 44

– Postgynum ridge thick and sclerotized (figs. 870, 934)................... 47

44. Abdomen round and flattened (figs. 391, 396, 890)........................... 45

– Abdomen round but not flattened (figs. 515, 618)............................. 46

45. Parmula on top of a thickened arclike structure (fig. 902)............ O. tsingy

– Parmula almost invisible and not on top of a thickened arclike structure (fig. 409).......................... O. berenty

46. Body small (1.0– 1.5 mm), orange. Pedicel paired extensions well developed and close to each other, groove connecting posterior spiracles sclerotized (figs. 616, 624, 635)................... O. manomgarivo

– Body large (1.85–2 mm), dark orange. Pedicel paired extensions small and widely separated from each other, groove connecting posterior spiracles not sclerotized (figs. 513, 521, 532)..................... O. kirindy

47. Postgynum depression as wide as the distance between the lateral apodemes and almost as long as the distance between epigastric furrow and the shallow groove connecting the tracheal spiracles (figs. 772, 773).......................... O. torotorofotsy

– Postgynum depression width less than the distance between the lateral apodemes and length less than the distance between epigastric furrow and the shallow groove connecting the tracheal spiracles (figs. 808, 870, 934)...................... 48

48. Paired scutal ridges medially joined (figs. 859, 923)........................... 49

– Paired scutal ridges not medially joined (figs. 694, 796)................... 50

49. Abdomen flat, postgynum depression ridge extremely thick (figs. 921, 934)............................. O. tsinjoriaky

– Abdomen not flattened, postgynum depression ridge not extremely thick (figs. 853, 870)................... O. tsimembo

50. Paired triangular extensions on pedicel area barely visible or less than two times higher than pedicel border and widely separated from each other, postgynum depression ridge medially thickened (figs. 796, 808)........................... O. tsimaloto

– Paired triangular extensions on pedicel area two times higher than pedicel border and close to each other, postgynum depressionridge not medially thickened (figs. 694, 706)................... O. namoroka .

Tables

TABLE 2 The top four predictor variables and their average (from 10 replicates) percent contribution for Opopaea distribution in Madagascar (from MAXENT model’s internal jackknife test of variable importance)

Contribution Predictor variables (%)
Precipitation of driest month (bio14) Precipitation of wettest quarter (bio16)	24.2 17.8
Max temperature of the warmest month (bio5) Precipitation of the driest month (bio17)	12.6 7.2