A Phylogeny and Classification of the Senticaudata subord. nov (Crustacea: Amphipoda)

Abstract

The Amphipoda includes a large clade defined by the presence of a previously unrecognised synapomorphy, apical robust setae on the rami of uropods 1-2. We term this clade the Senticaudata subord. nov. (Latin: sentis = thorn). It includes almost all freshwater species as well as a number of marine benthic taxa, formerly part of the 'Gammaridea'. The phylog-eny of the senticaudates was determined by cladistic analysis of morphological characters and character states. Within the suborder Senticaudata there are six infraorders: Carangoliopsida, Talitrida, Hadziida, Corophiida, Bogidiellida and Gam-marida. A classification is provided and all the senticaudate families are diagnosed. We introduce for the first time in am-phipod classification, the level parvorder between infraorder and superfamily. Four new families are described: Kairosidae; Eriopisidae; Nuuanuidae and Kergueleniolidae.

Full text

Accepted by S. Gerken: 27 Feb. 2017; published: 11 May 2017
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ZOOTAXA
A Phylogeny and Classification of the Amphipoda with the establishment of
the new order Ingolfiellida (Crustacea: Peracarida)
J.K. LOWRY
1
& A.A. MYERS
2
1
Australian Museum Research Institute, 1 William Street, Sydney, NSW 2010, Australia. E-mail: stephonyx@gmail.com
2
School of Biological, Earth and Environmental Sciences, University College Cork, Cork Enterprise Centre, Distillery Fields,
North Mall, Cork, Ireland. E-mail: bavayia@gmail.com
Magnolia Press
Auckland, New Zealand
4265

LOWRY & MYERS
2
·
Zootaxa 4265 (1) © 2017 Magnolia Press
J.K. LOWRY & A.A. MYERS
A Phylogeny and Classification of the Amphipoda with the establishment of the new order Ingolfiellida
(Crustacea: Peracarida)
(Zootaxa 4265)
89 pp.; 30 cm.
11 May 2017
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3
AMPHIPODA PHYL OGENY AND CLASSIFICATION
Table of contents
Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .7
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
Materials and methods. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
Characters used in the analysis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
Phylogenetic analysis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
Tree description . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
Tree discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .32
Classification . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
Superorder Peracarida . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
Order Ingolfiellida Hansen, 1903 stat. nov.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
Suborder Ingolfiellidea Hansen, 1903 (Ruffo, 1970) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
Infraorder Ingolfiellidamorpha Hansen, 1903 stat. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
Parvorder Ingolfiellidira Hansen, 1903 stat. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
Superfamily Ingolfielloidea Hansen, 1903 stat. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
Family Ingolfiellidae Hansen, 1903 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
Parvorder Metaingolfiellidira Ruffo, 1969 stat. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .37
Superfamily Metaingolfielloidea Ruffo, 1969 stat. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
Family Metaingolfiellidae Ruffo, 1969 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
Order Amphipoda Latreille, 1816 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
Suborder Pseudingolfiellidea Lowry & Myers, 2012a stat. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
Infraorder Pseudingolfiellida Lowry & Myers, 2012a stat. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
Parvorder Pseudingolfiellidira Lowry & Myers, 2012a stat. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
Superfamily Pseudingolfielloidea Lowry & Myers, 2012a stat. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
Family Pseudingolfiellidae Lowry & Myers, 2012a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
Suborder Hyperiidea H. Milne Edwards, 1830 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
Infraorder Physosomata Pirlot, 1929 (Bowman & Gruner, 1973) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
Parvorder Physosomatidira Pirlot, 1929 stat. nov.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
Superfamily Lanceoloidea Bovallius, 1887 (Bowman & Gruner, 1973) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
Family Chuneolidae Woltereck, 1909 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
Family Lanceolidae Bovallius, 1887 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
Family Megalanceolidae Zeidler, 2009 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
Family Metalanceolidae Zeidler, 2009 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
Family Microphasmidae Stephensen & Pirlot, 1931 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .39
Family Mimonecteolidae Zeidler, 2009. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
Family Prolanceolidae Zeidler, 2009 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40
Superfamily Scinoidea Stebbing, 1888 (Bowman & Gruner, 1973) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40
Family Archaeoscinidae Stebbing, 1904. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40
Family Microscinidae Zeidler, 2012 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40
Family Mimonectidae Bovallius, 1885 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40
Family Mimoscinidae Zeidler, 2012 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40
Family Scinidae Stebbing, 1888 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40
Infraorder Physocephalata Bowman & Gruner, 1973 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40
Parvorder Physocephalatidira Bowman & Gruner, 1973 stat. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41
Superfamily Vibilioidea Dana, 1852 (Bowman & Gruner, 1973) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41
Family Cyllopodidae Bovallius, 1887. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41
Family Paraphronimidae Bovallius, 1887 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41
Family Vibiliidae Dana, 1852 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41
Superfamily Phronimoidea Dana, 1852 (Bowman & Gruner, 1973) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41
Family Bougisidae Zeidler, 2004 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41
Family Cystisomatidae Willemöes-Suhm, 1875 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42
Family Dairellidae Bovallius, 1887 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42
Family Hyperiidae Dana, 1852 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42
Family Iulopididae Zeidler, 2004 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42
Family Lestrigonidae Zeidler, 2004 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42
Family Phronimidae Dana, 1852 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42
Family Phrosinidae Dana, 1852 (Stebbing, 1888) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .42
Superfamily Platysceloidea Spence Bate, 1862 (Bowman & Gruner, 1973) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42
Family Amphithyridae Zeidler, 2016 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43
Family Anapronoidae Bowman & Gruner, 1973 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43
Family Brachyscelidae Stephensen, 1923 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43
Family Eupronoidae Zeidler, 2016 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43

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Zootaxa 4265 (1) © 2017 Magnolia Press
Family Lycaeidae Claus, 1879 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43
Family Lycaeopsidae Chevreux, 1913 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43
Family Oxycephalidae Spence Bate, 1862 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43
Family Parascelidae Bovallius, 1887 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43
Family Platyscelidae Spence Bate, 1862. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43
Family Pronoidae Claus, 1879. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44
Family Thamneidae Zeidler, 2016 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44
Family Tryphanidae Boeck, 1871 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44
Suborder Colomastigidea Stebbing, 1899 stat. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44
Infraorder Colomastigida Stebbing, 1899 stat. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44
Parvorder Colomastigidira Stebbing, 1899 stat. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44
Superfamily Colomastigoidea Stebbing, 1899 stat. nov.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44
Family Colomastigidae Stebbing, 1899 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44
Parvorder Pagetinidira K.H. Barnard, 1931 stat. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .45
Superfamily Pagetinoidea K.H. Barnard, 1931 stat. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
Family Pagetinidae K.H. Barnard, 1931 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
Suborder Hyperiopsidea Bovallius, 1886, stat. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
Infraorder Hyperiopsida Bovallius, 1886, stat. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
Parvorder Hyperiopsidira Bovallius, 1886, stat. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
Superfamily Hyperiopsoidea Bovallius, 1886, stat. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .45
Family Hyperiopsidae Bovallius, 1886 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
Family Vitjazianidae Birstein & Vinogradov, 1955 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46
Parvorder Podosiridira Lowry & Myers, 2012b stat. nov.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46
Superfamily Podosiroidea Lowry & Myers, 2012b stat. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46
Family Podosiridae Lowry & Myers, 2012b . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46
Suborder Senticaudata Lowry & Myers, 2013 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46
Suborder Amphilochidea Boeck, 1871 stat. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46
Infraorder Amphilochida Boeck, 1871 stat. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46
Parvorder Maxillipiidira Ledoyer, 1973 stat. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47
Superfamily Maxillipioidea Ledoyer, 1973 stat. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47
Family Maxillipiidae Ledoyer, 1973 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47
Parvorder Oedicerotidira Ledoyer, 1973, stat. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .47
Superfamily Oedicerotoidea Lilljeborg, 1865b (Bousfield, 1979) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47
Family Exoedicerotidae Barnard & Drummond, 1982a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47
Family Oedicerotidae Lilljeborg, 1865b . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47
Family Paracalliopiidae Barnard & Karaman, 1982 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48
Parvorder Eusiridira Stebbing, 1888 stat. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48
Superfamily Eusiroidea Stebbing, 1888 (Bousfield, 1979) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48
Family Bateidae Stebbing, 1906 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .48
Family Eusiridae Stebbing, 1888. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48
Family Miramarassidae Lowry, 2006 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48
Family Thurstonellidae Lowry & Zeidler, 2008 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48
Superfamily Liljeborgioidea Stebbing, 1899 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49
Family Liljeborgiidae Stebbing, 1899 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .49
Subfamily Idunellinae d’Udekem d’Acoz, 2010. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49
Subfamily Liljeborgiinae Stebbing, 1899 (d’Udekem d’Acoz, 2010) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .49
Family Pseudamphilochidae fam. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49
Parvorder Amphilochidira Boeck, 1871 stat. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50
Superfamily Amphilochoidea Boeck, 1871 stat. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50
Family Amphilochidae Boeck, 1871 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50
Family Bolttsiidae Barnard & Karaman, 1987 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50
Family Cressidae Stebbing, 1899 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
Family Cyproideidae J.L. Barnard, 1974 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .51
Family Didymocheliidae Bellan-Santini & Ledoyer, 1986 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
Family Nihotungidae J.L. Barnard, 1972a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
Family Pleustidae Buchholz, 1874 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .51
Subfamily Atylopsinae Bousfield & Hendrycks, 1994a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
Subfamily Austropleustinae Bousfield & Hendrycks, 1994a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .51
Subfamily Dactylopleustinae Bousfield & Hendrycks, 1994a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
Subfamily Eosymtinae Bousfield & Hendrycks, 1994a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52
Subfamily Mesopleustinae Bousfield & Hendrycks, 1994a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52
Subfamily Neopleustinae Bousfield & Hendrycks, 1994a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52
Subfamily Parapleustinae Bousfield & Hendrycks, 1994a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52
Subfamily Pleusirinae Bousfield & Hendrycks, 1994a. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52

