Phalangiidae

Lateral view of Odontobunus armatus, from Roewer (1923).


Belongs within: Phalangioidea.
Contains: Platybunus, Opilio, Egaenus, Homolophus, Odiellus, Mitopus, Lacinius, Phalangium, Rhampsinitus, Dasylobus, Zacheus, Metaphalangium, Cristina, Rilaena.

The Phalangiidae is a diverse group of long-legged harvestmen found in Eurasia, North America and Africa (with some species introduced to Australasia). Members of this group are mostly leathery-bodied, with spiny legs and a characteristic penis morphology with the glans bent dorsally from the shaft (Tsurusaki 2007). Members of the subfamily Gyinae, recently placed in Phalangiidae by molecular phylogenetic analysis (Hedin et al. 2012), possess well-developed lateral wings on the penis close behind the shaft-glans junction.

Characters (from Tsurusaki 2007): Body generally soft and leathery, rarely sclerotised; coloration often forming medial 'saddle' marking. Ocularium relatively high, usually covered dorsally with several tubercles. Ozopores situated near bases of legs I, visible from above. Leg coxae without marginal rows of denticles, with only sparse setae; genital operculum and opisthosomal sternites also with only sparse setae; lateral opisthosomal sclerites absent. Pedipalps with claw usually smooth, rarely slightly toothed. Legs cylindrical, often pentagonal or hexagonal in cross section, with rows of spines or setae on edges; femora without nodules; tibiae with two accessory spiracles, one near proximal and one near apical end. Penis with glans movable, articulated, bent dorsally in resting position (usually at acute angle at base); stylus well delimited and movable. Ovipositor long and segmented with bifurcated tip.

