Oryzomys

Coues' rice rat Oryzomys couesi, copyright Paulina Arias Caballero de Miguel.


Belongs within: Cricetidae.

Oryzomys, the rice rats, are a genus of semiaquatic rodents found in southern North and South America. Members of this genus have short and coarse fur, relatively long tails and broad hind feet (Goodwin 1969).

<==Oryzomys
    |--O. (Oryzomys) G69
    |    |--O. (O.) alfaroi G69
    |    |    |--O. a. alfaroi G69
    |    |    |--O. a. chapmani G69
    |    |    |--O. a. gloriaensis G69
    |    |    |--O. a. guerrerensis G69
    |    |    `--O. a. palatinus G69
    |    |--O. (O.) caudatus [=G. alfaroi caudatus] G69
    |    |--O. (O.) couesi G69
    |    |    |--O. c. couesi G69
    |    |    |--O. c. aztecus G69
    |    |    `--O. c. mexicanus G69
    |    `--O. (O.) melanotis G69
    |         |--O. m. melanotis G69
    |         `--O. m. rostratus G69
    `--O. (Oligoryzomys) G69
         |--‘Oligoryzomys’ andinus IT07
         |--‘Oligoryzomys’ arenalis IT07
         |--‘Oligoryzomys’ chacoensis IT07
         |--‘Oligoryzomys’ delticola IT07
         |--‘Oligoryzomys’ destructor IT07
         |--‘Oligoryzomys’ eliurus IT07
         |--‘Oligoryzomys’ flavescens IT07
         |--O. (O.) fulvescens G69
         |    |--O. f. fulvescens G69
         |    |--O. f. lenis G69
         |    `--O. f. pacificus G69
         |--‘Oligoryzomys’ griseolus IT07
         |--‘Oligoryzomys’ longicaudatus IT07
         |--‘Oligoryzomys’ magellanicus IT07
         |--‘Oligoryzomys’ microtis IT07
         |--‘Oligoryzomys’ nigripes IT07
         |--‘Oligoryzomys’ vegetus IT07
         `--O. (O.) victus Thomas 1898 IT07, AC98

Oryzomys incertae sedis:
  O. albigularis IT07
  O. aphrastus BP87
  O. argentatus BP87
  O. auriventer IT07
  O. balneator IT07
  O. bauri BP87
  O. bolivaris IT07
  O. buccinatus IT07
  O. capito IT07
  O. cozumelae BP87
  O. devius IT07
  O. dimidiatus IT07
  O. emmonsae LMZ05
  O. fernandinae BP87
  O. galapagoensis IT07
  O. gorgasi IT07
  O. hammondi IT07
  O. intectus IT07
  O. intermedius IT07
  O. keaysi IT07
  O. kelloggi IT07
  O. lamia IT07
  O. legatus IT07
  O. levipes IT07
  O. macconnelli PCS05
  O. megacephalus PCS05
  O. narboroughi BP87
  O. nelsoni BP87
  O. nitidus PCS05
  O. oniscus IT07
  O. palustris IT07
  O. peninsulae BP87
  O. polius IT07
  O. ratticeps IT07
  O. rhabdops IT07
  O. saturatior IT07
  O. subflavus IT07
  O. talamancae IT07
  O. xantheolus IT07
  O. yunganus IT07

*Type species of generic name indicated

REFERENCES

[AC98] Alcover, J. A., X. Campillo, M. Macias & A. Sans. 1998. Mammal species of the world: additional data on insular mammals. American Museum Novitates 3248: 1–29.

[BP87] Burton, J. A., & B. Pearson. 1987. Collins Guide to the Rare Mammals of the World. Collins: London.

[G69] Goodwin, G. G. 1969. Mammals from the State of Oaxaca, Mexico, in the American Museum of Natural History. Bulletin of the American Museum of Natural History 141 (1): 1–269, 40 pls.

[IT07] Isaac, N. J. B., S. T. Turvey, B. Collen, C. Waterman & J. E. M. Baillie. 2007. Mammals on the EDGE: conservation priorities based on threat and phylogeny. PloS One 2 (3): e296.

[LMZ05] Lambert, T. D., J. R. Malcolm & B. L. Zimmerman. 2005. Variation in small mammal species richness by trap height and trap type in southeastern Amazonia. Journal of Mammalogy 86 (5): 982–990.

[PCS05] Percequillo, A. R., A. P. Carmignotto & M. J. de J. Silva. 2005. A new species of Neusticomys (Ichthyomyini, Sigmodontinae) from central Brazilian Amazonia. Journal of Mammalogy 86 (5): 873–880.

Neotoma

Bushy-tailed woodrat Neotoma cinerea, from here.


Belongs within: Neotominae.

Neotoma, the woodrats or pack rats, are a genus of rodents found in North America. Members of this genus are relatively large, with conspicuously hairy and elongate tails (Goodwin 1969).

<==Neotoma
    |--N. cinerea RC06
    |    |--N. c. cinerea M-PV-G01
    |    `--N. c. acraia (Elliot 1904) M-PV-G01
    `--+--N. leucodon Merriam 1894 RC06, MB86 [=N. albigula leucodon MB86]
       `--N. mexicana Baird 1855 RC06, MB86
            |--N. m. mexicana MB86
            |--N. m. isthmica G69
            |--N. m. parvidens G69
            |--N. m. picta G69
            |--N. m. tenuicauda Merriam 1892 MB86
            `--N. m. tropicalis G69

Neotoma incertae sedis:
  N. albigula Hartley 1894 M-PV-G01
  N. angustapalata IT07
  N. anthonyi IT07
  N. bryanti IT07
  N. bunkeri IT07
  N. chrysomelas IT07
  N. floridana MHL03
    |--N. f. floridana BP87
    `--N. f. smalli BP87
  N. fuscipes IT07
  N. goldmani Merriam 1903 MB86
  N. lepida Thomas 1893 M-PV-G01
    |--N. l. lepida H42
    |--N. l. devia H42
    |--N. l. felipensis H42
    |--N. l. gilva H42
    |--N. l. grinnelli Hall 1942 H42
    |--N. l. latirostra MBJ83
    |--N. l. monstrabilis H42
    `--N. l. nevadensis H42
  N. martinensis IT07
  N. micropus IT07
  N. nelsoni IT07
  N. ozarkensis MHL03
  N. palatina Goldman 1905 MB86
  N. phenax IT07
  N. stephensi IT07
  N. taylori MH03
  N. varia IT07

*Type species of generic name indicated

REFERENCES

[BP87] Burton, J. A., & B. Pearson. 1987. Collins Guide to the Rare Mammals of the World. Collins: London.

[G69] Goodwin, G. G. 1969. Mammals from the State of Oaxaca, Mexico, in the American Museum of Natural History. Bulletin of the American Museum of Natural History 141 (1): 1–269, 40 pls.

[H42] Hall, E. R. 1942. A new race of wood rat (Neotoma lepida). University of California Publications in Zoology 46 (5): 369–370.

[IT07] Isaac, N. J. B., S. T. Turvey, B. Collen, C. Waterman & J. E. M. Baillie. 2007. Mammals on the EDGE: conservation priorities based on threat and phylogeny. PloS One 2 (3): e296.

[MH03] Martin, R. A., R. T. Hurt, J. G. Honey & P. Peláez-Campomanes. 2003. Late Pliocene and Early Pleistocene rodents fom the northern Borchers Badlands (Meade County, Kansas), with comments on the Blancan-Irvingtonian boundary in the Meade Basin. Journal of Paleontology 77 (5): 985–1001.

[MB86] Matson, J. O. & R. H. Baker. 1986. Mammals of Zacatecas. Special Publications, Museum of Texas Tech University 24: 1–88.

[MHL03] Meng, J., Y. Hu & C. Li. 2003. The osteology of Rhombomylus (Mammalia, Glires): Implications for phylogeny and evolution of Glires. Bulletin of the American Museum of Natural History 275: 1–247.

[M-PV-G01] Montiel-Parra, G., G. A. Villegas-Guzman, M. Vargas & O. J. Polaco. 2001. Mites associated with nests of Neotoma albigula Hartley, 1894 (Rodentia: Muridae) in Durango, México. In: Halliday, R. B., D. E. Walter, H. C. Proctor, R. A. Norton & M. J. Colloff (eds) Acarology: Proceedings of the 10th International Congress pp. 586–593. CSIRO Publishing: Melbourne.

[MBJ83] Munger, J. C., M. A. Bowers & W. T. Jones. 1983. Desert rodent populations: factors affecting abundance, distribution, and genetic structure. Great Basin Naturalist Memoirs 7: 91–116.

[RC06] Reeder, S. A., D. S. Carroll, C. W. Edwards, C. W. Kilpatrick & R. D. Bradley. 2006. Neotomine-peromyscine rodent systematics based on combined analyses of nuclear and mitochondrial DNA sequences. Molecular Phylogenetics and Evolution 40 (1): 251–258.

Peromyscus (Peromyscus) maniculatus

North American deer mouse Peromyscus maniculatus, copyright J. N. Stuart.


Belongs within: Peromyscus.

The North American deer mouse Peromyscus maniculatus is a variable species found over the greater part of North America.

See also: The gordian mouse.