Zootaxa 4265 (1) © 2017 Magnolia Press
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AMPHIPODA PHYL OGENY AND CLASSIFICATION
Subfamily Pleustinae Buchholz, 1874 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52
Subfamily Pleustoidinae Bousfield & Hendrycks, 1994a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52
Subfamily Pleusymtinae Bousfield & Hendrycks, 1994a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52
Subfamily Stenopleustinae Bousfield & Hendrycks, 1994a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .52
Family Sebidae Walker, 1907 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53
Family Seborgiidae Holsinger, 1980 stat. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53
Family Stenothoidae Boeck, 1871 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53
Superfamily Leucothoidea Dana, 1852 (Bousfield, 1979) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53
Family Leucothoidae Dana, 1852 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53
Superfamily Iphimedioidea Boeck, 1871 stat. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53
Family Acanthonotozomatidae Stebbing, 1906 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54
Family Acanthonotozomellidae Coleman & Barnard, 1991a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54
Family Amathillopsidae Pirlot, 1934 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .54
Subfamily Amathillopsinae Pirlot, 1934 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .54
Subfamily Cleonardopsinae Lowry, 2006 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54
Subfamily Parepimeriinae Lowry, 2006 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54
Family Dikwidae Coleman & Barnard, 1991a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54
Family Epimeriidae Boeck, 1871 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54
Family Iphimediidae Boeck, 1871. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54
Family Lafystiidae Sars, 1893 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55
Family Laphystiopsidae Stebbing, 1899 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55
Family Ochlesidae Stebbing, 1910 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55
Family Odiidae Coleman & Barnard, 1991a. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55
Family Sicafodiidae Just, 2004 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55
Family Stilipedidae Holmes, 1908 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55
Subfamily Astryinae Pirlot, 1934 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55
Subfamily Alexandrellinae Holman & Watling, 1983 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .55
Subfamily Stilipedinae Holmes, 1908 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55
Family Vicmusiidae Just, 1990 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56
Infraorder Lysianassida Dana, 1849 stat. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .56
Parvorder Synopiidira Dana, 1852 stat. nov.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56
Superfamily Dexaminoidea Leach, 1814 (Bousfield, 1979) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56
Family Atylidae Lilljeborg, 1865a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56
Subfamily Anatylinae Bulycheva, 1955 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56
Subfamily Atylinae Lilljeborg, 1865a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56
Subfamily Lepechinellinae Schellenberg, 1926a (Bousfield & Kendall, 1994) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57
Subfamily Nototropiinae Costa 1853 (Bousfield & Kendall, 1994) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57
Family Dexaminidae Leach, 1814. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57
Subfamily Dexamininae Leach, 1814 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57
Subfamily Dexaminoculinae Bousfield & Kendall, 1994 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57
Subfamily Polycheriinae Bousfield & Kendall, 1994 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57
Subfamily Prophliantinae Nicholls, 1939 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57
Family Melphidippidae Stebbing, 1899 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .57
Family Pardaliscidae Boeck, 1871 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57
Superfamily Synopioidea Dana, 1852 (Bousfield, 1979) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58
Family Ampeliscidae Krøyer, 1842 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58
Family Argissidae Walker, 1904 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58
Family Synopiidae Dana, 1852 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58
Parvorder Haustoriidira Stebbing, 1906 stat. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59
Superfamily Haustorioidea Stebbing, 1906 (Barnard & Drummond, 1982b). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59
Family Cheidae Thurston, 1982 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59
Family Condukiidae Barnard & Drummond, 1982b. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59
Family Haustoriidae Stebbing, 1906 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59
Family Ipanemidae Barnard & Thomas, 1988b . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59
Family Otagiidae Hughes & Lörz, 2013 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59
Family Phoxocephalidae G.O. Sars, 1891 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .60
Subfamily Harpiniinae Barnard & Drummond, 1978 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 60
Subfamily Phoxocephalinae G.O. Sars, 1891 (Barnard & Drummond, 1978) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 60
Family Phoxocephalopsidae Barnard & Drummond, 1982b. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61
Family Platyischnopidae Barnard & Drummond, 1979 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61
Family Pontoporeiidae Dana, 1853 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61
Family Priscillinidae d’Udekem d’Acoz, 2007 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61
Family Sinurothoidae Ren, 1999. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61
Family Urohaustoriidae Barnard & Drummond, 1982b . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61

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Family Urothoidae Bousfield, 1979 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61
Family Zobrachoidae Barnard & Drummond, 1982b . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 62
Parvorder Lysianassidira Dana, 1849 stat. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 62
Superfamily Alicelloidea Lowry & De Broyer, 2008 stat. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 62
Family Alicellidae Lowry & De Broyer, 2008 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .62
Family Parargissidae fam. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 62
Family Podoprionidae Lowry & Stoddart, 1996 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63
Family Valettidae Stebbing, 1888 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63
Family Valettiopsidae Lowry & De Broyer, 2008 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63
Vemanidae fam. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63
Superfamily Stegocephaloidea Dana, 1852 (Bousfield, 1979) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .63
Family Stegocephalidae Dana, 1852 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .63
Subfamily Andaniexinae Berge & Vader, 2001 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .64
Subfamily Andaniopsinae Berge & Vader, 2001 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64
Subfamily Bathystegocephalinae Berge & Vader, 2001 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64
Subfamily Parandaniinae Berge & Vader, 2001 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64
Subfamily Stegocephalinae Dana, 1852 (Berge & Vader, 2001) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64
Superfamily Lysianassoidea Dana, 1849 (Bousfield, 1979) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .64
Family Adeliellidae fam. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64
Family Amaryllididae Lowry & Stoddart, 2002 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .65
Subfamily Amaryllidinae Lowry & Stoddart, 2002 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65
Subfamily Vijayiinae Lowry & Stoddart, 2002 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .65
Family Cebocaridae Lowry & Stoddart, 2011a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65
Family Cyclocaridae Lowry & Stoddart, 2011a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65
Family Cyphocarididae Lowry & Stoddart, 1997 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .65
Family Eurytheneidae Stoddart & Lowry, 2004 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65
Family Hirondelleidae Lowry & Stoddart, 2010a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .65
Family Lysianassidae Dana, 1849 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65
Subfamily Lysianassinae Dana, 1849 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66
Subfamily Waldeckiinae Lowry & Kilgallen, 2014a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66
Family Opisidae Lowry & Stoddart, 1995b . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .66
Family Scopelocheiridae Lowry & Stoddart, 1997 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .66
Subfamily Paracallisominae Kilgallen & Lowry, 2015a. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66
Subfamily Scopelocheirinae Lowry & Stoddart, 1997 (Kilgallen & Lowry, 2015a) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66
Family Tryphosidae Lowry & Stoddart, 1997 stat. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66
Family Uristidae Hurley, 1963 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67
Superfamily Aristioidea Lowry & Stoddart, 1997 stat. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67
Family Acidostomatidae Stoddart & Lowry, 2012 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67
Family Ambasiidae fam. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67
Family Aristiidae Lowry & Stoddart, 1997 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 68
Family Conicostomatidae Lowry & Stoddart, 2012b stat. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 68
Family Derjugianidae fam. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 68
Family Endevouridae Lowry & Stoddart, 1997 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .68
Family Izinkalidae Lowry & Stoddart, 2010c . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 68
Family Kergueleniidae Lowry & Stoddart, 2010d . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
Family Lepidepecreellidae Stoddart & Lowry, 2010b . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .69
Family Pakynidae nom. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
Family Sophrosynidae Lowry & Stoddart, 2010b . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
Family Thoriellidae Lowry & Stoddart, 2011a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .69
Family Trischizostomatidae Lilljeborg, 1865b . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
Family Wandinidae Lowry & Stoddart, 1990 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70

