Pseudoscorpiones

Pseudogarypus bicornis, copyright Mark Harvey.

Belongs within: Arachnida.
Contains: Chthoniidae, Iocheirata.

The Pseudoscorpiones are a group of small arachnids, superficially resembling scorpions due to the possession of large chelate pedipalps but lacking the elongate metasoma or posterior sting of a scorpion.

Characters (from Muchmore 1990): Body divided between anterior prosoma (cephalothorax) and posterior opisthosoma (abdomen), with the two sections broadly connected. Dorsal surface of prosoma covered by single large carapace; one or two pairs of eyes often present laterally toward the front. Venter of prosoma covered by coxae of palps and legs. Chelicerae two-segmented, chelate, attached close together under anterior margin of carapace; several setae present on chelicera with cluster of setae on medial side forming flagellum; comblike or membranous serrula present along side of each finger; spinneret (galea) present near end of movable finger. Pedipalps six-segmented, chelate; trichobothria present on chela. Legs six- or seven-segmented; pretarsus bearing two claws and arolium. Opisthosoma with twelve tergites; usually only eleven visible sternites; eleventh tergite and sternite sometimes fused as circumanal ring; twelfth tergite and sternite each very small anal plate. Spiracles present at lateral edges of third and fourth sternites.

<==Pseudoscorpiones [Cheliferinea, Chelonethi, Epiocheirata, Homosphyronida, Monosphyronida]
    |--+--Chthoniidae MHG08
    |  `--Iocheirata MHG08
    `--Feaelloidea MHG08
         |--Feaella [Feaellidae] MHG08
         |    `--F. anderseni MHG08
         `--Pseudogarypidae MHG08
              |--Neopseudogarypus scutellatus MHG08
              `--Pseudogarypus MHG08
                   |--P. banksi M90
                   `--P. bicornis MHG08

Pseudoscorpiones incertae sedis:
  ‘Acarus’ scorpioides Linnaeus 1758 H02
  Opsochernes carbophilus FF99
  Thelassochernes pallipes FF99
  Paratemnoides nidificator Balzan 1888 WR08
  Ectoceras T89
    |--E. bidens T89
    `--E. helferi T89
  Megathis T89
    |--M. desiderata T89
    `--M. kochii T89
  Obisium T89
    |--O. pallipes Lucas 1846 E12
    `--O. trifidum T89
  Dracochela Schawaller, Shear & Bonamo 1991 H02
    `--D. deprehendor Schawaller, Shear & Bonamo 1991 H02

*Type species of generic name indicated

REFERENCES

[E12] Evenhuis, N. L. 2012. Publication and dating of the Exploration Scientifique de l'Algérie: Histoire Naturelle des Animaux Articulés (1846–1849) by Pierre Hippolyte Lucas. Zootaxa 3448: 1–61.

[FF99] Forster, R., & L. Forster. 1999. Spiders of New Zealand and their World-wide Kin. University of Otago Press: Dunedin (New Zealand).

[H02] Harvey, M. S. 2002. The neglected cousins: what do we know about the smaller arachnid orders? Journal of Arachnology 30 (2): 357–372.

[M90] Muchmore, W. B. 1990. Pseudoscorpionida. In: Dindal, D. L. (ed.) Soil Biology Guide pp. 503–527. John Wiley & Sones: New York.

[MHG08] Murienne, J., M. S. Harvey & G. Giribet. 2008. First molecular phylogeny of the major clades of Pseudoscorpiones (Arthropoda: Chelicerata). Molecular Phylogenetics and Evolution 49: 170–184.

[T89] Thorell, T. 1889. Viaggio di Leonardo Fea in Birmania e regioni vicine. XXI.—Aracnidi Artrogastri Birmani raccolti da L. Fea nel 1885–1887. Annali del Museo Civico di Storia Naturale di Genova, Serie 2a 7: 521–729.

[WR08] Walsh, R. E., & L. S. Rayor. 2008. Kin discrimination in the amblypygid, Damon diadema. Journal of Arachnology 36 (2): 336–343.

Olpioidea

Synsphyronus sp., from Western Australian Museum.

Belongs within: Panctenata.
Contains: Olpiinae.

The Olpioidea was recognised by Harvey (1992) for the families Menthidae and Olpiidae on the basis of the presence of trichobothria on the pedipalpal femur. However, molecular phylogenetic analysis suggests that the Garypidae also fall within the clade formed by these families (Murienne et al. 2008).