<==Phalangiidae [Phalangidae, Phalangita]
    |--Opilioninae GV09
    |    |--Opilio S03
    |    |--Egaenus ZZ12
    |    |--Homolophus T07
    |    |--Himalphalangium Martens 1973 ZZ12
    |    |    |--*H. palpale (Roewer 1956) C92 [=Opilio palpalis S03]
    |    |    |--H. dolpoense Martens 1973 S03
    |    |    |--H. nepalense (Suzuki 1972) S03
    |    |    |--H. spinulatum (Roewer 1911) (see below for synonymy) S03
    |    |    |--H. suzukii Martens 1973 S03
    |    |    `--H. unistriatum Martens 1973 S03
    |    `--Scleropilio Roewer 1911 S03 [incl. Scutopilio Roewer 1956 TTL00b, Udezatus Nakatsudi 1943 TTL00b]
    |         |--*S. coriaceus Roewer 1911 TTL00b, S03
    |         |--S. armatus (Roewer 1911) (see below for synonymy) CL02
    |         |--S. insolens (Simon 1895) S03 (see below for synonymy)
    |         `--S. tibialis (Roewer 1956) CL02 [=*Scutopilio tibialis TTL00b]
    `--+--Oligolophinae [Oligolophini] GV09
       |    |  i. s.: ‘Phalangium’ bilineatum Fabricius 1779 (n. d.) R23
       |    |         ‘Phalangium’ conigerum Sørensen 1912 (n. d.) R23
       |    |         ‘Phalangium’ crassum Dufour 1831 (n. d.) R23
       |    |         ‘Phalangium’ cristatum Olivier 1791 (n. d.) R23
       |    |         ‘Phalangium’ lineola Dufour 1831 (n. d.) R23
       |    |         ‘Phalangium’ lupatum Eichwald 1830 (n. d.) R23
       |    |         ‘Phalangium’ mucronatum Müller 1776 (n. d.) R23
       |    |         ‘Phalangium’ muscorum Latreille 1798 (n. d.) R23, L98
       |    |         ‘Phalangium’ rubens Hermann 1804 (n. d.) R23 [=P. rubicundum Hermann 1804 G00]
       |    |         ‘Phalangium’ tricuspidatum Dufour 1831 (n. d.) R23
       |    |         ‘Acantholophus’ aculeatus Koch 1839 (n. d.) R23
       |    |         ‘Opilio’ acanthopus Rossi 1847 (n. d.) R23
       |    |         ‘Opilio’ scabripes Walker 1860 (n. d.) R23
       |    |--Odiellus T07
       |    |--Mitopus ZZ12
       |    |--Lacinius S84
       |    |--Mitopiella Banks 1930 C92
       |    |    `--*M. cinctipes Banks 1930 C92
       |    |--Paralacinius Morin 1934 C92
       |    |    `--*P. podoliensis Morin 1934 C92
       |    |--Roeweritta Šilhavý 1965 C92
       |    |    `--*R. carpentieri (Roewer 1953) [=Lacinius carpentieri] C92
       |    |--Paroligolophus Lohmander 1945 C92
       |    |    |--*P. agrestis (Meade 1855) C92 (see below for synonymy)
       |    |    `--‘Opilio’ laevis Roewer 1956 S03
       |    `--Oligolophus Koch 1872 ZZ12
       |         |--*O. tridens (Koch 1836) C92 (see below for synonymy)
       |         |--O. aspersus Karsch 1881 ZZ12 (see below for synonymy)
       |         |--O. hanseni (Kraepelin 1896) ZZ12 (see below for synonymy)
       |         |--O. mollis Koch 1871 (n. d.) R23
       |         `--O. tienmushanensis Wang 1941 ZZ12
       `--+--Gyinae [Gyantinae, Gyidae] HT12
          |    |--Gyoides Martens 1982 C92
          |    |    `--*G. maximus Martens 1982 C92
          |    |--Rongsharia Roewer 1957 C92
          |    |    `--*R. singularis Roewer 1957 C92
          |    |--Amilenus Martens 1969 M78, C92
          |    |    `--*A. aurantiacus (Simon 1881) C92 (see below for synonymy)
          |    `--Gyas Simon 1879 C92
          |         |--*G. annulatus (Olivier 1792) C92 (see below for synonymy)
          |         `--G. titanus Simon 1879 R23
          `--Phalangiinae [Dentizacheinae, Leptobuninae, Leptobunini, Phalangiini] GV09
               |--Phalangium T07
               |--Rhampsinitus S84
               |--Dasylobus D04
               |--Zacheus S84
               |--Metaphalangium S84
               |--Cristina S84
               |--Rilaena S84
               |--Camerobunus Staręga & Snegovaya 2008 DM10
               |    `--*C. okucola Staręga & Snegovaya 2008 SS08
               |--Bactrophalangium Šilhavý 1966 C92
               |    `--*B. jakesi Šilhavý 1966 C92
               |--Coptophalangium Staręga 1984 S84
               |    `--*C. bunigerum (Roewer 1956) S84 [=Opilio buniger S03]
               |--Liropilio Gritsenko 1979 C92
               |    `--*L. przhevalskii Gritsenko 1979 C92
               |--Parascleropilio Rambla 1975 S84
               |    `--*P. fernandezi Rambla 1975 S84
               |--Tchapinius Roewer 1929 [=Tschapinius (l. c.)] C92
               |    `--*T. malaisei Roewer 1929 C92
               |--Ramblinus Staręga 1984 S84
               |    `--*R. spinipalpis (Roewer 1911) [=Dasylobus spinipalpis] S84
               |--Liopilio Schenkel 1951 C92
               |    |--*L. glaber Schenkel 1951 C92
               |    `--L. yukon Cokendolpher 1981 BH09
               |--Toragrella Roewer 1955 C92
               |    |--*T. longipes Roewer 1955 C92
               |    `--T. normalis (Banks 1930) [=Hologagrella normalis] R55
               |--Bunochelis Roewer 1923 S84 [=Bunostomum Roewer 1911 non Railliet 1902 C92]
               |    |--B. spinifera (Lucas 1839) S84 (see below for synonymy)
               |    `--B. canariana (Strand 1911) S03 (see below for synonymy)
               |--Platybunoides Šilhavý 1955 ZZ12
               |    |--*P. argaea Šilhavý 1955 ZZ12 [=P. argaeus C92]
               |    `--P. songi Zhang & Zhang 2012 ZZ12
               |--Hindreus Kauri 1985 C92
               |    |--*H. leleupi (Roewer 1961) C92 [=Cheops leleupi C92, Odontobunus leleupi S84]
               |    |--H. crucifer Kauri 1985 K85
               |    `--H. elegans Kauri 1985 K85
               |--Megistobunus Hansen 1921 S84
               |    |--*M. longipes Hansen 1921 [incl. M. minutus Roewer 1956] S84
               |    |--M. funereus Lawrence 1962 S84
               |    `--M. lamottei (Roewer 1959) [=Dasylobus lamottei] S84
               |--Dacnopilio Roewer 1911 S84
               |    |--*D. armatus Roewer 1911 S84 (see below for synonymy)
               |    |--D. insularis Hansen 1921 S84 S84
               |    |--D. kraepelini (Roewer 1911) [=Egaenus kraepelini] S84
               |    `--D. scopulatus Lawrence 1962 S84
               |--Guruia Loman 1902 S84
               |    |--G. africana (Karsch 1878) (see below for synonymy) S84
               |    |--G. longipes Roewer 1911 S84
               |    |--G. quadrispina Roewer 1911 S84
               |    |--G. talboti Roewer 1911 S84
               |    `--G. ultima Caporiacco 1949 S84
               |--Leptobunus Banks 1893 [=Leptobrunus (l. c.); incl. Liomitopus Schenkel 1951] C84
               |    |--*L. californicus Banks 1893 C84 (see below for synonymy)
               |    |--L. aureus Cokendolpher 1984 C84
               |    |--L. borealis Banks 1899 C84
               |    |--L. pallidus Cokendolpher 1984 C84
               |    `--L. parvulus (Banks 1894) [=Liobunum parvulum, Leuronychus parvulus] C84
               |--Metadasylobus Roewer 1911 S84 [incl. Dentizacheus Rambla 1956 C92]
               |    |--*M. fuscoannulatus (Simon 1883) C92 (see below for synonymy)
               |    |--M. echinifrons (Simon 1879) D04 (see below for synonymy)
               |    `--‘Dentizacheus’ ibericus Rambla 1967 S73
               |--Graecophalangium Roewer 1923 C92
               |    |--*G. atticum Roewer 1923 C92
               |    |--G. cretaeum Martens 1966 M15
               |    |--G. drenskii Mitov 1995 M15
               |    |--G. karakalensis Tcheremis & Snegovaya 2010 M15
               |    |--G. marenzelleri (Nosek 1905) [=Egaenus marenzelleri] M15
               |    |--G. militare (Koch 1839) [=Opilio militaris, Phalangium militare] R23
               |    |--G. punicum Staręga 1973 M15
               |    `--G. turcicum Mitov 2009 K14
               |--Mizozatus Nakatsudi 1937 N37
               |    `--*M. flavidus Nakatsudi 1937 N37
               `--Odontobunus Roewer 1910 S84 (see below for synonymy)
                    |--*O. africanus Roewer 1910 [incl. Chelibunus furciger Roewer 1956, C. ruwenzorius Roewer 1956] S84
                    |--O. armatus (Sørensen 1910) S84 (see below for synonymy)
                    |--O. elegans (Roewer 1956) [=Megistobunus elegans] S84
                    |--O. kenianus Roewer 1957 S84
                    |--O. longipes (Lawrence 1962) [=Cheops longipes] S84
                    |--O. niger (Roewer 1956) S84 [=Ruandella nigra S84; incl. *R. coronata Roewer 1956 C92]
                    |--O. punctatus (Roewer 1956) (see below for synonymy) S84
                    `--O. pupillaris (Lawrence 1962) [=Chelibunus pupillaris] S84