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AMPHIPODA PHYL OGENY AND CLASSIFICATION
Abstract
A classification is proposed for the order Amphipoda. The Amphipoda includes six suborders, the Pseudingolfiellidea,
Hyperiidea, Colomastigidea, Hyperiopsidea, Senticaudata (described in a previous contribution (Lowry & Myers 2013))
and Amphilochidea. The suborder Ingolfiellidea is raised to order status. A cladistic tree, based on morphology, is pre-
sented illustrating the relationships of the Amphipoda at parvorder level. A tree for the families of the Physomatidira and
Physocephalatidira, a tree for the Maxillipiidira, Oedicerotidira, Eusiridira and Amphilochidira and a tree for the Synopi-
idira, Haustoriidira and Lysianassidira, are provided. Families are listed together with their included genera. New families
are diagnosed.
Key words: Phylogeny, Classification, Crustacea, Amphipoda, Ingolfiellida, New Suborders, New Infraorders, New Par-
vorders, New Superfamilies, New Families
Introduction
The Amphipoda is one of the largest orders of the Crustacea, Malacostraca with 223 families, 1618 genera and a
little less than 10,000 species. Species diversity is greatest in the newly created parvorder Lysianassidira, followed
closely by the newly created parvorders Amphilochidira, Caprellidira and Hadziidira (Fig. 1). The majority of
amphipods (about 81%) are marine or estuarine. Freshwater amphipods make up about 19%, of which the vast
majority (about 17%) are found in the parvorders Crangonyctidira or Gammaridira. About 3% of amphipods, all in
the Talitridira, are supralittoral or terrestrial. Most marine amphipods are benthic detrital-feeders, deposit-feeders,
suspension-feeders or predators, although information is often not precise enough to put them into specific groups.
Among the lysianassidirans, a little more than half are scavengers, making up about 6% of all amphipod species.
Parasitism is rare in amphipods. Laval (1980) reviewed hyperiidean amphipods as crustacean parasitoids
associated with gelatinous zooplankton. In his opinion all hyperiideans are parasitoids, but he found evidence for
parasitism mainly in the Physocephalatidira (about 2% of amphipods). If all hyperiideans are considered as
parasitoids then the figure is nearly 3%.
FIGURE 1. The relative species contribution of each parvorder to the Amphipoda. Excluded from pie chart: Carangoliopsidira
(2 species); Maxillipiidira (3 species); Pagetinidira (4 species); Podosiridira (1species); Pseudingolfiellidira (4 species).

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The current higher classification of the Amphipoda is not phylogenetic and consequently is in need of revision.
Families have traditionally been placed in alphabetical order (Barnard & Karaman 1991, Martin & Davis 2001).
This paper completes the work reported earlier by Myers & Lowry (2003) and Lowry & Myers (2013) on the
higher classification of the Amphipoda. A full classification of the Ingolfiellida and the Amphipoda is provided.
Previous morphological phylogenetic analyses of amphipod higher taxa have been either ‘phenetic’ (Bousfield
& Kendall 1994, Bousfield & Hendrycks 1995a, 1995b, 1997, 2002, Wang & Holsinger 2001), a now almost
universally discarded method; ‘phyletic’, a somewhat unclear method espoused by E.L. Bousfield (2001) that
creates trees based on ill-defined “first principles”; or ‘cladistic’ (Berge et al. 2000; Serejo 2004; Krapp-Schickel
2009, 2011). The methodology of these cladistic analyses has been to establish a list of characters with character
states, process them through a phylogenetic inference package and then publish the resulting tree. In our studies, by
contrast, the first tree is merely a starting point for our work. We use successively modified trees to determine
possible synapomorphies, delete uninformative characters and re-examine homoplastic characters. We identify and
examine character conflict and weigh evidence for homology. Only after many iterations do we come up with a
phylogenetic hypothesis expressed as a tree.
Statistical methods have played an increasingly important role in phylogenetic studies. Bootstrap, % jackknife
resampling, and other statistical measures are used at nodes to support cladistics trees. These internal tests of
robustness are, however, fundamentally phenetic and more importantly they do not test the quality of
synapomorphies. As Buhay (2009) has demonstrated, pumpkin pie can be shown to be the sister taxon to a crayfish
with 100% bootstrap support.
There have also been several molecular genetic studies on amphipod higher taxa e.g. Englisch et al. (2003),
MacDonald et al. (2005), Ito et al. (2007), Havermans et al. (2010), Hurt et al. (2013) and Verheye et al. (2015).
Each new molecular phylogeny produced for any given set of taxa is not built on any previous hypotheses but is
separate and unrelated (Mooi et al. 2011). There is no progress in understanding—only a different and ever-
changing picture of relationships that is tied to the ingenuity of the mathematics (Mooi et al. 2011). Molecular
studies rely on similarity between aligned sequences. Similarity is nothing more than two objects compared in
some way. Homology, by contrast, is a three-item relationship in which two homologs are more closely related to
each other than they are to a third. When homology is not addressed in molecular systematics there is a risk of
making unsupportable claims of relationship (Ebach et al. 2011). Many published molecular trees, unlike
morphology based trees, are not falsifiable, because their synapomorphies are not retrievable by the reader.
Molecular studies such as those of Verheye et al. (2015) use retrospective evidence, selecting only those
morphological synapomorphies that support their molecular tree while dismissing as homoplasies, any
morphological synapomorphies that do not support their tree. We determine synapomorphies on the basis of
carefully assessed morphological evidence, we do not pick and choose only those synapomorphies that fit some
preconceived notion of a correct tree. Molecular phylogeneticists also tend to ignore the fact that genes are not free
from homoplasy which can be viewed as noise that hides phylogenetic information (Hassanin et al. 1998).
Verheye et al. (2015) dismiss the senticaudate character of apical robust setae on uropods 1 and 2 (established
by Lowry & Myers, 2013) as convergent on the basis of a small number of taxa outside the Senticaudata that
exhibit this character. Our work is based on the entire known world families of Amphipoda. Our studies revealed a
very strong synapomorphy for the senticaudates (apical robust setae on uropods 1–2) and Lowry & Myers (2013)
clearly stated that a few species outside the senticaudate clade have secondarily developed robust setae on the
apices of uropods 1 and 2 rami.
Molecular studies rarely justify unexpected relationships revealed by their hypothesised phylogenies. For
example, Verheye et al. (2015) show Salentinella Ruffo, 1948 (Salentinellidae) to be a sister taxon to Haustorius
Müller, 1775 (Haustoriidae) and Syrrhoe Goës, 1866 (Synopiidae) to be a sister taxon to Bactrurus Hay, 1902 and
Crangonyx Spence Bate, 1859 (Crangonyctidae). These relationships do not fit any previous understanding of
amphipod family relationships. They are extraordinary to say the least and require discussion, but none is provided.
Molecular studies question morphologically based classifications, but their studies are based on extremely few
taxa and as a consequence they are not able to propose any new classification to replace the existing classifications.
We do not decry molecular methods, indeed we believe that morphological and molecular data should be
viewed as complementary. Myers & Lowry (2003) reduced the suborder Caprellidea to the status of a superfamily
within the suborder Corophiidea based on a morphological cladistic study. Later, Ito et al. (2011) came to the same
conclusion based on molecular studies. When morphological and molecular results coincide, it increases the