<==Olpioidea
    |--Menthidae H06
    |    |--Oligomenthus H06
    |    |--Paramenthus H06
    |    |--Thenmus Harvey in Harvey & Muchmore 1990 H06
    |    |    |--*T. aigialites Harvey in Harvey & Muchmore 1990 H06
    |    |    `--T. augustus Harvey 2006 H06
    |    `--Menthus Chamberlin 1930 H06
    |         |--*M. rossi (Chamberlin 1923) [=Minniza rossi] H06
    |         |--M. californicus Chamberlin 1930 H06
    |         |--‘Atemnus’ gracilis [=Lustrochernes gracilis; incl. Menthus lindahli] H06
    |         `--M. mexicanus Hoff 1945 H06
    `--Olpiidae HL08
         |--+--Olpiinae HL08
         |  `--Garypidae HL08
         |       |  i. s.: Archeolarca rotunda M90
         |       |--Garypus Koch 1873 H10
         |       `--+--Ammogarypus Beier 1962 H10
         |          |--Anagarypus Chamberlin 1930 H10
         |          |    |--A. australianus OW01
         |          |    `--A. heatwolei Muchmore 1982 HL08
         |          |--Elattogarypus Beier 1964 H10
         |          |    |--E. cicatrosus Mahnert 2007 H10
         |          |    |--E. cruciatus Beier 1964 H10
         |          |    `--E. somalicus Mahnert 1984 H10
         |          |--Eremogarypus Beier 1955 H10
         |          |    `--E. eximius Beier 1973 H10
         |          |--Meiogarypus Beier 1955 H10
         |          |    `--M. mirus H10
         |          |--Neogarypus Vachon 1937 H10
         |          |--Paragarypus Vachon 1937 H10
         |          |--Thaumastogarypus Beier 1947 H10
         |          |    `--T. mancus Mahnert 1982 H10
         |          `--Synsphyronus Chamberlin 1930 H10 (see below for synonymy)
         |               |--*S. paradoxus Chamberlin 1930 H12
         |               |--S. apimelus Harvey 1987 HL08
         |               |--S. attiguus Harvey 1987 H12
         |               |--S. christopherdarwini Harvey 2012 H12
         |               |--S. elegans Beier 1954 H10
         |               |--S. ellenae Harvey 2010 H10
         |               |--S. francesae Harvey 2010 H10
         |               |--S. hansenii (With 1908) [=Garypus hansenii, *Idiogarypus hansenii] H10
         |               |--S. leo Harvey 1987 H10
         |               |--‘*Maorigarypus’ melanochelatus Chamberlin 1930 H12
         |               |--S. mimulus Chamberlin 1943 H12
         |               `--S. nullarborensis Beier 1969 H10
         `--Hesperolpiinae [Hesperolpiini] HL08
              |  i. s.: Aphelolpium Hoff 1964 HL08
              |           `--A. cayanum M90
              |         Calocheirus Chamberlin 1930 HL08
              |           `--C. tenerifae Mahnert 2002 HL08
              |         Cardiolpium Mahnert 1986 HL08
              |         Ectatolpium Beier 1947 HL08
              |         Hesperolpium Chamberlin 1930 HL08
              |           |--H. andrewsi M90
              |           `--H. slevini M90
              |         Planctolpium Hoff 1964 HL08
              |           |--P. peninsulae CS90
              |           `--P. suteri M90
              |         Stenolpiodes Beier 1959 HL08
              |         Stenolpium Beier 1955 HL08
              |--Nanolpium Beier 1947 MHG08, HL08
              `--+--Progarypus Beier 1931 MHG08, HL08
                 |    |--P. gracilis Mahnert 2001 HL08
                 |    |--P. liliae Mahnert 2001 HL08
                 |    |--P. nigrimanus Mahnert 2001 HL08
                 |    `--P. setifer Mahnert 2001 HL08
                 `--+--Apolpium Chamberlin 1930 MHG08, HL08
                    |    `--A. parvum Hoff 1945 HL08
                    `--Pachyolpium Beier 1931 MHG08, HL08
                         |--P. atlanticum Mahnert & Schuster 1981 H09
                         `--P. monae M90

Synsphyronus Chamberlin 1930 H10 [incl. Idiogarypus Chamberlin 1930 H12, Maorigarypus Chamberlin 1930 H12; Synsphyronidae]

*Type species of generic name indicated

REFERENCES

[CS90] Corey, D. T., & I. J. Stout. 1990. Ground surface arachnids in sandhill communities of Florida. Journal of Arachnology 18: 167–172.