Phalangiidae incertae sedis:
  Cheiromachus Menge in Koch & Berendt 1854 S02, C92
    `--*C. coriaceus Menge in Koch & Berendt 1854 C92
  Platybuninae ZZ12
    |--Platybunus ZZ12
    |--Lophopilio Hadži 1931 ZZ12 [incl. Emonia Hadži 1973 C92]
    |    `--L. palpinalis (Herbst 1799) N05 (see below for synonymy)
    |--Bolea Hadži 1973 ZZ12
    |    `--*B. ephippiata Hadži 1973 C92
    |--Stankiella Hadži 1973 ZZ12
    |    `--*S. montana Hadži 1973 C92
    |--Acanthomegabunus Tsurusaki, Tchemeris & Logunov 2000 ZZ12
    |    `--*A. sibiricus Tsurusaki, Tchemeris & Logunov 2000 TTL00a
    |--Buresilia Šilhavý 1966 ZZ12, K14
    |    |--*B. nigerrimus (Roewer 1956) C92, K14 [=Metadasylobus nigerrimus C92]
    |    `--B. macirna (Roewer 1956) K14
    |--Rafalskia Staręga 1963 ZZ12
    |    |--R. olympica (Kulczyński 1903) C92 (see below for synonymy)
    |    |    |--R. o. olympica K14
    |    |    `--*R. o. bulgarica (Staręga 1963) K14 [=Paropilio (*Rafalskia) bulgaricus C92]
    |    `--R. cretica (Roewer 1923) K14
    |--Megabunus Meade 1855 ZZ12
    |    |--M. diadema (Fabricius 1779) DM10 (see below for synonymy)
    |    |--M. armatus (Kulczyński 1887) [=Platybunus armatus] R23
    |    |--M. lesserti Schenkel 1927 [incl. Platybunus femoralis Roewer 1956, Metaplatybunus femoralis] M78
    |    |--M. rhinoceros (Canestrini 1872) [=Platylophus rhinoceros] D04
    |    `--M. vignai Martens 1978 M78
    `--Metaplatybunus Roewer 1911 C92
         |--*M. grandissimus (Koch 1839) C92 (see below for synonymy)
         |--M. atroluteus Roewer 1915 R23
         |--M. carneluttii Hadži 1973 N05 [incl. M. carneluttii montenegrinus Hadži 1973 M78]
         |--M. corcyraeus Roewer 1956 R56
         |--M. creticus Roewer 1923 [=Platybunus strigosus Kulczynski 1903 non Koch 1867] R23
         |--M. filipes Roewer 1956 K14
         |--M. hypanicus Šilhavý 1966 CK05
         |--M. petrophilus Martens 1965 K14
         `--M. rhodiensis Roewer 1924 WS16
  Stephanobunus Dunlop & Mammitzsch 2010 DM10
    `--*S. mitovi Dunlop & Mammitzsch 2010 DM10
  Dicranopalpus Doleschall 1852 C92 (see below for synonymy)
    |--*D. gasteinensis Doleschall 1852 C92 (see below for synonymy)
    |--D. bolivari (Dresco 1949) [=*Fagea bolivari; incl. *Egaenasser extraordinarius Roewer 1953] C92
    |--D. insignipalpis (Simon 1879) [=Prosalpia insignipalpis] D04
    |--D. larvatus (Canestrini 1874) [=Liodes larvatus, Prosalpia larvata] D04
    |--D. martini (Simon 1878) [=Prosalpia martini] R23
    |--D. pyrenaeus Dresco 1948 D04
    |--D. ramiger (Koch & Berendt 1854) S02 (see below for synonymy)
    `--D. ramosus (Simon 1909) D04 [=*Dichranochirus ramosus C92; incl. Dicr. caudatus Dresco 1948 D04]

