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Eupnoi

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Eupnoi
Phalangium opilio Linnaeus, 1758 male by Silhavy 1956a
Phalangium opilio Linnaeus, 1758 male by Šilhavý (1956)

Order

Opiliones

Suborder

Eupnoi

Diversity

1807 spp



The Eupnoi are a suborder of harvestmen, with around 240 genera, and about 1,820 described species.

They consist of two superfamilies, the Phalangioidea with many long-legged species common to northern temperate regions, and the small group Caddoidea, which have prominent eyes and spiny pedipalps.

DistributionEdit

Caddoidea are mostly found in temperate zones of both hemispheres; however, they are known from the Palearctic only from Japan, and from Baltic amber (about 40 million years old). One species known from Japan is also found in North America, where several more species are found. Other species occur in Southern South America, Venezuela, New Zealand and Australia.

Neopilionidae show a Gondwanan distribution, with species found in South America, South Africa and Australia.

Sclerosomatidae are divided into several subfamilies, with Gagrellinae found in the Indo-Malayan and neotropical region, Gyinae at high elevations in the Caucasus, Alps and Nepal, Leiobuninae in the holarctic region down to Costa Rica, and Sclerosomatinae only in the Palearctic.

In the Phalangiidae, the Phalangiinae are most diverse in the Mediterranean, with several endemic genera in nearby coastal Africa. Although there are several Phalangiinae in the Nearctic, they are all introduced. Opilioninae are mostly palearctic, with a few species found in Southeast Asia. Oligolophinae are holarctic, with most species found in Europe. Platybuninae are also found from Europe to the Caucasus, with one find from Sumatra.[2]

RelationshipsEdit

Two different views exist at the moment: in the traditional view Eupnoi and Dyspnoi form a clade, with Laniatores as a sister group; however recently it has been proposed that Eupnoi are sister to a clade formed by Dyspnoi and Laniatores (the Dyspnolaniatores hypothesis).

SystematicsEdit

Although the two recognized superfamilies Caddoidea and Phalangioidea are probably monophyletic, the limits of families and subfamilies are uncertain in many cases, and are in urgent need of further study.

Caddidae (6 genera, 21 species)
Neopilionidae (20 genera, 64 species)
Phalangiidae (55 genera, 382 species)
Protolophidae Banks, 1893 (Nearctic – 8 species)
Sclerosomatidae (153 genera, 1337 species)

The supposed family "Stygophalangiidae", with its sole described species Stygophalangium karamani Oudemans, 1933 from former Yugoslavia is not a harvestman, but very likely a species of mite.

Key to Eupnoi families (subfamilies) [from Cokendolpher et al. 2007]Edit

1. Eyes enormous and ocularium occupies most of the prosoma (Figures 4.7a,b);
legs II shorter than legs IV.......................................................................................................................................................... Caddidae....2
.Eyes and ocularium normal; legs II longer than legs IV........................................................................................................................... 3

2. Cheliceral segment I with ventral spur; pedipalpal tarsus longer than tibia (Figure 4.7b)......................... Caddidae (Caddinae)
. Cheliceral segment I without ventral spur; pedipalpal tibia longer than tarsus (Figure 4.7a)..... Caddidae (Acropsopilioninae)

3. Spiracle of opisthosoma covered by grate of spines (Figure 4.9d) (Distribution: temperate regions of Southern Hemisphere) 4
. Spiracle of opisthosoma with entapophysis and no grill....................................................................................................................... 8

4. Penis with paired lateral bristle groups at articulation of glans and shaft (Figure 4.8d); Australia and New Zealand............
Monoscutidae....5
. Penis without paired lateral bristle groups, at most a pair of spines, at articulation of glans and shaft...... Neopilionidae....6