Harvey, M. S. 1992. The phylogeny and classification of the Pseudoscorpionida (Chelicerata: Arachnida). Invertebrate Taxonomy 6: 1373–1435.

[H06] Harvey, M. S. 2006. New species and records of the pseudoscorpion family Menthidae (Pseudoscorpiones). Records of the Western Australian Museum 23 (2): 167–174.

[H09] Harvey, M. S. 2009. The first Australasian species of the halophilic pseudoscorpion genus Paraliochthonius (Pseudoscorpiones: Chthoniidae). Records of the Western Australian Museum 25 (3): 329–344.

[H10] Harvey, M. S. 2010. Two new species of Synsphyronus (Pseudoscorpiones: Garypidae) from southern Western Australian granite landforms. Records of the Western Australian Museum 26 (1): 11–22.

[H12] Harvey, M. S. 2012. A new species of Synsphyronus (Pseudoscorpiones: Garypidae) from Western Australia. Records of the Western Australian Museum 27 (1): 55–61.

[HL08] Harvey, M. S., & M. C. Leng. 2008. The first troglomorphic pseudoscorpion of the family Olpiidae (Pseudoscorpiones), with remarks on the composition of the family. Records of the Western Australian Museum 24 (4): 387–394.

[M90] Muchmore, W. B. 1990. Pseudoscorpionida. In: Dindal, D. L. (ed.) Soil Biology Guide pp. 503–527. John Wiley & Sones: New York.

[MHG08] Murienne, J., M. S. Harvey & G. Giribet. 2008. First molecular phylogeny of the major clades of Pseudoscorpiones (Arthropoda: Chelicerata). Molecular Phylogenetics and Evolution 49: 170–184.

[OW01] Otto, J. C., & K. J. Wilson. 2001. Assessment of the usefulness of ribosomal 18S and mitochondrial COI sequences in Prostigmata phylogeny. In: Halliday, R. B., D. E. Walter, H. C. Proctor, R. A. Norton & M. J. Colloff (eds) Acarology: Proceedings of the 10^th International Congress pp. 100–109. CSIRO Publishing: Melbourne.

Panctenata

Larca lata, copyright John Hallmén.

Belongs within: Iocheirata.
Contains: Olpioidea, Elassommatina.

The Panctenata are a clade of pseudoscorpions characterised by features of the chelicerae, including the presence of a velum (though this is absent in some olpioids) and lamina exterior (Harvey 1992).

Synapomorphies (from Harvey 1992): Velum present; lamina exterior present; movable cheliceral finger with one or two distal teeth or lobes; seta gs of movable cheliceral finger subdistal; serrula exterior completely fused to movable cheliceral finger.

Panctenata [Cheliferoidae, Diplochelonthi]
    |--+--Olpioidea MHG08
    |  `--Elassommatina MHG08
    `--Garypinidae [Garypininae] MHG08
         |  i. s.: Neominniza Beier 1930 HL08
         |         Oreolpium Benedict & Malcolm 1978 HL08
         |           `--O. nymphum M90
         |         Teratolpium Beier 1959 HL08
         |         Thaumatolpium Beier 1931 HL08
         |         Amblyolpium Simon 1898 HL08
         |         Neoamblyolpium Hoff 1956 HL08
         |           |--N. alienum M90
         |           `--N. giulianii M90
         |         Serianus Chamberlin 1930 H09
         |           `--S. carolinensis M90
         |         Aldabrinus floridanus M90
         |         Solinellus simberloffi M90
         |--+--Syarinus [Syarininae] MHG08
         |  `--Pseudogarypinus MHG08
         |       |--P. cooperi MHG08
         |       `--P. frontalis M90
         `--+--+--Neochelanops MHG08
            |  `--Protogarypinus Beier 1954 MHG08, ML08
            |       `--P. giganteus MHG08
            `--Larca [Larcidae] MHG08
                 |--L. fortunata Zaragoza 2005 F05
                 |--L. granulata M90
                 |--L. lata MHG08
                 |--L. lucentina Zaragoza 2005 F05
                 `--L. notha M90

*Type species of generic name indicated

REFERENCES

[F05] Fernández, J. 2005. Noticia de nuevos táxones para la ciencia en el ámbito Íbero-Balear y Macaronésico. Nuevos táxones animales descritos en la península Ibérica y Macaronesia desde 1994 (IX). Graellsia 61 (2): 261–282.