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confidence in the hypothesis. We reject the concept that molecular phylogenies are implicitly ‘correct’ and
morphological phylogenies by inference are incorrect and should consequently be abandoned. As pointed out by
Dilman & Hilton (2011), “there is no “truth” in systematics—morphological or molecular—which, in part, is what
makes it a science”.
In our analysis of parvorders we chose Metaingolfiella as our outgroup. The Ingolfiellida are the sister group to
the Amphipoda. Wilson (2009) in his paper on the phylogenetic position of the Isopoda places Ingolfiella as an
outgroup to the four diverse Amphipoda included in his analysis. According to Ruffo (1969) and Vonk & Schram
(2003), Metaingolfiella lies at the base of the ingolfiellidan clade. For the analyses of the Amphilochidira,
Physomatidira and Physocephalatidira we chose Pseudingolfiella as our outgroup because it was shown in our
parvorder analysis to be at the base of the amphipod clade. In our analysis of one infraorder (Lysianassida) we used
Lundberg rooting. In Lundberg rooting the shortest ingroup network is rooted at the internode to which the
hypothetical ancestor attaches most parsimoniously (Lundberg, 1972). This gave us much better resolution on the
tree than did Pseudingolfiella.
The problem with large analyses of many taxa is that characters and character states have to be chosen to
represent adequately all taxa in the analysis. This leads to redundant characters for some clades in the tree. Also
characters/states that have high phylogenetic value to some clades may have low phylogenetic inference to others.
We decided that we would achieve higher resolution of family relationships by analysing infraorders separately.
Accordingly, our all-family tree was used simply as a basis for determining parvorders and infraorders for the next
analyses. Infraorders were then analysed individually using a subset of relevant characters/states. Accordingly
separate analyses were carried out for the Hyperiidea, Amphilochida and Lysianassida in order to introduce greater
resolution to the classification of those infraorders. Within the Lysianassida, a separate analysis was carried out for
the Haustoriidira and the results are shown here as a composite infraorder cladogram.
In this contribution, one new order Ingolfiellida Hansen, 1903 stat. nov., and four new suborders:
Pseudingolfiellidea Lowry & Myers, 2012a stat. nov.; Colomastigidea Stebbing, 1899 stat. nov.; Hyperiopsidea
Bovallius, 1886 stat. nov. and Amphilochidea Boeck, 1871 stat. nov. are erected. These join the existing suborders
Hyperiidea H. Milne Edwards, 1830 and Senticaudata Lowry & Myers 2013. Within these six amphipod suborders
there are 13 infraorders: Pseudingolfiellida Lowry & Myers, 2012a stat. nov., Physosomata Pirlot, 1929;
Physocephalata Bowman & Gruner, 1973; Colomastigida Stebbing, 1899 stat. nov.; Hyperiopsida Bovallius, 1886
stat. nov.; Carangoliopsida Lowry & Myers, 2013; Talitrida Rafinesque, 1815 (Serejo, 2004); Corophiida Leach,
1814 (Lowry & Myers 2013); Hadziida S. Karaman, 1943 (Lowry & Myers 2013); Bogidiellida Hertzog, 1936
(Lowry & Myers 2013); Gammarida Latreille, 1802 (Lowry & Myers 2013); Amphilochida Boeck, 1871 stat. nov.
and Lysianassida Dana, 1849 stat. nov. There are 23 amphipod parvorders, 15 of which are new:
Metaingolfiellidira Ruffo, 1969 stat. nov., Pseudingolfiellidira Lowry & Myers, 2012a stat. nov., Physomatidira
Pirlot, 1929 stat. nov.; Physocephalatidira Bowman & Gruner, 1973 stat. nov.; Colomastigidira Stebbing, 1899
stat. nov.; Pagetinidira K.H. Barnard, 1931 stat. nov.; Hyperiopsidira Bovallius, 1886 stat. nov.; Podosiridira
Lowry & Myers, 2012b stat. nov.; Carangoliopsidira Lowry & Myers, 2013; Talitridira Rafinesque, 1815 (Lowry
& Myers, 2013); Corophiidira Leach, 1814 (Lowry & Myers 2013); Caprellidira Leach, 1814 (Lowry & Myers,
2013); Hadziidira S. Karaman, 1943 (Lowry & Myers 2013); Bogidiellidira Herztog, 1936 (Lowry & Myers 2013);
Crangonyctidira Bousfield, 1973 (Lowry & Myers 2013); Gammaridira Latreille, 1802 (Lowry & Myers 2013);
Maxillipiidira Ledoyer, 1973 stat. nov.; Oedicerotidira Lilljeborg, 1865b (Bousfield, 1979) stat. nov.; Eusiridira
Stebbing 1888 stat. nov.; Amphilochidira Boeck, 1871 stat. nov.; Synopiidira Dana, 1852 stat. nov.; Haustoriidira
Stebbing, 1906 stat. nov. and Lysianassidira Dana, 1849 stat. nov. In addition two recently described senticaudate
families and three recently described hyperiidean families are included for completeness: Australomicroprotopidae
Myers, Lowry & Billingham, 2016, Zaramillidae Lowry & Myers, 2016, Amphithyridae Zeidler, 2016,
Eupronoidae Zeidler, 2016 and Thamneidae Zeidler, 2016.
The relative contribution of each parvorder to the Amphipoda is shown in Fig. 1.
Materials and methods
The basis of our analyses was an unpublished database to the families of world Amphipoda, based on a wide
representative selection of genera, built up over many years by one of us (JKL). A DELTA (Dallwitz 2005)