Characters (from Animal Diversity Web): Body round and slender; size small, typically 119 to 222 mm long, weighing between 10 and 24 grams; tail length variable, ranging from 45 mm to 105 mm. Woodland forms typically larger with larger tails and feet than prairie forms. Coloration grayish to reddish brown with white underparts. Fur short, soft, dense. Nose pointed; eyes large, black, beady; ears large with little fur. Vibrissae long and prominent. Forelimbs shorter than hind limbs; hind feet 22 mm or less. Tail finely-haired, bicolored, the darker top half and lighter bottom sharply differentiated. Dental formula 1/1 0/0 0/0 3/3; molars low-crowned, cuspidate; third upper molar less wide than first two.

<==Peromyscus (Peromyscus) maniculatus
    |--P. m. maniculatus CZ75
    |--P. m. algidus CZ75
    |--P. m. artemisiae CZ75
    |--P. m. austerus CZ75
    |--P. m. bairdii (Hoy & Kennicott 1857) [=Mus bairdii] B75
    |--P. m. blandus Osgood 1904 MB86
    |--P. m. borealis CZ75
    |--P. m. fulvus G69
    |--P. m. gambeli CZ75
    |--P. m. gracilis CZ75
    |--P. m. keeni J68
    |--P. m. labecula Elliot 1903 MB86
    |--P. m. luteus CZ75
    |--P. m. macrorhinus CZ75
    |--P. m. nebrascensis (Coues in Coues & Allen 1877) [=Hesperomys sonoriensis var. nebrascensis] J61
    |--P. m. oreas CZ75
    |--P. m. osgoodi CZ75
    |--P. m. ozarkiarum CZ75
    |--P. m. pallescens CZ75
    |--P. m. prevostensis J68
    `--P. m. sonoriensis HD37

*Type species of generic name indicated

REFERENCES

[B75] Bowles, J. B. 1975. Distribution and biogeography of mammals of Iowa. Special Publications, The Museum, Texas Tech University 9: 1–184.

[CZ75] Caire, W. & E. G. Zimmerman. 1975. Chromosomal and morphological variation and circular overlap in the deer mouse, Peromyscus maniculatus, in Texas and Oklahoma. Systematic Zoology 24 (1): 89–95.

[G69] Goodwin, G. G. 1969. Mammals from the State of Oaxaca, Mexico, in the American Museum of Natural History. Bulletin of the American Museum of Natural History 141 (1): 1–269, 40 pls.

[HD37] Hall, E. R., & S. D. Durrant. 1937. A new kangaroo mouse (Microdipodops) of Utah and Nevada. Journal of Mammalogy 18 (3): 357–359.

[J68] Johnson, M. L. 1968. Application of blood protein electrophoretic studies to problems in mammalian taxonomy. Systematic Zoology 17 (1): 23–30.

[J61] Jones, J. K., Jr. 1961. A syntype of Peromyscus maniculatus nebrascensis (Coues). Proceedings of the Biological Society of Washington 74: 101–104.

[MB86] Matson, J. O. & R. H. Baker. 1986. Mammals of Zacatecas. Special Publications, Museum of Texas Tech University 24: 1–88.

Tylomyinae

Watson's climbing rat Tylomys watsoni, copyright D. H. Janzen.


Belongs within: Cricetidae.

The Tylomyinae, climbing rats and vesper rats, are a group of arboreal rodents found in Mexico and Central America.

Characters (from Animal Diversity Web): Medium to large-sized; 95 to 255 mm in length; tail 85 to 250 mm in length, tail usually slightly longer body; weight 29 to 280 grams. Fur short or long; cinnamon, buff, tawny, gray, russet, or brown above, white below. Ears nearly naked, short or long. Whiskers long and black. Eyes quite large, sometimes each surrounded by a dark ring. Tail either covered with long hairs and tufted at tip, or nearly naked. Hind feet modified for climbing. Two pairs of mammae present in the inguinal region. Molars brachydont, cuspidate, with major cusps lying opposite one another; cheek teeth bearing well-developed mesolophs and mesolophids; second upper molar with four roots; third lower molar relatively large with a crown pattern like that of the second lower molar. Interorbital region cuneate, with prominant supraorbital shelves continuing posteriorly as pronounced temporal ridges. Interparietal bone large, contacting the squamosal, to which the tegmen tympani are united. Zygomatic plates narrow, usually without a dorsal notch. Alisphenoid strut present; subsquamosal fenestra absent; postglenoid foramen quite small. Mesopterygoid fossa usually completely ossified; parapterygoid fossa shallow and slender. Sphenopalatine vacuities, if present, tiny slits. First rib attaching to only first thoracic vertebra. Humerus with entepicondylar foramen. Calcaneum with wide, proximally-positioned trochlear process. Stomach single-chambered; gall bladder absent. Caecum long and complex. Glans penis wide and short (though longer than baculum), with large, well-spaced spines.

<==Tylomyinae [Tylomyini]
    |--Ototylomys phyllotis RC06
    |--Otonyctomys hatti RC06, IT07
    |--Nyctomys sumichrasti RC06, G69
    |    |--N. s. sumichrasti G69
    |    |--N. s. colimensis G69
    |    `--N. s. pallidulus G69
    `--Tylomys RC06
         |--T. bullaris G69
         |--T. fulviventer IT07
         |--T. mirae IT07
         |--T. nudicaudus RC06
         |    |--T. n. nudicaudus G69
         |    |--T. n. gymnurus G69
         |    |--T. n. microdon G69
         |    |--T. n. tumbalensis G69
         |    `--T. n. villai Schaldach 1966 [=T. n. villae] G69
         |--T. panamensis IT07
         `--T. watsoni IT07

*Type species of generic name indicated

REFERENCES

[G69] Goodwin, G. G. 1969. Mammals from the State of Oaxaca, Mexico, in the American Museum of Natural History. Bulletin of the American Museum of Natural History 141 (1): 1–269, 40 pls.

[IT07] Isaac, N. J. B., S. T. Turvey, B. Collen, C. Waterman & J. E. M. Baillie. 2007. Mammals on the EDGE: conservation priorities based on threat and phylogeny. PloS One 2 (3): e296.

[RC06] Reeder, S. A., D. S. Carroll, C. W. Edwards, C. W. Kilpatrick & R. D. Bradley. 2006. Neotomine-peromyscine rodent systematics based on combined analyses of nuclear and mitochondrial DNA sequences. Molecular Phylogenetics and Evolution 40 (1): 251–258.

Swertia

Many-stem swertia Swertia multicaulis, copyright Pema G. Bhutia.


Belongs within: Gentianoideae.

Swertia, the felworts, is a cosmopolitan genus of herbs, most diverse in Asia and Africa.

Characters (from Flora of China): Herbs annual or perennial. Roots fibrous or woody; primary roots with few secondary rootlets or rhizomes short and with few fleshy adventitious rootlets. Stems absent, scapiform, or well developed, ascending or erect, terete, striate or angled, simple or branched. Leaves opposite, rarely alternate or whorled, margin entire. Inflorescences cymose, usually grouped into simple or paniculate thyrses, rarely strictly dichotomous, sometimes reduced to single flowers and inflorescences racemelike or flowers solitary and terminal. Flowers 4- or 5-merous. Calyx and corolla rotate, lobed to base, tubes less than 3 mm. Nectaries 1 or 2 per corolla lobe, with fimbriate margin or represented by naked spotlike gland patches. Stamens inserted at base of corolla tube. Ovary 1-celled. Style short to elongate. Capsules 2-valved, few to many seeded. Seeds small.

Swertia
    |--S. acaulis O88
    |--S. alternifolia O88
    |--S. candelabrum O88
    |--S. ciliata O88
    |--S. cuneata O88
    |--S. dilatata O88
    |--S. hispidicalyx O88
    |--S. hookeri O88
    |--S. kingii O88
    |--S. multicaulis O88
    |--S. petiolata O88
    |--S. racemosa O88
    |--S. radiata RS99
    |--S. ramosa O88
    |--S. speciosa O88
    `--S. staintonii O88

*Type species of generic name indicated

REFERENCES

[O88] Ohba, H. 1988. The alpine flora of the Nepal Himalayas: An introductory note. In: Ohba, H. & S. B. Malla (eds) The Himalayan Plants vol. 1. The University Museum, University of Tokyo, Bulletin 31: 19–46.

[RS99] Rossman, A. Y., G. J. Samuels, C. T. Rogerson & R. Lowen. 1999. Genera of Bionectriaceae, Hypocreaceae and Nectriaceae (Hypocreales, Ascomycetes). Studies in Mycology 42: 1–248.

Anapidae

Comaroma simoni, copyright Pierre Oger.


Belongs within: Araneoidea.

The Anapidae are a group of minute spiders found in litter and moss, usually in moist forests. Many species construct small horizontal orb webs (Jocqué & Dippenaar-Schoeman 2007). Females lack palpi and also have secondarily haplogyne genitalia without surface expression on the abdomen (Paquin et al. 2010).