Nomen nudum: Oligolophus obsoletus Anon. in Roewer 1923 [=Opilio obsoletus] R23

*Amilenus aurantiacus (Simon 1881) C92 [=Liobunum aurantiacum C92, Nelima aurantiaca N05; incl. L. signatum Kulczynski 1909 R23]

Bunochelis canariana (Strand 1911) S03 [=Phalangium canarianum S84, Metaphalangium canarianum S84, M. canariarum (l. c.) S84; incl. Bunochelis altenai Spoek 1959 S84, Opilio kochi Rambla 1956 S03, O. leiobuniformis Koch 1872 (n. n.) R23, Paropilio leiobuniformis Koch ex Roewer 1911 S84]

Bunochelis spinifera (Lucas 1839) S84 [=Phalangium spiniferum S84, Metaphalangium spiniferum S84, Opilio spiniferus S84, Phalangium spinipes R12; incl. Bunochelis brevipes Roewer 1923 S84, O. canariensis Koch 1872 S84, *Bunostomum crassipes Roewer 1911 C92, Bunochelis longipes Roewer 1923 S84]

*Dacnopilio armatus Roewer 1911 S84 [incl. D. armatus obscurior Roewer 1958 S84, Opilio punctatissimus Roewer 1959 S03]

Dicranopalpus Doleschall 1852 C92 [incl. Dichranochirus Simon 1909 C92, Egaenasser Roewer 1953 C92, Fagea Dresco 1949 non Buen 1940 C92, Liodes Koch 1869 non Heyden 1826 C92, Prosalpia Koch 1872 C92; Dicranopalpinae D04]