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database of 300 characters each with two or more character states was analysed in PAUP version 4.0b8a (Swofford
2003) using heuristic searches and the criterion of parsimony, to give a first tree. This tree was then analysed in
MacClade (version 3.08) to determine which characters were potentially useful synapomorphies, which were
uninformative and which appeared to be homoplastic. Characters that appeared to be homoplastic were re-
examined to see if a supposed character state was actually two or more characters that could be redefined with
separate states. If so, the newly defined characters and states were then put back into the analysis and another run
was performed. Synapomorphies were challenged by each successive tree. This process of reanalysis and
reiteration was then continued until the shortest tree with the most resolved synapomorphies resulted. In this
analysis almost 100 iterations were performed. In the analyses, only synapomorphies were used. For the taxon
diagnoses, each character state was checked using the intkey option in the Delta database of world families.
Diagnostic descriptions sometimes used characters that were autapomorphies, and therefore could not be used in
the phylogenetic analyses.
A synapomorphy is defined as a shared uniquely derived character state. In the process of evolution a character
state can be transformed. One possible transformation is ‘lost’. For example, in insects a synapomorphy is ‘wings’,
but fleas have no wings because in the process of evolution, the wings have been lost. It follows that not all
members of a clade may necessarily share a given synapomorphy. In determining synapomorphies we record a
character state as synapomorphic if it is present in the majority, not necessarily all, of the members of a family or
higher taxon. If a character state is present in a number of members of a clade, the most parsimonious solution is
that it has been derived through an evolutionary lineage from an ancestor and is thus a synapomorphy. The
alternative is that is has been evolved independently in each taxon that exhibits the character state and is therefore
an autapomorphy for each taxon. This alternative is much less parsimonious.
An important synapomorphy for the Iphimedioidea is the acuminate pereopod 4 coxa, even though this state
does not occur in a few small specialised families, the Laphystiopsidae, Sicafodiidae, Vicmusiidae and some (but
not all) of the Acanthonotozomellidae. Here we are left with two hypotheses. Firstly, that the acuminate coxa 4 has
evolved independently in nine families (Amathillopsidae, Epimeriidae, Lafystiidae, Dikwidae, Iphimediidae,
Stilipedidae, Acanthonotozomatidae, Ochlesidae and the Odiidae, i.e. nine autapomorphies, or secondly that the
Iphimedioidea are monophyletic (i.e. the character state is derived from a common ancestor), and that the character
state has been modified (loss of acuminate tips) during descent in a few small aberrant families. We believe that the
second hypothesis (monophyly) is the most parsimonious.
An important synapomorphy for the parvorder Amphilochidira is the pereopod 4 carpus shorter than the
propodus. It has been modified in the Amphilochidae, Cyproideidae, Amathillopsidae and some Ochlesidae.
However, it remains a synapomorphy for 18 out of the 22 families in the Amphilochidira. The discovery of some
taxa that do not possess the character state, does not falsify our hypothesis.
A fundamental attribute of science is that each new scientific discovery builds on, and is informed by, past
scientific endeavours. Science advances by building on the bricks of past workers. When studying amphipod
phylogeny we can refer to the works of scientists such as G.O. Sars and T.R.R. Stebbing in the nineteenth century
and J.L Barnard and E.L Bousfield in the twentieth century along with hundreds of other amphipodologists who
have together established a broad framework for our understanding of amphipod relationships. We use this
accumulated knowledge to assist us in recognising homoplasies. If a character state is shared (synapomorphy) by
members of a clade, but is also present in a taxon outside that clade, in a ‘distant’ part of our cladistic tree, we use
the historical knowledge of amphipod relationships together with our cladistics analysis (showing the pattern of
synapomorphies), to determine whether the character state in the ‘distant’ taxon is a homoplasy. For example, the
presence of apical robust setae on uropods 1 and 2 is a synapomorphy for the Senticaudata (Lowry & Myers, 2013),
but we find this character state also in some Phoxocephalidae. The acquired knowledge from all past studies is that
the Phoxocephalidae are not closely related to any senticaudates, strengthening our assumption that this is a
homoplasy. The assumption is complemented by our finding of several other synapomorphies that place the
Phoxocephalidae in the Haustorioidea of the Lysianassida. The combination of synapomorphies from our analysis
and the perceived wisdom of past amphipodologists direct our assumptions. When we examine the phoxocephalids
in question we find that their appendages are covered in robust setae. This is due to their burrowing habits. If
numerous robust setae are developed over their appendages to assist burrowing, it is not unexpected that they have
also developed robust setae on the apices of their uropods.
It is worth noting that molecular studies depart from normal scientific progress by not building on, nor taking

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cognisance of, any previous work. Each molecular analysis stands in isolation and the results are taken as ‘correct’
without any reference to past understanding, including other molecular studies with which they may be conflicting.
The analysis of very large databases provides a useful overview of relationships but can give weak
phylogenetic relationship information below the infraorder level because some of the characters/states chosen for
the large analysis are not applicable to individual parvorders. Better results may be obtained at the family level by
analysing infraorder clades separately and choosing only the appropriate characters/states for each analysis.
Accordingly separate analyses were carried out for the Hyperiidea, Amphilochida and Lysianassida in order to
introduce greater resolution to the classification of those infraorders.
In the all-family analysis we used 118 characters (Figs 2–10) for 126 taxa. In the parvorder analysis we
analysed 21 taxa using 27 characters. In the infraorder Amphilochida analysis there were 34 families in two
parvorders and 72 characters were used. In the infraorder Lysianassida analysis there were 54 families in three
parvorders and 77 characters were used and in the suborder Hyperiidea analysis there were 32 families in two
infraorders and two parvorders using 80 characters. Matrices of the original character sets used in these analyses
are available as Supplementary files on the body page of this paper in the website of Zootaxa.
Two measures of tree performance are given. The consistency index (CI)—the minimum number of changes
divided by the number required on the tree (CI = 1 if there is no homoplasy) and the retention index (RI) calculated
by taking the maximum number of changes on a tree minus the number of changes on the tree and dividing by the
maximum number of changes on the tree minus the minimum number of changes in the dataset.
The terminology for spines and setae follows Watling (1989). The classification of calceolus types follows
Lincoln & Hurley (1981). Also we are using terminology more in line with that used for other peracaridan groups.
For mouthparts, the inner and outer plates of the maxilla 1 and maxilla 2 are termed the basal endite and ischial
endite respectively. Jaume et al. (2009) interpret the maxillae plates as coxal and basal endites. This confusion can
only be rectified by embryological studies and for the moment we prefer to use a terminology that is consistent
between maxillae and maxilliped. We also prefer to retain the terms antenna 1 and antenna 2 instead of antennules
and antennae and likewise maxilla 1 and maxilla 2 instead of maxillule and maxilla, simply because we feel these
terms are less subject to confusion.
Characters used in the analysis
(Figs 2–10)
Subequal = within 1.1% difference.
1. Body
1. laterally compressed
2. subcylindrical
3. laterally compressed with small coxae
4. small, coxae often fused to body
5. vermiform
6. dorsoventrally flattened
7. subglobular in males and females
8. subglobular in males, globular in females
2. Head
1. as long as deep
2. longer than deep
3. deeper than long
4. much deeper than long (at least 1.5 ×)
3. Head anteroventral margin
1. recessed
2. concave
3. rounded
4. vertical
5. oblique
6. acute/subacute
7. produced
4. Head rostrum
1. vestigial or absent
2. present

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FIGURE 2. Body shape and head anteroventral margin. 1.1 laterally compressed (after Krapp-Schickel, 2006a); 1.2
subcylindrical (after LeCroy 1995); 1.6 dorsoventrally flattened (after Barnard & Drummond 1981); 1.7 subglobular (after
Zeidler 2012); 3.1 recessed (after d’Udekem d’Acoz 2010); 3.2 concave (after Myers et al. 1987); 3.3 rounded (after Zeidler
2003b); 3.4 vertical (after Coleman 2010); 3.5 oblique (after King 2009); 3.6 acute (after Coleman 1994); 3.7 produced (after
Just 1990).

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FIGURE 3. Rostrum. 5.2 cylindrical (after Souza-Filhol 2011) 5.3 cylindrical with anteroventral keel (after J.L. Barnard &
Drummond 1979); 5.4 dorsoventrally flattened (after J.L. Barnard 1999); 5.5 laterally flattened (after White & Thomas 2009);
5.6 visor-like (after J.L. Barnard & Drummond 1978); 5.7 lanceolate (after Zeidler 2012); 5.8 spine-like (after Lowry & Myers
2003); 5.9 recurved (after J.L. Barnard 1970). Eye. 6.2 coalesced (after Thomas & Barnard 1985); 7.4 ventrally tapered (after
Zeidler 2003b); 7.5 occupying most of lateral surface of head (after Zeidler 2004).