Characters (from Jocqué & Dippenaar-Schoeman 2007): Body size <2 mm. Carapace with ocular region elevated; pedicel arising from circular rimmed cavity on posterior declivity; glandular openings often present, situated in hollow pits on edge of carapace at anterolateral corners. Six or eight eyes present; anterior median eyes often reduced or absent; other eyes arranged in three pairs. Chelicerae with teeth on both margins. Labrum bearing anterior spur projecting forward between chelicerae, sometimes reduced to a small crest. Female palp reduced in size or absent, without claw. Sternum as broad as long, broadly fused to carapace. Relative leg lengths (longest to shortest) 4123 or 1423; legs short, without spines; metatarsi shorter than tarsi; tarsi with three claws; tarsal organ capsulate and domed; femora I of male with short spine ventrally. Abdomen of male with large dorsal scutum, or two or three scuta, or dorsal region with scattered small sclerotised areas; ventral scutum surrounding pedicel. Abdomen of female soft or with small, brown sclerotised ring. Two booklungs present or anterior tracheae opening through spiracles on either side of genital opening. Anterior spinnerets small, close together; spinnerets together with colulus surrounded by a sclerotised ring; posterior lateral spinnerets with triplet of one flagelliform and two aggregate gland spigots in both sexes, aggregate gland spigot larger than accompanying flagelliform spigot; piriform gland spigots with reduced base. Genitalia haplogyne; spermathecae well separated; male palp variable, femur and patella sometimes with bristles or apophyses, embolus varying from spiniform to short and wide.

<==Anapidae [Amazulinae, Anapeae, Anapinae]
    |--Comaroma Bertkau 1889 S03
    |    `--C. simoni Bertkau 1889 S03
    `--+--Anapis Simon 1895 S03
       |    `--A. amazonas Platnick & Shadab 1978 S03
       `--Crozetulus Hickman 1939 S03
            |--*C. minutus Hickman 1939 H39
            `--C. rhodesiensis Brignoli 1981 S03

Anapidae incertae sedis:
  Sofanapis antillanca Platnick & Forster 1989 A02
  Crassanapis chilensis R14
  Pseudanapis Simon 1905 S03
    `--P. burra Forster 1959 W76
  Chasmocephalon O.P.-Cambridge 1889 S03
    `--C. crassifemoralis Wunderlich 1976 W76
  Anapisona S03
  Novanapis Platnick & Forster 1989 PVD10
    `--*N. spinipes Platnick & Forster 1989 PVD10
  Gertschanapis shantzi BS09
  Paranapis Platnick & Forster 1989 PVD10
    |--*P. insula (Forster 1951) PVD10
    `--P. isolata Platnick & Forster 1989 PVD10
  Zealanapis Platnick & Forster 1989 PVD10
    |--*Z. armata (Forster 1951) PVD10
    |--Z. australis (Forster 1951) PVD10
    |--Z. conica (Forster 1951) PVD10
    |--Z. insula Platnick & Forster 1989 PVD10
    |--Z. kuscheli Platnick & Forster 1989 PVD10
    |--Z. matua Platnick & Forster 1989 PVD10
    |--Z. montana Platnick & Forster 1989 PVD10
    |--Z. otago Platnick & Forster 1989 PVD10
    |--Z. punta Platnick & Forster 1989 PVD10
    `--Z. waipoua Platnick & Forster 1989 PVD10
  Palaeoanapis nana Wunderlich 1988 S93

*Type species of generic name indicated

REFERENCES

[A02] Agnarsson, I. 2002. Sharing a web—on the relation of sociality and kleptoparasitism in theridiid spiders (Theridiidae, Araneae). Journal of Arachnology 30 (2): 181–188.

[BS09] Blackledge, T. A., N. Scharff, J. A. Coddington, T. Szüts, J. W. Wenzel, C. Y. Hayashi & I. Agnarsson. 2009. Reconstructing web evolution and spider diversification in the molecular era. Proceedings of the National Academy of Sciences of the USA 106 (13): 5229–5234.

[H39] Hickman, V. V. 1939. Opiliones and Araneae. British, Australian and New Zealand Antarctic Research Expedition Reports Series B 4: 157–188.

[PVD10] Paquin, P., C. J. Vink & N. Dupérré. 2010. Spiders of New Zealand: annotated family key and species list. Manaaki Whenua Press: Lincoln (New Zealand).

[R14] Ramírez, M. J. 2014. The morphology and phylogeny of dionychan spiders (Araneae: Araneomorphae). Bulletin of the American Museum of Natural History 390: 1–374.

[S03] Schütt, K. 2003. Phylogeny of Symphytognathidae s.l. (Araneae, Araneoidea). Zoologica Scripta 32 (2): 129–151.

[S93] Selden, P. A. 1993. Arthropoda (Aglaspidida, Pycnogonida and Chelicerata). In: Benton, M. J. (ed.) The Fossil Record 2 pp. 297–320. Chapman & Hall: London.

[W76] Wunderlich, J. 1976. Spinnen aus Australien. 1. Uloboridae, Theridiosomatidae und Symphytognathidae (Arachnida: Araneida). Senckenbergiana Biologica 57 (1–3): 113–124.

Last updated: 15 January 2022.

Araneus

European garden spider Araneus diadematus, copyright Böhringer Friedrich.


Belongs within: Araneini.

Araneus is a genus of orb-weaving spiders including the European garden spider Araneus diadematus and related species. The name has been used in a much broader sense in the past (at its broadest, as a generic name for all spiders); many of the species listed below would now be placed in segregate genera. The type species, A. angulatus, is a widespread species in the Palaearctic region characterised by angular lateral humps on the opisthosoma, which is mostly dark with a light cross-shaped marking between the humps. The European garden spider A. diadematus has a similar colour pattern but lacks the prominent angular humps.