5. Body rounded; only carapace sclerotized; opisthosoma smooth. Legs relatively long. Chelicerae sexually dimorphic; those of male enormously enlarged (Figure 4.8g)....................................................................................................... Monoscutidae (Megalopsalidinae)
. Body dorsoventrally flattened, heavily sclerotized; dorsum ornamented. Legs short (femur I less than body width). Sexual dimorphism absent (Figure 4.8a,b)................................................................................................................................... Monoscutidae (Monoscutinae)

6. Pedipalpal patella very long (patella about twice as long as tibia); tibia reflexed on the patella so that dorsal angle is less than 180° (Figure 4.9b); Australia, Chile............................................................................................................................ Neopilionidae (Ballarrinae)
. Pedipalpal patella shorter (patella less than 15% longer than tibia); tibia not reflexed upward on patella (Figure 4.9e)....... 7

7. Pedipalpal patella and tibia with dense pile of setae; patella slightly longer than tibia; pedipalpal claw reduced and smooth (Figure 4.9a); South Africa......................................................................................................................................... Neopilionidae (Neopilioninae)
. Pedipalps without dense pile of setae; patella much shorter than tibia; claw large and toothed (Figure 4.9c); Chile, Argentina, Brazil ........................................................................................................................................................................ Neopilionidae (Enantiobuninae)

8. Male pedipalp almost always greatly thickened (Figure 4.8h); tarsus shorter than tibia; western North America Protolophidae
. Pedipalpal tarsus longer than tibia............................................................................................................................................................ 9

9. Pedipalpal claw smooth.................................................................................................................................................. Phalangiidae...10
. Pedipalpal claw toothed............................................................................................................................................................................. 13

10. Basal segment of chelicera with ventral spur (Figure 4.10f); chelicerae not strikingly different between the sexes ...............
Phalangiidae (Oligolophinae) . Basal segment of chelicera without ventral spur; chelicerae sometimes differ between the sexes (Figure 4.10h)........................ .......
Phalangiidae (Phalangiinae sensu lato) 11

11. Pedipalpal femur (often also remaining segments) thorned ventrally (Figure 4.10j); chelicerae of both sexes similar; penis rodlike, with slightly widened base .............................................................................................................................. Phalangiidae (Platybuninae)
. Pedipalp lacks ventral thorns; male chelicera tends to be modified (Figure 4.10h)........................................................................ 12

12. Pedipalpal patella and tibia of adults without distomesal apophyses; chelicerae of both sexes basically the same in structure, though often enormously expanded in male; leg I of male tends to be enormously swollen; shaft of penis dorsoventrally flattened (Figure 4.10k). . . . . . . . . . . . . . . . . Phalangiidae (Opilioninae)
. Pedipalpal patella and tibia with distomesal apophyses; male chelicera tends to be modified in morphology (especially in the distal segment); femur of male pedipalp usually with ventral rows of denticles; shaft of penis generally more rounded.......................
Phalangiidae (Phalangiinae sensu stricto)

13. Coxae II showing distally blunt lobes........................................................................................................ Sclerosomatidae (Gyinae)
. Coxae II not as above................................................................................................................................................................................... 14

14. Leg femora with noduli (at least one on femur II, Figure 4.11p); dorsum sclerotized............... Sclerosomatidae (Gagrellinae)
. Leg femora usually without noduli............................................................................................................................................................ 15

15. Dorsum heavily sclerotized, often with spines or humps (Figure 4.11c); penis tapering toward tip (Figure 4.11l)..................
Sclerosomatidae (Sclerosomatinae)

. Dorsum less sclerotized, usually smooth; penis not tapering, often with alate part (distribution: southern part of Eurasia)...
Sclerosomatidae (Leiobuninae)

ReferencesEdit

Cokendolpher, J.C., Tsurusaki, N., Tourinho, A.L., Taylor, C.K., Gruber, J. & Pinto-da-Rocha, R. (2007) Suborder Eupnoi. In: R.Pinto-da-Rocha, G. Machado & G.Giribet (eds). The Harvestmen: The Biology of Opiliones. 597 pp. Harvard University Press, Cambridge, pp. 108–131.


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