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5. Head rostrum
1. absent
2. cylindrical
3. cylindrical with anteroventral keel
4. dorsoventrally flattened
5. laterally flattened
6. visor like
7. lanceolate
8. spine-like
9. recurved
6. Eyes
1. not coalesced
2. coalesced
3. absent
7. Eye shape
1. round
2. ovoid
3. reniform
4. ventrally tapered
5. occupying most of lateral surface of head
6. vestigial or absent
8. Antenna 1–2
1. type 3 calceolus absent (lysianassoid-type)
2. type 3 calceolus present (lysianassoid-type)
9. Antenna 1–2
1. type 7 calceolus absent (oedicerotoid-type)
2. type 7 calceolus present (oedicerotoid-type)
10. Antenna 1–2
1. type 8 calceolus absent (haustorioid-type)
2. type 8 calceolus present (haustorioid-type)
11. Antenna 1
1. subequal in length to antenna 2
2. longer than (more than 1.1 ×) antenna 2
3. shorter than (less than 0.9 ×) antenna 2
4. shorter than peduncle of antenna 2
11A. Antenna 1
1. inserted on anterior surface of head
2. inserted on anteriolateral surface of head
3. inserted on ventral surface of head
12. Antenna 1
1. not spear-like
2. spear-like
13. Antenna 1 peduncle
1. 3-articulate
2. 1- or 2-articulate
14. Antenna 1 peduncular article 1
1. subequal to article 2
2. longer than (more than 1.1 ×) article 2
3. shorter than (less than 0.9 ×) article 2
15. Antenna 1 peduncle
1. with sparse robust and slender setae
2. with many robust and slender setae
3. with many slender setae only
4. with sparse slender setae only
16. Antenna 1 primary flagellum
1. 5- or more articulate
2. less than 5-articulate
17. Antenna 1 callynophore
1. well-developed (2-field)
2. weakly developed (1-field)
3. absent
18. Antenna 1 accessory flagellum
1. present
2. vestigial or absent (1-articulate or scale-like)
19. Antenna 1 accessory flagellum
1. not forming cap
2. forming cap
20. Antenna 2
1. medium length (0.33 to 0.66 × body length)
2. short (less than 0.33 × body length)
3. long (0.66 × to subequal body length)
4. greater than body length
5. absent
20A. Antenna 2
1. inserted on anterior surface of head
2. inserted on ventral surface of head
3. absent in female
21. Antenna 2
1. articles not folded in zigzag fashion
2. male peduncular and flagellar articles folded in
zigzag fashion
22. Antenna 2 peduncle
1. with sparse robust and slender setae
2. with sparse or no slender setae
3. with many slender setae
4. with many robust and slender setae
5. absent
23. Antenna 2 peduncle
1. article 1 not enlarged
2. article 1 bulbous
24. Antenna 2 brush setae—adult male
1. absent
2. present
25. Antenna 2 flagellum
1. subequal with peduncle
2. shorter than peduncle
3. longer than peduncle
4. absent

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FIGURE 4. Antennae. 8.2 calceolus type 3 (after Lincoln & Hurley 1981); 9.2 calceolus type 7 (after Lincoln & Hurley 1981);
10.2 calceolus type 8 (after Lincoln & Hurley 1981); 12.2 antenna 1 spear-like (after Chang-tai Shih & Hendrycks 1996); 15.2
antenna 1 peduncle with many robust or slender setae (after Thurston 1982); 17.1 antenna 1 callynophore well developed (after
Lowry & Stoddart 1995b); 17.2 antenna 1 callynophore weakly developed (1-field) (after LeCroy 1995); 19.2 antenna 1
accessory flagellum forming a cap (unpublished); 21.2 antenna 2 male peduncular and flagellar articles folded in zigzag fashion
(after Vinogradov, Volkov & Semenova 1982); 24.2 antenna 2 with brush setae (after Myers & LeCroy 2009).

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FIGURE 5. Mouthparts. 28.1 mouthpart bundle subquadrate (after Lowry & Springthorpe 2005); 28.2 mouthpart bundle
subconical (after Coleman & Lowry 2006); 29.2 labrum, epistome complex separate (after Lowry & Stoddart 1997); 30.1
mandible incisor dentate (after Coleman 1998); 30.2 mandible incisor smooth (after Lowry & Stoddart 1994); 30.3 mandible
incisor minutely serrate (after Zeidler 2004); 30.4 mandible incisor absent (after Lowry & Stoddart 1994); 33.2 mandible molar
non-setose, smooth (after Azman 2009); 33.3 mandible molar a fully setose tongue (after Stoddart & Lowry 2004); 33.5
mandible molar a broad setose flap (after Ruffo 1978); 33.6 mandible molar a smooth protuberance with large robust setae
(after d’Udekem d’Acoz 2010b); 36.1 mandible accessory setal row present (after J.L. Barnard & Drummond 1978); 36.2
mandible accessory setal row absent (after Berge et al. 2000): 40.3 maxilla 1 basal endite covered in setae (after Zeidler 2009);
44.2 maxilla 2 basal endite with oblique setal row (after Hughes 2009); 45.3 maxilla 2 basal endite covered in setae (after
Zeidler 2009).

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FIGURE 6. Mouthparts. 31.1 molar triturative (after Barnard 1970); 33.1 molar not setose (after Bousfield & Hendrycks
1994): 33.2 molar non-setose, smooth (after Holsinger 1980); 33.3 molar with fully setose tongue (after Lowry & Stoddart
1995); 33.4 molar with smooth setose protuberance (after d’Udekum d’Acoz 2010b); 33.5 molar with broad setose flap (after
Lowry & Stoddart 1996); 33.6 molar protuberance with large robust setae (after Barnard & Drummond 1978); 34.1 mandibular
molar medium (after Moore 1992b); 34.2 mandibular molar large (after Barnard & Thomas 1998); 34.3 mandibular molar
small (after Myers 1974); 34.4 mandibular molar completely dominating mandible (after Barnard 1972).

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26. Antenna 2 flagellum
1. 5 or more articulate
2. less than 5-articulate
3. absent
27. Antenna 2 flagellum—adult male
1. not elongated compared with female
2. greatly elongated compared with female
3. absent
28. Mouthpart bundle
1. subquadrate
2. subconical
29. Labrum, epistome complex
1. entire
2. separate
30. Mandible incisor
1. dentate
2. smooth
3. minutely serrate
4. vestigial or absent
31. Mandible molar
1. triturative
2. non-triturative
3. vestigial or absent
32. Mandible molar
1. strongly triturating
2. weakly triturating
3. not triturating
33. Mandible molar
1. not setose
2. non-setose smooth molar
3. fully setose tongue
4. a smooth setose protuberance
5. a broad setose flap
6. a smooth protuberance with large robust setae
34. Mandible molar
1. medium
2. large
3. small
4. completely dominating mandible
5. vestigial or absent
35. Mandible lacinia mobilis
1. present on both sides
2. present on left side only
3. absent
36. Mandible, accessory setal row
1. well-developed
2. vestigial or absent
37. Mandible molar
1. setal patch absent
2. patch of fine setae
38. Mandible palp
1. present in both sexes
2. present in male, absent in female
3. vestigial or absent in both sexes
39. Maxilla 1 basal endite (inner plate)
1. present
2. absent
40. Maxilla 1 basal endite (inner plate)
1. setose apically
2. strongly setose along medial margin
3. covered in setae
4. without setae
5. absent
41. Maxilla 1 ischial endite (outer plate) (used only for
Aristioidea and Lysianassoidea)
1. with 7/4 setal-tooth arrangement
2. with 6/5 setal-tooth arrangement
3. with 7/4 crown setal-tooth arrangement
4. with 8/3 setal-tooth arrangement
5. with 3–6 in one row
6. with complex arrangement
7. with 2 + 3–5 vestigial setal-teeth
8. setal-teeth absent
42. Maxilla 1 palp
1. well-developed
2. vestigial or absent
43. Maxilla 2 basal endite (inner plate)
1. present, endites free
2. present, fused with ischial endite
3. absent
44. Maxilla 2 basal endite (inner plate)
1. without oblique setal row
2. with oblique setal row
3. absent
45. Maxilla 2 basal endite (inner plate)
1. without setae on medial margin
2. with strongly setose medial margin
3. covered in setae
4. absent
46. Maxilliped basal endite (inner plate) fusion
1. separate
2. fused medially
3. vestigial or absent