<==Araneus Clerck 1758 F11 (see below for synonymy)
    |--*A. angulatus Clerck 1757 G76 [=Aranea angulata AM11, Epeira angulata K01]
    |--A. acuminatus (K. 1871) [=Epeira acuminata] R13
    |--‘Epeira’ albomaculata Lucas 1846 E12
    |--‘Epeira’ albotriangula Koch & Keyserling 1887 KK90
    |--‘Epeira’ alpica K01
    |--A. alsine Walckenaer 1802 G76 [=Epeira alsine K01]
    |--A. anantnagensis Tikader & Bal 1981 TB81
    |--‘Epeira’ anatipes Koch & Keyserling 1887 KK90
    |--‘Epeira’ annulata Koch & Keyserling 1886 KK90
    |--‘Epeira’ anseripes Walckenaer 1841 [incl. Cyclosa melanura Simon 1877] KK90
    |--‘Epeira’ arenacea Koch & Keyserling 1886 KK90
    |--‘Epeira’ armida K01
    |--‘Aranea’ arundinacea Linnaeus 1758 L58
    |--‘Epeira’ balanus Doleschall 1859 D59
    |--‘Epeira’ bertkani Koch & Keyserling 1890 KK90
    |--A. biapicatifer Strand 1907 [=Aranea biapicatifera] J98
    |--A. bicentenarius DC01
    |--‘Epeira’ bifida Doleschall 1859 D59
    |--A. bilunifer (Pocock 1900) HU08
    |--‘Aranea’ bipunctata Linnaeus 1758 L58
    |--A. bituberculatus (Walckenaer 1802) TB81 (see below for synonymy)
    |--‘Epeira’ bogoriensis Doleschall 1859 D59
    |--‘Epeira’ bradleyi Koch & Keyserling 1887 KK90
    |--A. brisbanae (Koch 1867) PVD10 [=Epeira brisbanae KK90]
    |--A. brounii Urquhart 1844 S99 [=Aranea brouni M54]
    |--‘Aranea’ caementaria L02a
    |--‘Aranea’ calophylla Walckenaer 1802 G76
    |--‘Epeira’ caputlupi Doleschall 1859 D59
    |--‘Epeira’ carbonaria K01
    |--A. cavaticus (Keyserling 1881) B11
    |--A. cingulatus DC01
    |--A. circe Savigny & Audouin 1825 G76 [=Epeira circe K01]
    |--‘Epeira’ circulissparsa Koch & Keyserling 1887 KK90
    |--‘Epeira’ collina Koch & Keyserling 1886 KK90
    |--‘Aranea’ corollata Linnaeus 1758 L58
    |--‘Epeira’ coronata Rainbow 1897 R97
    |--‘Epeira’ cossoni Sim. 1885 K01
    |--A. cucurbitinus Clerck 1757 TB81 [=Aranea cucurbitina L58, Araniella cucurbitina G76, Epeira cucurbitina V09]
    |--‘Epeira’ cyrtarachnoides Koch & Keyserling 1887 KK90
    |--‘Epeira’ dehaanii Doleschall 1859 D59
    |--A. diadematus Clerck 1757 G76 (see below for synonymy)
    |--‘Aranea’ dorsalis L02a
    |--A. eburniventris MC13
    |--A. eburnus (Keyserling 1886) F11 [=Epeira eburna KK90]
    |--A. ejusmodi Bösenberg & Strand 1906 AM11
    |--‘Epeira’ enucleata Karsch 1879 K92
    |--‘Epeira’ ephippiata K92
    |--‘Epeira’ exanthematica Doleschall 1859 KK90
    |--A. exsertus MC13
    |--‘Aranea’ exusta Petrunkevitch 1963 P92
    |--‘Epeira’ fastidiosa Koch & Keyserling 1887 KK90
    |--‘Epeira’ ficta Rainbow 1896 R96
    |--‘Aranea’ flavissima Linnaeus 1758 L58
    |--‘Epeira’ furcifera Koch & Keyserling 1886 KK90
    |--‘Epeira’ geniculata D57
    |--A. gibbosus Walckenaer 1802 G76 [=Epeira gibbosa V09, Gibbaranea gibbosa G76]
    |--A. grossus Koch 1845 G76 [=Epeira grossa K01]
    |--‘Epeira’ heroine Koch 1871 KK90
    |--‘Aranea’ heteropthalma L02b
    |--A. himalayaensis Tikader 1975 TB81
    |--‘Epeira’ himalayana Sim. 1889 K01
    |--‘Epeira’ hirsutula Stoliczka 1869 S69
    |--‘Epeira’ hispida Doleschall 1859 D59
    |--‘Epeira’ idonea Koch & Keyserling 1887 KK90
    |--‘Epeira’ inconspicua K01
    |--‘Epeira’ indistincta Doleschall 1859 D59
    |--‘Epeira’ inquieta Koch & Keyserling 1887 KK90
    |--‘Epeira’ insularis K18
    |--A. iviei DC01
    |--‘Epeira’ junghuhnii Doleschall 1859 D59
    |--‘Epeira’ juniperi K18
    |--‘Epeira’ kuhlii Doleschall 1859 D59
    |--‘Epeira’ labyrinthea B06
    |--‘Epeira’ lancearia Koch & Keyserling 1887 KK90
    |--‘Aranea’ lapidaria L02b
    |--A. lathyrinus (Holmberg 1875) [=Metepeira lathyrina] AM11
    |--‘Aranea’ levipes Linnaeus 1758 L58
    |--‘Aranea’ lineata Linnaeus 1758 L58
    |--‘Epeira lineata’ Lucas 1846 non Aranea lineata Linnaeus 1758 E12
    |--‘Epeira’ lodicula Koch & Keyserling 1887 KK90
    |--A. lutulentus (Koch & Keyserling 1886) MC13, KK90 [=Epeira lutulenta KK90]
    |--‘Epeira’ malabarica D57
    |--‘Epeira’ mamillana Koch & Keyserling 1887 KK90
    |--‘Epeira’ manipa Doleschall 1857 D57
    |--‘Epeira’ margaritacea Doleschall 1859 D59
    |--A. marmoreus Clerck 1757 G76 [=Epeira marmorea K01]
    |--‘Epeira’ mediocris Kulczyński 1901 K01
    |--A. miniatus DC01
    |--A. mitificus (Simon 1886) AM11 [=Epeira mitifica TB81; incl. Zilla nawazi Dyal 1935 TB81]
    |--‘Epeira’ moluccensis Doleschall 1857 D57
    |--A. mongolicus Simon 1895 [=Araneus (Epeira) mongolicus] S95
    |--‘Epeira’ mulieraria Koch & Keyseling 1887 KK90
    |--‘Epeira’ necopina Koch & Keyserling 1887 KK90
    |--‘Aranea’ nidulans L99
    |--‘Epeira’ nigropunctata KK90
    |--‘Epeira’ nigrotarsalis Doleschall 1859 D59
    |--A. nordmanni Thorell 1856 G76
    |--‘Aranea’ notata Linnaeus 1758 L58
    |--A. nympha Simon 1889 TB81
    |--‘Epeira’ obtusata Karsch 1892 K92
    |--‘Aranea’ ocellata Linnaeus 1758 L58
    |--A. omoeda Thorell 1856 [=Gibbaranea omoeda] G76
    |--‘Epeira’ orichalcea Doleschall 1857 D57
    |--‘Epeira’ packardii Thorell 1875 K01
    |--A. pahalgaonensis Tikader & Bal 1981 TB81
    |--A. panchganiensis Tikader & Bal 1981 TB81
    |--‘Epeira’ paradoxa Lucas 1846 E12
    |--A. pegnia DC01
    |--‘Aranea’ perita Latreille 1799 L99
    |--‘Epeira’ phthisica KK90
    |--A. potanini Simon 1895 [=Araneus (Epeira) potanini] S95
    |--A. pratensis DC01
    |--‘Epeira’ proxima K01
    |--‘Epeira’ psittacina Koch in Koch & Keyserling 1887 KK90
    |--‘Epeira’ pyramidata K92
    |--A. quadratus Clerck 1757 G76 [=Epeira quadrata K01]
    |--‘Aranea’ quadripunctata Linnaeus 1758 L58
    |--‘Epeira’ quaesita Koch & Keyserling 1887 KK90
    |--‘Epeira’ queribunda Koch & Keyserling 1887 KK90
    |--‘Epeira’ quieta Koch & Keyserling 1887 KK90
    |--‘Epeira’ rabiosula Koch & Keyserling 1887 KK90
    |--‘Epeira’ radja Doleschall 1857 D57
    |--‘Epeira’ ramulosa Koch & Keyserling 1887 KK90
    |--‘Epeira’ rara Koch & Keyserling 1887 KK90
    |--‘Aranea’ raui Strand 1907 J98
    |--‘Aranea’ recherchensis Main 1954 M54
    |--A. redii (Scopoli 1763) R14, G76 [=Aranea redii G76, Agalenatea redii G76]
    |--‘Aranea’ redimita Linnaeus 1758 L58
    |--‘Aranea’ relucens L02a
    |--‘Aranea’ reticulata Linnaeus 1758 L58
    |--‘Epeira’ rhodosternon Doleschall 1859 D59
    |--A. rhombocephalus MC13
    |--‘Aranea’ riparia Linnaeus 1758 L58
    |--‘Epeira’ rotundula Koch in Koch & Keyserling 1887 KK90
    |--‘Epeira’ rubicundula Koch & Keyserling 1887 KK90
    |--‘Aranea’ rufipes Linnaeus 1758 L58
    |--‘Aranea’ rupestris Linnaeus 1758 L58
    |--A. saevus Koch 1872 G76 [=Epeira saeva K01]
    |--‘Epeira’ scutifera Koch & Keyserling 1886 KK90
    |--‘Epeira’ sericea S69
    |--‘Aranea’ sexpunctata Linnaeus 1758 L58
    |--‘Epeira’ similaris Rainbow 1896 R96
    |--‘Epeira’ solers K18
    |--‘Epeira’ solitaria Emerton 1884 K01
    |--‘Epeira’ spectabilis Doleschall 1859 D59
    |--‘Aranea’ spinosa Linnaeus 1758 L58
    |--‘Epeira’ squamifera Koch & Keyserling 1886 KK90
    |--‘Epeira’ stigmatisata Karsch 1878 K92
    |--‘Epeira’ stolida Koch & Keyserling 1887 KK90
    |--‘Epeira’ strix K18
    |--A. sturmi V09
    |--A. sublutia (Urquhart 1892) [=Epeira sublutia] NS00
    |--‘Epeira’ suspicax Cambridge 1876 K01
    |--‘Epeira’ sydneyica Koch & Keyserling 1887 KK90
    |--‘Epeira’ tabida KK90
    |--‘Epeira’ talipedata Koch & Keyserling 1887 KK90
    |--A. tartaricus [=A. (Epeira) tartaricus] S95
    |--‘Epeira’ theodori Thorell 1894 [=E. (Singa) theodori] T94
    |--‘Epeira’ thomisoides Duf. 1855 K92
    |--A. transmarinus (Keyserling 1865) G84, KK90 [=Epeira transmarina KK90; incl. E. producta Koch 1867 KK90]
    |--A. trifolium DC01
    |--‘Epeira’ trituberculata Lucas 1846 E12
    |--‘Aranea’ truncorum Linnaeus 1758 L58
    |--A. ullrichi (Hahn 1834) [=Epeira ullrichii, Gibbaranea ullrichii] G76
    |--A. undatus V09
    |--‘Epeira’ unicolor Doleschall 1857 D57
    |--‘Epeira’ urbana Koch & Keyserling 1887 KK90
    |--‘Epeira’ usualis Koch & Keyserling 1887 KK90
    |--‘Epeira’ vallata Koch & Keyserling 1886 KK90
    |--A. ventricosus (Koch 1878) AM11 (see below for synonymy)
    |--A. verrucosus (Walckenaer 1841) [=Epeira verrucosa] S99
    |--‘Aranea’ viatica Linnaeus 1758 L58
    |--‘Epeira’ victoria [incl. E. victoria var. orientalis Kulczyński 1901 non E. orientalis Urquhart 1887] K01
    |--‘Epeira’ viriditus M54
    |--‘Epeira’ vulgaris K18
    |--‘Epeira’ wagneri Rainbow 1896 R96
    |--‘Epeira’ westringii K01
    `--‘Epeira’ wittfeldae B06

Araneus Clerck 1758 F11 [incl. Agalenatea Archer 1951 G76, Aranea Linnaeus 1758 G76, Burgessia McCook 1894 TB81, Conepeira Archer 1951 TB81, Epeira Latreille 1810 G76, Gibbaranea Archer 1951 G76, Neopora Simon 1864 TB81]

Araneus bituberculatus (Walckenaer 1802) TB81 [=Aranea bituberculata TB81, *Gibbaranea bituberculata G76; incl. Aranea alboarcuata Panzer 1804 TB81, Epeira bicornis Menge 1866 TB81, E. dromedaria Koch 1845 TB81, Araneus dromedarius TB81]

Araneus diadematus Clerck 1757 G76 [=Araneus (Epeira) diadematus S95, Aranea diademata C01, Epeira diademata K01; incl. *Aranea diadema Linnaeus 1758 G76, Araneus diadema AM11, *Epeira diadema G76]

Araneus ventricosus (Koch 1878) AM11 [=Aranea ventricosus AM11, Epeira ventricosa K01; incl. E. senta Karsch 1879 K01, Araneus (Epeira) sentus S95]

*Type species of generic name indicated

REFERENCES

[AM11] Araujo, D., V. F. Mattos, A. M. Giroti, M. G. Kraeski, L. S. Carvalho & A. D. Brescovit. 2011. Cytogenetical characterization of six orb-weaver species and review of cytogenetical data for Araneidae. Journal of Arachnology 39 (2): 337–344.