*Dicranopalpus gasteinensis Doleschall 1852 C92 [incl. Leiobunum bibrachiatum Koch 1869 D04, *Liodes bibrachiatus C92, *Prosalpia bibrachiata C92; incl. D. fraternus Szalay 1950 D04]

Dicranopalpus ramiger (Koch & Berendt 1854) S02 [=Opilio ramiger S02; incl. D. palmnickensis Roewer 1939 S02, O. corniger Menge in Koch & Berendt 1854 S02, D. corniger D04]

Guruia africana (Karsch 1878) [=Zacheus africanus, Egaenus africanus; incl. *Guruia frigescens Loman 1902, Rhampsinitus lettowi Roewer 1923, G. leucobunus Roewer 1917, G. levis Loman 1902, G. matengona Roewer 1956, G. mecutinea Roewer 1956, Phalangium montanum Sørensen 1910, Rhampsinitus montanus, Guruia mossambica Lawrence 1957, G. nigra Roewer 1912, G. obsti Roewer 1912, Phalangium (Guruia) palmatimanus Pocock 1902, G. palmatimana, G. palpinalis Roewer 1911, G. tetea Roewer 1956, G. ukrewia Roewer 1956] S84

*Gyas annulatus (Olivier 1792) C92 [=Phalangium annulatum C92; incl. P. bicolor Fabricius 1793 R23, Opilio nigricans Koch in Hahn & Koch 1848 R23, Leiobunum nigricans R23]

Himalphalangium spinulatum (Roewer 1911) [=Opilio spinulatus; incl. O. nigerrimus Schenkel 1953, O. pentaspinulatus Suzuki 1950] S03

*Leptobunus californicus Banks 1893 C84 [=Leptobrunus (l. c.) californicus C84; incl. *Liomitopus laevis Schenkel 1951 C92]

Lophopilio palpinalis (Herbst 1799) N05 [=Opilio palpinalis D04, Acantholophus palpinalis D04, Odiellus palpinalis D04, Odius palpinalis D04, Oligolophus palpinalis D04, Phalangium palpinale R23; incl. Opilio fuscatus Koch in Hahn & Koch 1848 R23, *Emonia labacensis Hadži 1973 C92, Phalangium spinulosum Hermann 1804 D04, Odiellus sublaevis Caporiacco 1940 M78, Acantholophus terricola Koch 1836 D04, Oligolophus terricola D04, Opilio terricola D04, Phalangium terricola R23, *Lophopilio tridentatus Hadži 1931 C92, Megabunus trispinifrons Morin 1931 D04]

Megabunus diadema (Fabricius 1779) DM10 [=Phalangium diadema D04; incl. M. grouvellei Simon 1881 D04, *M. insignis Meade 1855 C92]

Metadasylobus echinifrons (Simon 1879) D04 [=Dasylobus echinifrons D04, Phalangium echinifrons S73; incl. Dentizacheus zuluetai Rambla 1959 D04]

*Metadasylobus fuscoannulatus (Simon 1883) C92 [=Dasylobus fuscoannulatus S84; incl. *Dentizacheus tinerfensis Rambla 1956 C92]

*Metadasylobus grandissimus (Koch 1839) C92 [=Platylophus grandissimus C92, Platybunus grandissimus R12; incl. Opilio instratus Koch 1867 R23, Dasylobus instratus R23, Metadasylobus instratus S73, Phalangium instratum R23, Opilio pristes Koch 1867 R23, Dasylobus pristes R23, Metadasylobus pristes S73, Opilio vorax Koch 1867 R23, Metadasylobus vorax S73]

Odontobunus Roewer 1910 S84 [incl. Chelibunus Roewer 1915 C92, Cheops Sørensen 1910 C92, Ruandella Roewer 1956 C92; Odontobuninae]

Odontobunus armatus (Sørensen 1910) S84 [=*Cheops armatus C92; incl. *Chelibunus africanus Roewer 1915 non O. africanus Roewer 1910 C92, Cheops albidorsum Roewer 1915 S84, Cheops laevipes Roewer 1913 S84, Cheops minor Roewer 1913 S84]