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47. Maxilliped ischial endite (outer plate)
1. small (longer than palp article 1, not longer than
palp article 2)
2. large (longer than palp article 2, not longer than
palp article 3)
3. very large (longer than palp article 3)
4. vestigial or absent (not longer than palp article
1)
47A. Maxilliped
1. palps present
2. palps absent
48. Pereonites
1. separate
2. 1–2, 1–3, 1–4 or 1–5 fused
49. Coxal gills
1. unstalked
2. stalked
50. Coxae 1–3
1. none vestigial or reduced
2. coxa 1 reduced or absent
3. coxa 2 reduced
4. coxae 1–2 vestigial
5. coxae 1–3 reduced or vestigial
6. fused to pereonites
51. Coxae 1–4 overlapping or not
1. overlapping
2. discontiguous
3. fused to pereonites
52. Coxae 1–4
1. longer than broad
2. subequal in length and breadth
3. broader than long
4. fused to pereonites
53. Coxae 1–4
1. not acuminate
2. acuminate
3. fused to pereonites
54. Gnathopod 1
1. similar in size to gnathopod 2
2. smaller (or weaker) than gnathopod 2
3. larger (or stouter) than gnathopod 2
4. absent
55. Gnathopod 1
1. subchelate
2. simple
3. carpochelate
4. parachelate
5. chelate
6. absent
56. Gnathopod 1 coxa
1. subequal to coxa 2
2. smaller than coxa 2
3. larger than coxa 2
4. vestigial, hidden or partially hidden by coxa 2
5. vestigial, not hidden by coxa 2
6. fused to pereonite
57. Gnathopod 1 dactylus
1. large
2. reduced
3. small to minute, strongly or weakly covered in
setae
4. absent
58. Gnathopod 2
1. subchelate
2. simple
3. carpochelate
4. parachelate
5. chelate
6. eucarpochelate
59. Gnathopod 2
1. not mitten-shaped
2. mitten-shaped
60. Gnathopod 2 coxa
1. subequal to coxa 3
2. smaller than but not hidden by coxa 3
3. smaller than, mostly hidden by coxa 3
4. larger than coxa 3
5. fused to pereonite
61. Gnathopod 2 ischium
1. short (length less than 2 × breadth)
2. long (length at least 2 × breadth)
62. Gnathopod 2 carpus
1. short (length less than 2 × breadth)
2. long (length at least 2 × breadth)
63. Gnathopod 2 carpus
1. subequal to propodus
2. shorter than (less than 0.9 ×) propodus
3. longer than (more than 1.1 ×) propodus
64. Gnathopod 2 carpus
1. not produced along posterior margin of
propodus
2. strongly produced along posterior margin of
propodus
3. produced along posterior margin of propodus
65. Gnathopod 2 dactylus
1. well-developed
2. reduced
3. absent

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FIGURE 7. Pereon. 58.1 gnathopod 2 subchelate (after Myers 2012); 58.2 gnathopod 2 simple (after Bellan-Santini &
Diviacco 1990); 58.3 gnathopod 2 carpochelate (after Myers 1977); 58.4 gnathopod 2 parachelate (after Barnard 1972b);
gnathopod 2 chelate (after Barnard 1970).

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66. Pereopods
1. 3–7 without hooded dactyli
2. some or all of 3–7 with hooded dactyli
67. Pereopods 3–4
1. without glands
2. glandular
68. Pereopod 3 coxa
1. subequal in length and breadth
2. longer than broad
3. broader than long
4. fused to pereonite
69. Pereopod 3 merus
1. subequal to propodus
2. shorter than (less than 0.9 ×) propodus
3. longer than (more than 1.1 ×) propodus
4. more than 2 × longer than propodus
70. Pereopod 3 carpus
1. subequal to propodus
2. shorter than (less than 0.9 ×) propodus
3. longer than (more than 1.1 ×) propodus
71. Pereopod 3 carpus
1. not produced
2. strongly produced posteriorly
3. produced anteriorly
72. Pereopod 3 dactylus
1. well-developed
2. small or poorly developed
3. minute or absent
73. Pereopod 4 coxa
1. subequal to coxa 3
2. smaller than coxa 3
3. larger than coxa 3
4. vestigial
5. fused to pereon
74. Pereopod 4 coxa
1. without posteroventral lobe
2. with well-developed posteroventral lobe
3. with small posteroventral lobe
74A. Pereopod 4 coxa
1. not acuminate
2. acuminate ventrally
75. Pereopods 5–6 basis
1. not or moderately enlarged
2. greatly enlarged
76. Pereopods 5–7
1. similar in length
2. pereopod 5 shorter, pereopods 6 and 7 similar
length
3. pereopods 5 to 7 progressively longer
4. pereopods 5 to 7 progressively shorter
5. pereopods 5 and 6 similar in length, pereopod 7
shorter
6. pereopods 5 and 6 same length, pereopod 7
much longer
7. pereopods 5 and 7 same length, pereopod 6
much longer
8. pereopods 5 and pereopod 7 same length,
pereopod 6 extremely long—whip-like
9. pereopod 5 longer than pereopods 6 to 7
77. Pereopods 5–7
1. with few or no robust or slender setae
2. with many rows of facial and/or marginal
robust and/or slender setae
78. Pereopods 5–7 merus
1. linear
2. strongly expanded
3. slightly expanded
4. with anterodistal and posterodistal lobes
79. Pereopods 5–7 merus
1. posterior margin weakly setose
2. posterior margin strongly setose
3. posterior margin without slender setae
80. Pereopod 5
1. simple
2. carpochelate
3. subchelate
81. Pereopod 5
1. subequal to pereopod 6
2. shorter than pereopod 6
3. longer than pereopod 6
82. Pereopod 5 coxa
1. subequal to coxa 4
2. smaller than coxa 4
3. larger than coxa 4
4. vestigial
5. fused to pereonite
83. Pereopod 5 coxa
1. without lobes
2. equilobate
3. with posteroventral lobe
4. with anteroventral lobe
5. with posterodorsal lobe

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FIGURE 8. Pereon. 59.2 gnathopod 2 mitten-shaped (after Lowry & Stoddart 1995b); 64.2 gnathopod 2 carpus strongly
produced along posterior margin of propodus (after J.L. Barnard & Drummond 1982); 66.2 pereopods 3–7 with hooded dactyli
(after Zeidler 2009); 74.1 pereopod 4 coxa without posteroventral lobe (after Coleman 1998); 74.2 pereopod 4 coxa with well-
developed posteroventral lobe (after Lowry & Stoddart 1994); 74.3 pereopod 4 coxa with small posteroventral lobe (after
Lowry & Stoddart 1994); 75.2 pereopods 5–6 basis greatly enlarged (after Zeidler 1999). 77.2 pereopods 5–7 with many rows
of facial and marginal robust and slender setae (after J.L. Barnard & Drummond 1982b).

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AMPHIPODA PHYL OGENY AND CLASSIFICATION
FIGURE 9. Pereon & Pleon. 83.2 pereopod 5 coxa equilobate (after Barnard & Drummond 1982a); 83.3 pereopod 5 coxa with
posteroventral lobe (after Azman 2009); 83.4 pereopod 5 coxa with anteroventral lobe (after Bousfield & Kendall 1994); 83.5
pereopod 5 coxa with posterodorsal lobe (after Bowman 1973); 92.2 epimeron 2 with plumose setae (after J.L. Barnard &
Drummond 1982b); 98.2 uropod 1–2 rami apices with embedded setae (after Lowry & Stoddart 1995b); 99.2 uropod 1
basofacial robust seta (after Myers 1985); 100.1 uropod 1 rami styliform (after Barnard 1970); 100.2 uropod 1 rami lanceolate
(after King 2009); 100.3 uropod 1 rami linguiform (after Barnard & Drummond 1982b); 100.4 uropod 1 rami ferrulate (after
Myers 1995); 100.5 uropod 1 rami stylospatulate (after Noodt 1959); 100.6 uropod 1 rami paddle-shaped (after Bowman 1978);
103.2 uropod 2 endopod with constriction (after Lowry & Stoddart 2009).