[B06] Banks, N. 1906. Arachnida from the Bahamas. Bulletin of the American Museum of Natural History 22: 185–189.

[B11] Blackledge, T. A. 2011. Prey capture in orb weaving spiders: are we using the best metric? Journal of Arachnology 39 (2): 205–210.

[C01] Cambridge, F. O. P. 1901. A revision of the genera of the Araneae or spiders with reference to their type species. Annals and Magazine of Natural History, series 7, 7: 51–65.

[DC01] Davis, M. J., & F. A. Coyle. 2001. Habitat distribution and life history of Araneus spider species in the Great Smoky Mountains National Park. Bulletin of the British Arachnological Society 12 (2): 49–57.

[D57] Doleschall, C. L. 1857. Bijdrage tot de kennis der Arachniden van den Indischen Archipel. Natuurkundig Tijdschrift voor Nederlandsch Indië, series 3, 3 (5–6): 399–434, pls 1–2.

[D59] Doleschall, C. L. 1859. Tweede Bijdrage tot de kennis der Arachniden van den Indischen Archipel. Verhandelingen der Natuurkundige Vereeniging in Nederlandsch Indie [Acta Societatis Scientiarum Indo-Neêrlandicae] 5 (5): 1–60, pls 1–18.

[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.

[F11] Framenau, V. W. 2011. Lariniophora, a new monotypic orb-weaving spider genus from Australia (Araneae: Araneidae: Araneinae). Records of the Western Australian Museum 26 (2): 191–201.

[G84] Gauld, I. D. 1984. An Introduction to the Ichneumonidae of Australia. British Museum (Natural History).

[G76] Grasshoff, M. 1976. Zur Taxonomie und Nomenklatur mitteleuropäischer Radnetzspinnen der Familie Araneidae (Arachnida: Araneae). Senckenbergiana Biologica 57 (1–3): 143–154.

[HU08] Hore, U., & V. P. Uniyal. 2008. Diversity and composition of spider assemblages in five vegetation types of the Terai Conservation Area, India. Journal of Arachnology 36 (2): 251–258.

[J98] Jäger, P. 1998. Das Typenmaterial der Spinnentiere (Arachnida: Acari, Amblypygi, Araneae, Opiliones, Pseudoscorpiones, Scorpiones, Uropygi) aus dem Museum Wiesbaden. Jahrbuecher des Nassauischen Vereins fuer Naturkunde 119: 81–91.

[K92] Karsch, F. 1892. Arachniden von Ceylon und von Minikoy gesammelt von den Herren Doctoren P. und F. Sarasin. Berliner Entomologische Zeitschrift 36 (2): 267–310.

[KK90] Koch, L., & E. Keyserling. 1884–1890. Die Arachniden Australiens nach der Natur beschrieben und abgebildet vol. 2. Bauer & Raspe: Nürnberg.

[K18] Kohl, F. F. 1918. Die Hautflügergruppe "Sphecinae". IV. Teil. Die natürliche Gattung Sceliphron Klug (Pelopoeus Latr.). Annalen des Naturhistorischen Museum in Wien 32: 1–171.

[K01] Kulczyński, V. 1901. Arachnoideák [Arachnoidea]. In: Horváth, G. (ed.) Zichy Jenő Gróf Harmadik Ázsiai Utazása [Dritte Asiatische Forschungsreise des Grafen Eugen Zichy] vol. 2. Zichy Jenő Gróf Harmadik Ázsiai Utazásának Állattani Eredményei [Zoologische Ergebnisse der Dritten Asiatischen Forschungsreise des Grafen Eugen Zichy] pp. 311–369. Victor Hornyánszky: Budapest, and Karl W. Hierseman: Leipzig.

[L99] Latreille, P. A. 1799. Extrait d'un mémoire sur la famille des araignées mineuses. Bulletin des Sciences, par la Societé Philomathique de Paris 1 (22): 169.

[L02a] Latreille, P. A. 1802a. Histoire Naturelle des Fourmis, et recueil de mémoires et d'observations sur les abeilles, les araignées, les faucheurs, et autres insectes. Théophile Barrois père: Paris.

[L02b] Latreille, P. A. 1802b. Histoire Naturelle, générale et particulière des crustacés et des insectes vol. 3. Familles naturelles des genres. F. Dufart: Paris.

[L58] Linnaeus, C. 1758. Systema Naturae per Regna Tria Naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata. Laurentii Salvii: Holmiae.

[M54] Main, B. Y. 1954. Spiders and Opiliones. In: Willis, J. H. (ed.) The Archipelago of the Recherche pp. 37–53. Australian Geographical Society: Melbourne.

[MC13] Majer, J. D., S. K. Callan, K. Edwards, N. R. Gunawardene & C. K. Taylor. 2013. Baseline survey of the terrestrial invertebrate fauna of Barrow Island. Records of the Western Australian Museum Supplement 83: 13–112.

[NS00] Nicholls, D. C., P. J. Sirvid, S. D. Pollard & M. Walker. 2000. A list of arachnid primary types held in Canterbury Museum. Records of the Canterbury Museum 14: 37–48.

[PVD10] Paquin, P., C. J. Vink & N. Dupérré. 2010. Spiders of New Zealand: annotated family key and species list. Manaaki Whenua Press: Lincoln (New Zealand).

[P92] Poinar, G. O., Jr. 1992. Life in Amber. Stanford University Press: Stanford.

[R96] Rainbow, W. J. 1896. Descriptions of some new Araneidae of New South Wales. No. 6. Proceedings of the Linnean Society of New South Wales 21 (3): 320–344, pls 18–20.

[R97] Rainbow, W. J. 1897. Descriptions of some new Araneidae of New South Wales, No. 7. Proceedings of the Linnean Society of New South Wales 21 (4): 628–633, pl. 49.

[R13] Rainbow, W. J. 1913. Arachnida from the Solomon Islands. Records of the Australian Museum 10 (1): 1–16.

[R14] Ramírez, M. J. 2014. The morphology and phylogeny of dionychan spiders (Araneae: Araneomorphae). Bulletin of the American Museum of Natural History 390: 1–374.

[S95] Simon, E. 1895. Arachnides recueillis par Mr G. Potanine en Chine et en Mongolie (1876–1879). Izviestĭia Imperatorskoi Akademĭi Nauk” (Bulletin de l’Academie Imperiale des Sciences de St. Pétersbourg), ser. 5, 4: 331–345.

[S99] Simon, E. 1899. Ergebnisse einer Reise nach dem Pacific (Schauinsland 1896–1897). Zoologische Jahrbücher, Abteilung für Systematik, Geographie un Biologie der Thiere 12 (4): 411–437.

[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.

[T94] Thorell, T. 1894. Förteckning öfver Arachnider från Java och närgränsande öar, insamlade af docenten D:r Carl Aurivillius; jemte beskrifingar å några sydasiatiska och sydamerikanska Spindlar. Bihang till K. Svenska Vetenskaps-Akademiens Handlingar 20 pt 4 (4): 1–63.

[TB81] Tikader, B. K., & A. Bal. 1981. Studies on some orb-weaving spiders of the genera Neoscona Simon and Araneus Clerck of the family Araneidae (=Argiopidae) from India. Records of the Zoological Survey of India, Miscellaneous Publication, Occasional Paper 24: 1–60.

[V09] Verdcourt, B. (ed.) 2009. Additions to the Wild Fauna and Flora of the Royal Botanic Gardens, Kew. XXVI. Miscellaneous records. Kew Bulletin 64 (1): 183–194.

Last updated: 15 January 2022.

Cyrtophora

Tent-web spider Cyrtophora citricola, copyright Ed Nieuwenhuys.


Belongs within: Araneidae.

Cyrtophora is a mostly Indo-Australian genus of spiders (with some species in Africa and the New World) whose members construct a horizontal orbweb with a tightly woven mesh and an extremely tight, permanent non-sticky spiral (Framenau & Scharff 2009).