Harvey, M. S. 1992. The phylogeny and classification of the Pseudoscorpionida (Chelicerata: Arachnida). Invertebrate Taxonomy 6: 1373–1435.

[H08] Harvey, M. S. 2009. The first Australasian species of the halophilic pseudoscorpion genus Paraliochthonius (Pseudoscorpiones: Chthoniidae). Records of the Western Australian Museum 25 (3): 329–344.

[HL08] Harvey, M. S., & M. C. Leng. 2008. The first troglomorphic pseudoscorpion of the family Olpiidae (Pseudoscorpiones), with remarks on the composition of the family. Records of the Western Australian Museum 24 (4): 387–394.

[M90] Muchmore, W. B. 1990. Pseudoscorpionida. In: Dindal, D. L. (ed.) Soil Biology Guide pp. 503–527. John Wiley & Sones: New York.

[MHG08] Murienne, J., M. S. Harvey & G. Giribet. 2008. First molecular phylogeny of the major clades of Pseudoscorpiones (Arthropoda: Chelicerata). Molecular Phylogenetics and Evolution 49: 170–184.

Solifugae

Galeodes sp., copyright Arthur Anker.

Belongs within: Arachnida.

The Solifugae, variously called sun spiders, camel spiders or wind scorpions, are fast-moving arachnids found primarily in arid environments. They have a body divided between the prosoma and a segmented opisthosoma, and have conspicuously large, chelate two-segmented chelicerae.

Characters (from International Society of Arachnology): Body divided into two tagmata, the cephalothorax (prosoma) and abdomen (opisthosoma); appendages attached to prosoma. Dorsal plate of cephalothorax divided into propeltidium, mesopeltidium and metapeltidium. Abdomen segmented. Two eyes present, centrally located on dorsum of propeltidium. Chelicerae two-segmented, enormous, projecting forward from cephalothorax; male chelicerae bearing flagella or flagelliform setae. Pedipalps covered in sensory setae; cuticular lips present at tips of pedipalps that open to expose an eversible suctorial organ. Feeding via external digestion; sieve-like rostrum inside mouth opening filters out particulate material. Alimentary system consists of foregut, midgut and hindgut; hindgut bears lateral branches. Extensive tracheal system present that opens via prosomal and opisthosomal spiracles. Heart located dorsally in the abdomen. Brain consists of supra- and subesophageal ganglia. Malleoli (racquet organs) present as fan-like processes extending below ventral surface of last pair of legs. Female solifuge reproductive system includes pair of ovaries with pair of oviducts leading to a single genital chamber. Internal male reproductive system consists of two pairs of testes merging into vasa deferentia, which then merge into the genital chamber.

<==Solifugae [Solpugida]
    |  i. s.: Ammotrechella stimpsoni CS90
    |         Eremocosta gigasella RS10
    |--Protosolpuga [Protosolpugidae] H02
    |    `--*P. carbonaria Petrunkevitch 1913 H02
    |--Ceromidae R52
    |    |--*Cratosolpuga wunderlichi Selden 1996 H02
    |    `--Ceroma ornatum Karsch 1885 R52
    |--Eusimonia [Karschiidae] AP02
    |    |--E. mirabilis Roewer 1934 AP02
    |    `--E. wunderlichi Pieper 1977 GE02
    |--Ammotrechidae R54
    |    |  i. s.: *Happlodontus proterus Poinar & Santiago-Blay 1989 H02
    |    |         Oltacola gomezi Roewer 1934 AP02
    |    |         Procleobis patagonicus (Holmberg 1876) AP02
    |    |         Nothopuga DW10
    |    `--Ammotrechinae R54
    |         |--Ammotrechula schusterae Roewer 1954 R54
    |         `--Ammotrecha R54
    |              |--A. limbata R54
    |              `--A. stolli R54
    |--Galeodidae AP02
    |    |--Aellopus S69
    |    |--Rhax S69
    |    `--Galeodes Olivier 1791 BR05
    |         |--G. arabs Koch 1842 S69
    |         |--G. araneoides (Pallas 1772) AP02 [=Phalangium araneoides H02]
    |         |--G. barbarus T09
    |         |--G. brevipes S69
    |         |--G. fatalis [incl. G. vorax] S69
    |         |--G. orientalis Stoliczka 1869 S69
    |         `--G. subulata W68
    |--Hexisopodidae KBT11
    |    |--Hexisopus Karsch 1879 KBT11
    |    |    `--H. psammophilus Wharton 1981 KBT11
    |    `--Chelypus Purcell 1902 KBT11
    |--Gluvia [Daesiidae] KBT11
    |    `--G. dorsalis (Latreille 1817) KBT11
    |--Daesiidae KBT11
    |--Solpuga Lichtenstein 1796 H02 [Solpugidae KBT11]
    |    `--S. barbara Lucas 1846 E12
    `--Chanbria [Eremobatidae] DW10
         `--C. regalis Muma 1951 GE02