Odontobunus punctatus (Roewer 1956) [=Odontosoma punctatum; incl. Odontosoma albofasciatum Roewer 1956, Odontosoma lossidanum Roewer 1956] S84

Oligolophus aspersus Karsch 1881 ZZ12 [=Lacinius aspersus TT05, Odiellus aspersus TT05; incl. Nelima melanodorsum Roewer 1911 ST83, Mitopus striatipes Roewer 1957 ST83]

Oligolophus hanseni (Kraepelin 1896) ZZ12 [=Acantholophus hanseni D04, A. ephippiger Hansen 1884 non Oligolophus ephippiger Simon 1879 D04]

*Oligolophus tridens (Koch 1836) C92 [=Opilio tridens BH09, Acantholophus tridens D04; incl. Oligolophus tridens f. obscura Lohmander 1945 M78]

*Paroligolophus agrestis (Meade 1855) C92 [=Opilio agrestis BH09, Acantholophus agrestis D04, Lophopilio agrestis D04, Oligolophus (Paroligolophus) agrestis D04; incl. Ol. ephippiger Simon 1879 D04]

Rafalskia olympica (Kulczyński 1903) C92 [=Platybunus strigosus olympicus C92, Metaplatybunus olympicus R23; incl. Phalangium werneri Kulczynski 1904 R23]

Scleropilio armatus (Roewer 1911) [=Opilio armatus; incl. Scutopilio diadema Gritsenko 1975, Scleropilio diadema] CL02

Scleropilio insolens (Simon 1895) S03 [=Egaenus insolens CL02; incl. Scutopilio elenae Gritsenko 1975 TTL00b, Scleropilio elenae TTL00b, Egaenus rugosus Schenkel 1963 S03, *Udezatus spinosus Nakatsudi 1943 TTL00b]

*Type species of generic name indicated

REFERENCES

[BH09] Bragg, P. D., & R. G. Holmberg. 2009. The harvestmen (Arachnida, Opiliones) of British Columbia. J. Entomol. Soc. Brit. Columbia 106: 29–37.

[CL02] Chemeris, A. N., & D. V. Logunov. 2002. Further taxonomic notes on the genus Scleropilio Roewer, 1911 (Arachnida: Opiliones: Phalangiidae). Arthropoda Selecta 11 (3): 209–222.

[C84] Cokendolpher, J. C. 1984. Revision of the harvestman genus Leptobunus and dismantlement of the Leptobunidae (Arachnida: Opiliones: Palpatores). Journal of the New York Entomological Society 92 (4): 371–402.

[C92] Crawford, R. L. 1992. Catalogue of the genera and type species of the harvestman superfamily Phalangioidea (Arachnida). Burke Museum Contributions in Anthropology and Natural History 8: 1–60.

[D04] Delfosse, E. 2004. Catalogue préliminaire des Opilions de France métropolitaine (Arachnida Opiliones). Bulletin de Phyllie 20: 34–58.

[DM10] Dunlop, J. A., & L. Mammitzsch. 2010. A new genus and species of harvestman from Baltic amber. Palaeodiversity 3: 23–32.

[G00] Giribet, G. 2000. Catalogue of the Cyphophthalmi of the world (Arachnida, Opiliones). Revista Ibérica de Aracnología 2: 49–76.

[GV09] Giribet, G., L. Vogt, A. Pérez González, P. Sharma & A. B. Kury. 2009. A multilocus approach to harvestman (Arachnida: Opiliones) phylogeny with emphasis on biogeography and the systematics of Laniatores. Cladistics 25: 1–30.

[HT12] Hedin, M., N. Tsurusaki, R. Macías-Ordóñez & J. W. Shultz. 2012. Molecular systematics of sclerosomatid harvestmen (Opiliones, Phalangioidea, Sclerosomatidae): geography is better than taxonomy in predicting phylogeny. Molecular Phylogenetics and Evolution 62 (1): 224–236.

[K85] Kauri, H. 1985. Opiliones from central Africa. Koninklijk Museum voor Midden-Afrika, Turveren, België, Zoologische Wetenschappen, Annalen 245: 1–167.

[K14] Kurt, K. 2014. Updated checklist of harvestmen (Arachnida: Opiliones) in Turkey. Arch. Biol. Sci. 66 (4): 1617–1631.

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Last updated: 29 November 2017.

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