Characters (from Framenau & Scharff 2009): Lateral eyes slightly separated; posterior eye row recurved or straight; legs relatively heavy; fourth leg with combined patella and tibia slightly shorter than femur, and shorter than combined metatarsus and tarsus; male pedipalp with embolus placed near median apophysis, supported by a conductor, basal cymbium concavity absent; female epigyne lacking a scape.

<==Cyrtophora Simon 1864 [incl. Euetria Thorell 1890, Suzumia Nakatsudi 1943] FS09
    |--*C. citricola (Forsskål 1775) [=Aranea citricola; incl. C. sculptilis Koch 1872] FS09
    |--C. beccarii (Thorell 1878) FS09
    |--C. cicatrosa (Stoliczka 1869) FS09
    |--C. cordiformis (Koch 1871) FS09
    |--C. crassipes (Rainbow 1897) [=Epeira crassipes, Aranea crassipes, Araneus crassipes] FS09
    |--C. cylindroides (Walckenaer 1842) FS09 (see below for synonymy)
    |--C. exanthematica (Doleschall 1859) FS09
    |--C. feai (Thorell 1887) AM11
    |--C. gazellae (Karsch 1878) [=Epeira gazellae, Aranea gazellae, Araneus gazellae] FS09
    |--C. hirta Koch 1872 FS09
    |--C. moluccensis (Doleschall 1857) FS09
    |--C. monulfi Chrysanthus 1960 FS09
    |--C. parnasia Koch 1872 FS09
    |--C. rainbowi (Roewer 1955) [=Cyclosa rainbowi, Epeira pallida Rainbow 1897 non Walckenaer 1805, Cyc. pallida] FS09
    |--C. trigona (Koch 1871) [=Epeira trigona, Aranea trigona, Araneus trigonus] FS09
    `--C. unicolor (Doleschall 1857) FS09

Cyrtophora cylindroides (Walckenaer 1842) FS09 [=Epeira cylindroides R13; incl. E. nephilina Koch 1871 FS09, E. viridipes Doleschall 1859 FS09]

*Type species of generic name indicated

REFERENCES

[AM11] Araujo, D., V. F. Mattos, A. M. Giroti, M. G. Kraeski, L. S. Carvalho & A. D. Brescovit. 2011. Cytogenetical characterization of six orb-weaver species and review of cytogenetical data for Araneidae. Journal of Arachnology 39 (2): 337–344.

[FS09] Framenau, V. W. & N. Scharff. 2009. Cyrtobill darwini, a new species in a new orb-weaving spider genus from Australia (Araneae: Araneidae: Cyrtophorinae). Records of the Western Australian Museum 25 (3): 315–328.

[R13] Rainbow, W. J. 1913. Arachnida from the Solomon Islands. Records of the Australian Museum 10 (1): 1–16.

Argiope

Wasp spider Argiope bruennichi, copyright Cinnamologus.


Belongs within: Araneidae.

Argiope, St Andrew's cross spiders, is a genus of often large and colourfully striped orbweb spiders that derive their vernacular name from the construction by some species of webs with an X-shaped stabilimentum radiating from the centre. The effect is enhanced by the spider sitting with the front and rear pairs of legs each closely held and aligned with the arms of the stabilimentum. Conversely, the Palaearctic species A. bruennichi is known as the 'wasp spider' in reference to its bright yellow and black coloration.

Characters (from Doleschall 1859, as section Argyopes of genus Epeira): Usually furry, ornately coloured. Cephalothorax very flat, anteriorly narrow. Anterior median eyes nearer to each other than are the posterior median eyes. Abdomen ovoid, flat-topped, angular. Legs moderately prolonged, radially extended at rest.

<==Argiope Audouin 1826 [Argiopinae] PVD10
    |--*A. lobata (Pallas 1772) [=Aranea lobata] G76
    |--A. aemula K92
    |--A. aetherea (Walck. 1837) [=Epeira aetherea] R13
    |--A. amoena Koch 1878 B11
    |--A. anasuja Thor. 1887 K92
    |--A. appensa E95
    |--A. argentata (Fabricius 1775) GA11
    |--A. aurantia Lucas 1833 B11
    |--A. aurocincta E95
    |--A. australis JD-S07
    |--A. avara Thorell 1868 S99
    |--A. bruennichi (Scopoli 1772) G76
    |--A. catenulata (Doleschall 1859) K92, D59 [=Epeira catenulata K92]
    |--A. crenulata (Doleschall 1857) [=Epeira (Argyopes) crenulata] D57
    |--A. curvipes Koch & Keyserling 1886 KK90
    |--A. cuspidata E95
    |--A. fasciata (Hentz 1847) [=Epeira fasciata; incl. A. plana Koch 1867, A. transversa Emerton 1884] KK90
    |--A. flavipalpis E95
    |--A. florida E95
    |--A. katherina Levi 1983 GA11
    |--A. keyserlingi (Karsch 1878) GA11
    |--A. leopardina Pocock 1898 R13
    |--A. luzona (Walckenaer 1841) AM11
    |--A. mammillaris (Stoliczka 1869) [=Epeira (Argyopes) mammillaris] S69
    |--A. minuta Karsch 1879 AM11
    |--A. ocyaloides E95
    |--A. ornata Simon 1884 K92
    |--A. pelewensis Koch & Keyserling 1886 KK90
    |--A. picta (K. 1871) [incl. A. principalis K. 1871] R13
    |--A. plagiata Karsch 1892 K92
    |--A. protensa Koch 1872 PVD10
    |--A. pulchella Thorell 1881 AM11
    |--A. radon Levi 1983 GA11
    |--A. reinwardtii (Doleschall 1859) [=Epeira reinwardtii; incl. A. doleschallii Thorell 1873] T94
    |--A. savignyi Levi 1968 B11
    |--A. stellata (Stoliczka 1869) [=Epeira (Argyopes) stellata] S69
    |--A. striata Doleschall 1857) [=Epeira (Argyopes) striata] D57
    |--A. taprobanica Thor. 1887 K92
    |--A. trifasciata (Forskål 1775) SC97 [=Epeira (Argyopes) trifasciata D57]
    |--A. trivittata Karsch 1892 K92
    `--A. versicolor (Doleschall 1859) [=Epeira versicolor] K92

*Type species of generic name indicated

REFERENCES

[AM11] Araujo, D., V. F. Mattos, A. M. Giroti, M. G. Kraeski, L. S. Carvalho & A. D. Brescovit. 2011. Cytogenetical characterization of six orb-weaver species and review of cytogenetical data for Araneidae. Journal of Arachnology 39 (2): 337–344.

[B11] Blackledge, T. A. 2011. Prey capture in orb weaving spiders: are we using the best metric? Journal of Arachnology 39 (2): 205–210.

[D57] Doleschall, C. L. 1857. Bijdrage tot de kennis der Arachniden van den Indischen Archipel. Natuurkundig Tijdschrift voor Nederlandsch Indië, series 3, 3 (5–6): 399–434, pls 1–2.

[D59] Doleschall, C. L. 1859. Tweede Bijdrage tot de kennis der Arachniden van den Indischen Archipel. Verhandelingen der Natuurkundige Vereeniging in Nederlandsch Indie [Acta Societatis Scientiarum Indo-Neêrlandicae] 5 (5): 1–60, pls 1–18.

[E95] Elgar, M. A. 1995. The duration of copulation in spiders: comparative patterns. Records of the Western Australian Museum Supplement 52: 1–11.

[G76] Grasshoff, M. 1976. Zur Taxonomie und Nomenklatur mitteleuropäischer Radnetzspinnen der Familie Araneidae (Arachnida: Araneae). Senckenbergiana Biologica 57 (1–3): 143–154.

[GA11] Gregorič, M., I. Agnarsson, T. A. Blackledge & M. Kuntner. 2011. Darwin's bark spider: giant prey in giant orb webs (Caerostris darwini, Araneae: Araneidae)? Journal of Arachnology 39 (2): 287–295.

[JD-S07] Jocqué, R., & A. S. Dippenaar-Schoeman. 2007. Spider Families of the World. Royal Museum for Central Africa: Tervuren (Belgium).

[K92] Karsch, F. 1892. Arachniden von Ceylon und von Minikoy gesammelt von den Herren Doctoren P. und F. Sarasin. Berliner Entomologische Zeitschrift 36 (2): 267–310.

[KK90] Koch, L., & E. Keyserling. 1884–1890. Die Arachniden Australiens nach der Natur beschrieben und abgebildet vol. 2. Bauer & Raspe: Nürnberg.

[PVD10] Paquin, P., C. J. Vink & N. Dupérré. 2010. Spiders of New Zealand: annotated family key and species list. Manaaki Whenua Press: Lincoln (New Zealand).

[R13] Rainbow, W. J. 1913. Arachnida from the Solomon Islands. Records of the Australian Museum 10 (1): 1–16.

[SC97] Scharff, N., & J. A. Coddington. 1997. A phylogenetic analysis of the orb-weaving spider family Araneidae (Arachnida, Araneae). Zoological Journal of the Linnean Society 120: 355–434.

[S99] Simon, E. 1899. Ergebnisse einer Reise nach dem Pacific (Schauinsland 1896–1897. Zoologische Jahrbücher, Abteilung für Systematik, Geographie un Biologie der Thiere 12 (4): 411–437.

[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.