*Type species of generic name indicated

REFERENCES

[AP02] Alberti, G., & A. V. Peretti. 2002. Fine structure of male genital system and sperm in Solifugae does not support a sister-group relationship with Pseudoscorpiones (Arachnida). Journal of Arachnology 30 (2): 268–274.

[BR05] Bouchet, P., J.-P. Rocroi, J. Frýda, B. Hausdorf, W. Ponder, Á. Valdés & A. Warén. 2005. Classification and nomenclator of gastropod families. Malacologia 47 (1–2): 1–397.

[CS90] Corey, D. T., & I. J. Stout. 1990. Ground surface arachnids in sandhill communities of Florida. Journal of Arachnology 18: 167–172.

[DW10] Dabert, M., W. Witalinski, A. Kazmierski, Z. Olszanowski & J. Dabert. 2010. Molecular phylogeny of acariform mites (Acari, Arachnida): strong conflict between phylogenetic signal and long-branch attraction artifacts. Molecular Phylogenetics and Evolution 56 (1): 222–241.

[E12] Evenhuis, N. L. 2012. Publication and dating of the Exploration Scientifique de l'Algérie: Histoire Naturelle des Animaux Articulés (1846–1849) by Pierre Hippolyte Lucas. Zootaxa 3448: 1–61.

[GE02] Giribet, G., G. D. Edgecombe, W. C. Wheeler & C. Babbitt. 2002. Phylogeny and systematic position of Opiliones: a combined analysis of chelicerate relationships using morphological and molecular data. Cladistics 18: 5–70.

[H02] Harvey, M. S. 2002. The neglected cousins: what do we know about the smaller arachnid orders? Journal of Arachnology 30 (2): 357–372.

[KBT11] Klann, A. E., T. L. Bird & G. Talarico. 2011. Ultrastructural characterization of Hexisopus psammophilus (Arachnida: Solifugae: Hexisopodidae) spermatozoa in comparison to other solifuge spermatozoal traits. Journal of Arachnology 39 (2): 280–286.

[RS10] Regier, J. C., J. W. Shultz, A. Zwick, A. Hussey, B. Ball, R. Wetzer, J. W. Martin & C. W. Cunningham. 2010. Arthropod relationships revealed by phylogenomic analysis of nuclear protein-coding sequences. Nature 463: 1079–1083.

[R52] Roewer, C. F. 1952. Opilioniden and Solifugen aus Ost-Afrika. Veröffentlichungen aus dem Deutschen Kolonial- und Uebersee-Museum in Bremen, Reihe A 2: 87–89.

[R54] Roewer, C. F. 1954. Spinnentiere aus El Salvador, I. (Arachnoidea: Pedipalpi, Solifuga, Opiliones-Laniatores). Senckenbergiana Biologica 35: 57–73.

[S69] Stoliczka, F. 1869. Contribution to the knowledge of Indian Arachnoidea. Journal of the Asiatic Society of Bengal 38 pt 2 (4): 201–251, pls 18–20.

[T09] Taylor, C. K. 2009. Australiscutum, a new genus of Monoscutidae (Arachnida: Opiliones) from eastern Australia, with the first record of asymmetrical chelicerae in Opiliones. Insect Systematics and Evolution 40: 319–332.

[W68] Wood, H. C., Jr. 1868. On the Phalangeae of the United States of America. Communications of the Essex Institute 6: 10–40.

Chthoniidae

Austrochthonius sp., copyright Kristi Ellingsen.

Belongs within: Pseudoscorpiones.
Contains: Tyrannochthoniini.

The Chthoniidae are a group of pseudoscorpions characterised by differing number of segments in the anterior and posterior pairs of legs (five post-coxal segments in legs I and II vs six in legs III and IV) (Muchmore 1990). Three families have been recognised within the superfamily Chthonioidea, the Chthoniidae, Lechytiidae and Tridenchthoniidae, but molecular phylogenetic analysis indicates that the first is paraphyletic to the latter two (Murienne et al. 2008).