[T94] Thorell, T. 1894. Förteckning öfver Arachnider från Java och närgränsande öar, insamlade af docenten D:r Carl Aurivillius; jemte beskrifingar å några sydasiatiska och sydamerikanska Spindlar. Bihang till K. Svenska Vetenskaps-Akademiens Handlingar 20 pt 4 (4): 1–63.

Last updated: 15 January 2022.

Uropygi

Hypoctonus sp., copyright Bernard Dupont.


Belongs within: Arachnida.
Contains: Thelyphoninae.

The Uropygi, whip-scorpions, are a group of tropical arachnids with raptorial pedipalps and an elongate filamentous post-pygidium (Harvey 2002). An alternative vernacular name, 'vinegaroon', comes from their possession of anal glands from which they spray unpleasant-smelling chemicals at threats.

<==Uropygi [Holopeltidia, Oxopoei, Thelyphonida]
    |  i. s.: Etiennus africanus (Hentschel 1984) H02
    |         Proschizomus Dunlop & Horrocks 1996 H02
    |         Mesoproctus Dunlop 1998 H02
    |--Hypoctonidae [Hypoctoninae, Hypoctonini] RC73
    |    |--Thelyphonellus Pocock 1894 RC73
    |    |    `--*T. amazonicus (Butler 1872) RC73
    |    |--Labochirus Pocock 1894 RC73
    |    |    |--*L. proboscideus (Butler 1872) RC73
    |    |    |--L. cervinus Pocock 1900 RC73
    |    |    `--L. tauricornis Pocock 1900 RC73
    |    `--Hypoctonus Thorell 1889 RC73
    |         |--*H. formosus (Butler 1872) RC73 [=Thelyphonus formosus T89; incl. T. angustus Stoliczka 1873 T89]
    |         |--H. africanus Hentschel 1899 RC73
    |         |--H. andersoni (Oates 1890) RC73
    |         |--H. binghami (Oates 1890) RC73
    |         |--H. birmanicus Hirst 1911 RC73
    |         |--H. browni Gravely 1912 RC73
    |         |--H. carmichaeli Gravely 1916 RC73
    |         |--H. clarki Cooke & Shadab 1973 RC73
    |         |--H. dawnae Gravely 1912 RC73
    |         |--H. ellisii Gravely 1912 RC73
    |         |--H. gastrostictus Kraepelin 1897 RC73
    |         |--H. granosus Pocock 1900 RC73
    |         |--H. kraepelini Simon 1901 RC73
    |         |--H. oatesi Pocock 1900 RC73
    |         |--H. proboscideus (Butler 1872) [=Thelyphonus proboscideus; incl. T. parvimanus Butler 1873] K92
    |         |--H. rangunensis (Oates 1890) RC73
    |         |--H. saxatilis (Oates 1890) RC73
    |         |--H. silvaticus (Oates 1890) RC73
    |         |--H. stoliczkae Gravely 1912 RC73
    |         `--H. woodmasoni (Oates 1890) RC73
    `--Thelyphonidae [Telyphonidae, Thelyphonoidae] H04b
         |  i. s.: Prothelyphonus Fritsch 1904 F04
         |           |--P. bohemicus (Kušta 1884) [=Thelyphonus bohemicus, Geralinura bohemica] F04
         |           `--P. cordai Fritsch 1904 F04
         |         Geralinura Scudder 1884 F04, H02
         |           |--G. crassa Kušta 1888 F04
         |           |--G. noctua Kušta 1888 F04
         |           `--G. scudderi Kušta 1888 F04
         |--Thelyphoninae RC73
         |--Typopeltis Pocock 1894 [incl. Gipopeltis Speijer 1934; Typopeltinae] RC73
         |    |--*T. crucifer Pocock 1900 RC73
         |    |--T. amurensis (Tarnani 1889) RC73
         |    |--T. cantonensis Speijer 1936 [incl. Mastigoproctus transoceanicus Lazell 2000] H04b
         |    |--T. dalyi Pocock 1900 RC73
         |    |--T. harmandi Kraepelin 1901 [=*Gipopeltis harmandi] RC73
         |    |--T. kamahouii Tarnani 1900 RC73
         |    |--T. kasnakowi Tarnani 1900 H04a
         |    |--T. magnificus Haupt 2004 H04a
         |    |--T. niger (Tarnani 1894) H04b
         |    |--T. sinensis Butler 1872 H04b
         |    |--T. stimpsoni (Wood 1862) RC73
         |    `--T. tarnanii Pocock 1902 RC73
         `--Mastigoproctinae [Mastigoproctidae, Uroproctinae] RC73
              |--Teltus Speijer 1936 RC73
              |    `--*T. vanoorti Speijer 1936 RC73
              |--Uroproctus Pocock 1894 RC73
              |    `--*U. assamensis (Stoliczka 1869) RC73
              |--Amauromastigon Mello-Leitão 1931 RC73
              |    `--*A. annectens (Werner 1916) RC73 [=Mastigoproctus annectens J98]
              |--Mimoscorpius Pocock 1894 RC73
              |    `--*M. pugnator (Butler 1872) RC73
              `--Mastigoproctus Pocock 1894 RC73
                   |--*M. giganteus (Lucas 1835) RC73
                   |--M. baracoensis Franganillo 1931 RC73
                   |--M. brasilianus (Koch 1843) H04b
                   |--M. butleri Pocock 1894 RC73
                   |--M. colombianus Mello-Leitao 1940 RC73
                   |--M. formidabilis Hirst 1912 RC73
                   |--M. liochirus Pocock 1902 RC73
                   |--M. maximus (Tarnani 1889) RC73
                   |--M. minensis Mello-Leitao 1931 RC73
                   |--M. perditus Mello-Leitao 1931 RC73
                   |--M. proscorpio (Latreille 1806) RC73
                   `--M. tantalus Roewer 1954 R54 RC73

*Type species of generic name indicated

REFERENCES

[F04] Fritsch, A. 1904. Palaeozoische Arachniden. Selestverlag: Prague.

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

[H04a] Haupt, J. 2004a. A new species of whipscorpion from Laos (Arachnida: Uropygi: Thelyphonidae). Senckenbergiana Biologica 83 (2): 151–155.

[H04b] Haupt, J. 2004b. Mastigoproctus transoceanicus a synonym of Typopeltis cantonensis (Arachnida, Uropygi, Thelyphonidae). Senckenbergiana Biologica 83 (2): 157–162.

[J98] Jäger, P. 1998. Das Typenmaterial der Spinnentiere (Arachnida: Acari, Amblypygi, Araneae, Opiliones, Pseudoscorpiones, Scorpiones, Uropygi) aus dem Museum Wiesbaden. Jahrbuecher des Nassauischen Vereins fuer Naturkunde 119: 81–91.

[K92] Karsch, F. 1892. Arachniden von Ceylon und von Minikoy gesammelt von den Herren Doctoren P. und F. Sarasin. Berliner Entomologische Zeitschrift 36 (2): 267–310.

[RC73] Rowland, J. M., & J. A. L. Cooke. 1973. Systematics of the arachnid order Uropygida (=Thelyphonida). Journal of Arachnology 1: 55–71.

[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.

Amblypygi

Charinus cf. africanus, copyright Arthur Anker.


Belongs within: Arachnida.
Contains: Neoamblypygi.

The Amblypygi, whip-spiders, are a group of flattened arachnids with raptorial pedipalps and extremely long, filamentous front legs that are used as tactile organs (Harvey 2002).

<==Amblypygi
    |  i. s.: Admetus Koch 1850 GE02, BR05
    |           |--A. pomilio GE02
    |           `--A. whitei B06
    |         Hemiphrynus viridiceps B06
    |--Paracharon [Paleoamblypygi, Paracharontidae] W99
    |    `--P. caecus Hansen 1921 W99
    `--Euamblypygi [Phrynoidae] W99
         |--Charinidae W99
         |    |--Catageus Thorell 1889 W99, H02
         |    |    `--*C. pusillus Thorell 1889 T89
         |    |--+--Phrynichosarax W99
         |    |  `--Sarax Simon 1892 W99, H02
         |    |       |--S. mediterraneus Delle Cave 1986 W05
         |    |       `--S. sarawakensis (Thorell 1888) W99
         |    `--Charinus Simon 1892 W05 (see below for synonymy)
         |         |--*C. australianus (Koch 1867) [=Phrynus australianus] VGF14
         |         |--C. abbatei Delle Cave 1986 W05
         |         |--C. acaraje Pinto-da-Rocha et al. 2002 VGF14
         |         |--C. acosta (Quintero 1983) W05
         |         |--C. africanus Hansen 1921 W99
         |         |--C. asturius Pinto-da-Rocha et al. 2002 VGF14
         |         |--C. bengalensis (Gravely 1911) [=*Charinides bengalensis] W05
         |         |--C. brasilianus Weygoldt 1972 W05
         |         |--C. dhofarensis Weygoldt, Pohl & Polak 2002 W05
         |         |--C. eleonorae Baptista & Giupponi 2003 VGF14
         |         |--C. ioanniticus (Kritscher 1959) W05 [=*Lindosiella ioannitica El-H02]
         |         |--C. jibaossu Vasconcelos, Giupponi & Ferreira 2014 VGF14
         |         |--C. koepckei Weygoldt 1972 W05
         |         |--C. madagascariensis Fage 1954 W05
         |         |--C. milloti Fage 1939 W05
         |         |--C. montanus Weygoldt 1972 W99
         |         |--C. mysticus Giupponi & Kury 2002 VGF14
         |         |--C. neocaledonicus Kraepelin 1895 W05
         |         |--C. pakistanus Weygoldt 2005 W05
         |         |--C. pescotti Dunn 1949 W05
         |         |--C. potiguar Vasconcelos et al. 2013 VGF14
         |         |--C. schirchii (Mello-Leitão 1931) (n. d.) VGF14
         |         |--C. seychellarum Kraepelin 1898 W05
         |         |--C. socotranus Weygoldt, Pohl & Polak 2002 W05
         |         |--C. stygochthobius Weygoldt & Van Damme 2004 W05
         |         |--C. troglobius Baptista & Giupponi 2002 VGF14
         |         `--C. vulgaris Miranda & Giupponi 2011 VGF14
         `--Neoamblypygi W99

Charinus Simon 1892 W05 [incl. Charinides Gravely 1911 W05, Lindosiella Kritscher 1959 El-H02, Tricharinus Quintero 1986 W05]

*Type species of generic name indicated

REFERENCES

[B06] Banks, N. 1906. Arachnida from the Bahamas. Bulletin of the American Museum of Natural History 22: 185–189.

[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.

[El-H02] El-Hennawy, H. K. 2002. The first record of Amblypygi from Egypt. Journal of Arachnology 30 (2): 452–453.

[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.

[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.

[VGF14] Vasconcelos, A. C. O., A. P. L. Giupponi & R. L. Ferreira. 2014. A new species of Charinus from Minas Gerais State, Brazil, with comments on its sexual dimorphism (Arachnida: Amblypygi: Charinidae). Journal of Arachnology 42 (2): 155–162.

[W99] Weygoldt, P. 1999. Spermatophores and the evolution of female genitalia in whip spiders (Chelicerata, Amblypygi). Journal of Arachnology 27: 103–116.

[W05] Weygoldt, P. 2005. Biogeography, systematic position, and reproduction of Charinus ioanniticus (Kritscher 1959) with the description of a new species from Pakistan (Chelicerata, Amblypygi, Charinidae). Senckenbergiana Biologica 85 (1): 43–56.

Leanchoiliida

Reconstruction of Leanchoilia, copyright Marianne Collins.


Belongs within: Euarthropoda.

The Leanchoiliida are a group of Cambrian 'great-appendage' arthropods that bore elongate flagella on the podomeres of the great appendage.

Synapomorphies (from Cotton & Braddy 2004): Head with four segments; distal spines on terminal podomeres of endopods of second segment appendages longer than entire podomere series, terminating in annulate flagellae.

<==Leanchoiliida [Alalcomenaeidea, Leanchoiliidacea, Leanchoiliidea]
    |--Alalcomenaeus Simonetta 1970 LSE13, CB04 [Alalcomenaeida]
    |    |--A. cambricus Simonetta 1970 CB04
    |    `--A. illecebrosus (Hou 1987) CB04
    `--+--Actaeus Simonetta 1970 LSE13, CB04 [Actaeida]
       `--+--Oestokerkus LSE13
          `--Leanchoilia Walcott 1912 CB04, B95
               |  i. s.: L. hanceyi CB04
               |--L. persephone LSE13
               `--+--L. illecebrosa LSE13
                  `--L. superlata Walcott 1912 LSE13, CB04

*Type species of generic name indicated

REFERENCES

[B95] Bousfield, E. L. 1995. A contribution to the natural classification of Lower and Middle Cambrian arthropods: Food-gathering and feeding mechanisms. Amphipacifica 2: 3–34.

[CB04] Cotton, T. J., & S. J. Braddy. 2004. The phylogeny of arachnomorph arthropods and the origin of the Chelicerata. Transactions of the Royal Society of Edinburgh: Earth Sciences 94: 169–193.

[LSE13] Legg, D. A., M. D. Sutton & G. D. Edgecombe. 2013. Arthropod fossil data increase congruence of morphological and molecular phylogenies. Nature Communications 4 (2485): 1–7.

Thelyphoninae

Thelyphonus sp., copyright Arthur Anker.


Belongs within: Uropygi.

The Thelyphoninae is a group of whipscorpions found in south-east Asia and Melanesia (Rowland & Cooke 1973).

Characters (from Rowland & Cooke 1973): Tarsal segments of first leg of female usually modified (except in Minbosius manilanus and Abaliella rohdei), variously incrassate or apophysate; pedipalpal patellar apophysis of male unmodified, occasionally longer than in female; abdominal sternites II and III modified; two, four or no ommatoids present; keel present between lateral and median eyes; posterior abdominal tergites divided.

<==Thelyphoninae [Thelyphonini]
    |--Minbosius Speijer 1936 RC73
    |    `--*M. manilanus (Koch 1843) RC73
    |--Chajnus Speijer 1936 RC73
    |    `--*C. renschi Speijer 1936 RC73
    |--Glyptogluteus Rowland 1973 RC73
    |    `--*G. augustus Rowland 1973 RC73
    |--Ginosigma Speijer 1936 RC73
    |    |--*G. schimkewitschi (Tarnani 1894) RC73
    |    `--G. lombokensis Speijer 1936 RC73
    |--Tetrabalius Thorell 1888 RC73, H02
    |    |--*T. seticauda (Doleschall 1857) RC73
    |    `--T. nasutus Thorell 1889 RC73
    |--Abaliella Strand 1928 [=Abalius Kraepelin 1897 (preoc.)] RC73
    |    |--*A. rohdei (Kraepelin 1897) RC73
    |    |--A. dicranotarsalis Rowland 1973 RC73
    |    |--A. gertschi Rowland 1973 RC73
    |    |--A. manilana (Kraepelin 1901) RC73
    |    |--A. samoana (Kraepelin 1879) RC73
    |    `--A. willeyi (Pocock 1898) RC73
    `--Thelyphonus Latreille 1802 H02 [=Telyphonus (l. c.) KK84]
         |--*T. caudatus (Linnaeus 1758) RC73 [=Phalangium caudatum H02; incl. T. thorellii Tarnani 1890 T94]
         |--T. anthracinus Pocock 1894 RC73
         |--T. antillanus T94
         |--T. asperatus Thorell 1888 RC73
         |--T. assamensis Stoliczka 1869 [=Telyphonus assamensis] S69
         |--T. billitonensis Speijer 1931 RC73
         |--T. borneensis Kraepelin 1897 RC73
         |--T. burchardi Kraepelin 1912 RC73
         |--T. celebensis Kraepelin 1897 RC73
         |--T. doriae Thorell 1888 RC73
         |--T. grandis Speijer 1931 RC73
         |--T. hanseni Kraepelin 1897 RC73
         |--T. ignotus T94
         |--T. insulanus Koch & Keyserling 1884 RC73, KK84
         |--T. javanus T94
         |--T. kinabaluensis Speijer 1933 RC73
         |--T. klugi Kraepelin 1897 RC73
         |--T. lawrencei Rowland 1973 RC73
         |--T. leucurus Pocock 1898 RC73
         |--T. linganus Koch 1843 RC73
         |--T. pococki Tarnani 1901 RC73
         |--T. proscorpius T94
         |--T. rufipes [=Telyphonus rufipes] D57
         |--T. schnehageni Kraepelin 1897 RC73
         |--T. semperi Kraepelin 1897 RC73
         |--T. sepiaris Butler 1873 RC73 [incl. T. nigrescens Butler 1873 T89]
         |--T. seticauda Doleschall 1857 [=Telyphonus seticauda] D57
         |--T. spinimanus [=Telyphonus spinimanus] S69
         |--T. stimpsonii Wood 1861 [=Telyphonus stimpsonii] S69
         |--T. sucki Kraepelin 1897 RC73
         |--T. sumatranus Kraepelin 1897 RC73
         |--T. vanoorti Speijer 1936 RC73
         `--T. wayi Pocock 1900 RC73

*Type species of generic name indicated

REFERENCES

[D57] Doleschall, C. L. 1857. Bijdrage tot de kennis der Arachniden van den Indischen Archipel. Natuurkundig Tijdschrift voor Nederlandsch Indië, series 3, 3 (5–6): 399–434, plates 1–2.

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

[KK84] Koch, L., & E. Keyserling. 1884-1890. Die Arachniden Australiens nach der Natur beschrieben und abgebildet vol. 2. Bauer & Raspe: Nürnberg.

[RC73] Rowland, J. M., & J. A. L. Cooke. 1973. Systematics of the arachnid order Uropygida (=Thelyphonida). Journal of Arachnology 1: 55–71.

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

[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.

[T94] Thorell, T. 1894. Förteckning öfver Arachnider från Java och närgränsande öar, insamlade af docenten D:r Carl Aurivillius; jemte beskrifingar å några sydasiatiska och sydamerikanska Spindlar. Bihang till K. Svenska Vetenskaps-Akademiens Handlingar 20 pt. 4 (4): 1–63.