<==Chthoniidae [Chthoniinea, Chthonioidea, Heterosphyronida]
    |--Pseudotyrannochthoniinae MHG08
    |    |--Afrochthonius godfreyi MHG08
    |    `--+--Pseudotyrannochthonius MHG08
    |       `--Selachochthonius MHG08
    `--Chthoniinae MHG08
         |--Chthonius Koch 1843 MHG08, H09
         |    |--C. californicus S62
         |    |--C. coecus Packard 1884 HM06
         |    `--C. tetrachelatus M90
         `--+--+--Apochthonius MHG08
            |  `--Kleptochthonius MHG08
            `--+--+--+--Sathrochthonius insulanus MHG08
               |  |  `--Lechytia [Lechytiidae, Lechytiini] MHG08
               |  |       `--L. hoffi MHG08
               |  `--+--Pseudochthonius squamosus MHG08, L88
               |     `--Tridenchthoniidae [Dithidae, Dithinae, Tridenchthoniinae] M90
               |          |--Anaulacodithella MHG08
               |          |--Verrucaditha spinosa M90
               |          `--Tridenchthonius M90
               `--+--Tyrannochthoniini MHG08
                  `--+--Drepanochthonius MHG08
                     `--Austrochthonius Chamberlin 1929 MHG08, HM06 (see below for synonymy)
                          |--*A. chilensis (Chamberlin 1923) [=Chthonius chilensis] HM06
                          |    |--A. c. chilensis HM06
                          |    |--A. c. magalhanicus HM06
                          |    `--A. c. transversus HM06
                          |--A. argentinae HM06
                          |--A. australis HM06
                          |--A. bolivianus HM06
                          |--A. cavicola HM06
                          |--A. easti HM06
                          |--A. iguazuensis HM06
                          |--A. insularis HM06
                          |--A. mordax HM06
                          |--A. muchmorei Harvey & Mould 2006 (see below for synonymy) HM06
                          |--A. paraguayensis HM06
                          |--A. parvus (Mello-Leitão 1939) [=*Cecoditha parva] HM06
                          |--A. persimilis HM06
                          |--A. rapax HM06
                          |--A. semiserratus HM06
                          |--A. strigosus Harvey & Mould 2006 HM06
                          |--A. tullgreni (Beier 1931) [=*Paraustrochthonius tullgreni] HM06
                          `--A. zealandicus HM06
                               |--A. z. zealandicus HM06
                               `--A. z. obscurus HM06

Chthoniidae incertae sedis:
  Chiliochthonius HM06
  Francochthonius HM06
  Malcolmochthonius HM06
  Mexichthonius HM06
  Tyrannochthoniella HM06
  Congochthonius HM06
  Sathrochthoniella HM06
  Aphrastochthonius M90
  Kewochthonius M90
    |--K. amplus Schuster 1962 S62
    |--K. paludis S62
    |--K. spingolus Schuster 1962 S62
    `--K. stanfordianus Chamberlin 1929 S62

Austrochthonius Chamberlin 1929 MHG08, HM06 [incl. Cecoditha Mello-Leitão 1939 HM06, Paraustrochthonius Beier 1931 HM06]

Austrochthonius muchmorei Harvey & Mould 2006 [=Chthonius caecus Tullgren 1909 non Simon 1885, Mundochthonius caecus, Sathrochthonius tullgreni Chamberlin 1962 non Paraustrochthonius tullgreni Beier 1931] HM06

*Type species of generic name indicated

REFERENCES

[H02] Harvey, M. S. 2009. The first Australasian species of the halophilic pseudoscorpion genus Paraliochthonius (Pseudoscorpiones: Chthoniidae). Records of the Western Australian Museum 25 (3): 329–344.

[HM06] Harvey, M. S., & L. G. Mould. 2006. A new troglomorphic species of Austrochthonius (Pseudoscorpiones: Chthoniidae) from Australia, with remarks on Chthonius caecus. Records of the Western Australian Museum 23 (2): 205–211.

[L88] Liebherr, J. K. 1988. General patterns in West Indian insects, and graphical biogeographic analysis of some circum-Caribbean Platynus beetles (Carabidae). Systematic Zoology 37 (4): 385–409.

[M90] Muchmore, W. B. 1990. Pseudoscorpionida. In: Dindal, D. L. (ed.) Soil Biology Guide pp. 503–527. John Wiley & Sones: New York.

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