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84. Pereopod 5 basis
1. subrectangular
2. subquadrate
3. subovate
4. subtriangular
5. round
85. Pereopod 5 basis
1. strongly expanded
2. slightly expanded
3. linear
86. Pereopod 5 carpus
1. linear
2. strongly expanded
3. slightly expanded
87. Pereopod 5 dactylus
1. well-developed
2. small or poorly developed
3. minute
4. absent
88. Pereopod 6 basis
1. strongly expanded
2. slightly expanded
3. linear
89. Pereopod 7
1. subequal to pereopod 5
2. shorter than pereopod 5
3. longer than pereopod 5
90. Pereopod 7
1. similar in structure to pereopod 6
2. different in structure from pereopod 6
91. Pleonites 1 to 3
1. without dorsal spines
2. with dorsal spines
92. Epimeron 2
1. without plumose setae
2. with plumose setae
3. epimeron 2 absent
93. Urosome
1. not carinate
2. carinate
94. Urosomites
1. 1 to 3 free
2. 1 to 2 free, urosomite 3 absent
3. 1 to 3 coalesced
4. 1–2 coalesced, 3 free
5. 1 free, 2–3 coalesced
95. Urosomite 1
1. longer than urosomite 2 or 2–3 (up to 2 ×)
2. shorter than (less than 0.9 ×) urosomite 2 or 2–3
3. subequal to urosomite 2 or 2–3
4. much longer than urosomite 2 or 2–3 (at least 3
×)
96. Urosomite 1
1. without distoventral robust seta
2. with distoventral robust seta
97. Uropods 1–2 rami
1. without apical robust setae
2. with apical robust setae
98. Uropods 1–2 rami
1. apices without embedded setae
2. apices with embedded setae
99. Uropod 1 peduncle
1. without basofacial robust seta
2. with basofacial robust seta
100. Uropod 1 rami
1. styliform
2. lanceolate
3. linguiform
4. ferrulate
5. stylospatulate
6. paddle-shaped
101. Uropod 2 rami
1. styliform
2. lanceolate
3. linguiform
4. ferrulate
5. stylospatulate
6. paddle-shaped
7. vestigial
8. absent
102. Uropod 2 endopod
1. subequal to exopod
2. shorter than (less than 0.9 ×) exopod
3. longer than (more than 1.1 ×) exopod
4. absent
103. Uropod 2 endopod
1. without constriction
2. with constriction
104. Uropod 3 peduncle
1. short (length less than 2 × breadth)
2. long (length at least 2 × breadth)
3. vestigial
4. fused to rami

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FIGURE 10. Uropods. 100.4 uropod 1 rami ferrulate (after Souza-Filhol 2011); 100.5 uropod 1 ramus stylospatulate (after
Ruffo 1969); 106.1 uropod 3 rami styliform (after Barnard 1970); 106.2, uropod 3 ramus linguiform (after Williams & Barnard
1988);106.3 uropod 3 rami lanceolate (after White & Reimer 2012); 106.4 uropod 3 rami paddle-shaped (after Krapp-Schickel
2009).
105. Uropod 3 rami
1. biramous
2. uniramous
3. rami absent
106. Uropod 3 rami
1. styliform
2. linguiform
3. lanceolate
4. paddle-shaped
5. vestigial
6. absent
107. Uropod 3 rami
1. without fringing plumose setae
2. with fringing plumose setae
3. absent
108. Uropod 3 endopod
1. not apically setose
2. apically setose
3. absent
109. Uropod 3 endopod
1. subequal to exopod
2. longer than (more than 1.1 ×) exopod
3. shorter than (less than 0.9 ×) exopod
4. minute
5. absent

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Phylogenetic analysis
All characters were unweighted with the exception of the calceoli types and the mitten-shaped gnathopod 2, each of
which was given a weight of 4. These are complex structures which should have a good index of monophyly and
we thus felt justified in giving a weighting factor to these unique characters.
Because the analysis employed an iterative process between DELTA, PAUP and MacClade, the choice of
starting tree was unimportant as long as it was a parsimoniously shortest tree. When a number of shortest trees
were available they were not significantly different from each other.
The all-family tree had a CI of 0.60 and a RI of 0.60. The parvorder tree had a CI of 0.76 and a RI of 0.70. The
Amphilochida tree had a CI of 0.66 and a RI of 0.44. The Lysianassida tree had a CI of 0.70 and a RI of 0.53, while
the Hyperiidea tree had a CI of 0.59 and a RI of 0.57.
Tree description
The Pseudingolfiellida is a small infraorder of free-living freshwater amphipods (Fig. 11) characterised by reduced
discontiguous coxae and vestigial epimeral plates. The Hyperiidea (Figs 11, 12) is a monophyletic suborder
characterized by having a minutely serrate mandibular incisor, no palps on the maxilliped and urosomite segments
2 and 3 coalesced. In addition most taxa (except Vibilioidea and some Platysceloidea) have subglobular to globular
bodies, a head deeper to much deeper than long, a mandible generally without molar, though it is present in some
Physocephalata; no mandibular accessory setal (raker) row, pereopod 4 coxa without posteroventral lobe and an
entire telson. The Colomastigida is a monophyletic infraorder (Fig. 11) characterized by having urosomites 2 and 3
coalesced, loss of the mandible lacinia mobilis on both sides and fused basal endites on the maxilliped. They also
have a subcylindrical body, antenna 1 and antenna 2 with less than 5-articulate flagellum and pereopod 4 coxa
lacking a posteroventral lobe. The Hyperiopsida (Fig. 11) is a monophyletic though poorly supported infraorder,
characterized by sub-cylindrical body, pereopod 3 merus longer than propodus and pereopod 5–7 bases linear. The
Senticaudata (Fig. 11) is a monophyletic suborder characterized by uropod 1 and 2 having apical robust setae and
uropod 3 with ferulate rami. The Amphilochidea is a monophyletic suborder containing the infraorders
Amphilochida and the Lysianassida and based on the weak synapomorphy, presence of brush setae in adult males.
The monophyletic infraorder Amphilochida (Fig. 13) is defined by coxae 1–4 overlapping and uropod 3 exopod
longer than peduncle. The Lysianassida (Fig. 14) is a large well-defined monophyletic infraorder characterised by a
short carpus on the second gnathopod, lanceolate rami on uropods 1 to 3.
110. Uropod 3 exopod
1. subequal to peduncle
2. longer than (more than 1.1 ×) peduncle
3. shorter than (less than 0.9 ×) peduncle
4. absent
111. Uropod 3 exopod
1. 2-articulate
2. 1-articulate
3. absent
112. Telson
1. entire (including notched or emarginate)
2. cleft
3. absent
113. Telson
1. laminar
2. dorsoventrally thickened
114. Telson
1. as long as broad
2. longer than broad
3. broader than long
4. absent
115. Telson
1. without apical robust setae
2. with apical robust setae

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AMPHIPODA PHYL OGENY AND CLASSIFICATION
Within the infraorder Colomastigida, members of the Colomastigidira sometimes possess a callynophore
which is always lacking in pagetinidirans. The colomastigidirans have a simple gnathopod 1 whereas it is
subchelate in pagetinidirans. In pagetinidirans, coxa 5 is smaller than coxa 4 but in colomastigidirans they are
subequal.
FIGURE 11. Cladogram of relationships of parvorders within the Amphipoda with Metaingolfiellidae as the outgroup. Cross-
lines represent significant synapomorphies with character state transformation. A1 = peduncular article 3 long. FT = fleshy
telson. GP = glandular pereopods 3–4.

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FIGURE 12. Cladogram of relationships of Hyperiidea with Pseudingolfiellidae as the outgroup. Cross-lines represent
significant synapomorphies with character state transformation.

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AMPHIPODA PHYL OGENY AND CLASSIFICATION
FIGURE 13. Cladogram of relationships of Amphilochida with Pseudingolfiellidae as the outgroup. Cross-lines represent
significant synapomorphies with character state transformation.

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FIGURE 14. Cladogram of relationships of Lysianassida with Lundberg rooting. Cross-lines represent significant
synapomorphies with character state transformation.

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AMPHIPODA PHYL OGENY AND CLASSIFICATION
The Hyperiopsidira have a callynophore, a long pereopod 3 merus, coxa 4 subequal with coxa 3 and without a
posteroventral lobe, coxa 5 smaller than coxa 4, linear bases on pereopods 5 and 6 and urosomite 1 much longer
than urosomite 2. The Podosiridira have a subcylindrical body, a head that is longer than deep, antenna 1 with
peduncular articles 1 and 2 subequal, antenna 2 flagellum longer than the peduncle, mandible molar small, coxae
1–4 broader than long, gnathopod 1 smaller than gnathopod 2 and subchelate, pereopod 3 merus longer than the
propodus, coxa 4 smaller than coxa 3, pereopod 7 basis linear, uropod 3 endopod and exopod subequal and telson
entire.
The Infraorders and Parvorders of the Senticaudata have already been discussed by Lowry & Myers (2013).
Within the Amphilochida (Fig. 13) there are four parvorders. The Maxillipiidira is a well-defined
monophyletic parvorder based on a laterally compressed body with short coxae, second antennae much longer than
the body and sixth pereopods extremely elongate. The Oedicerotidira is a small group of three families defined by
having many rows of robust or slender setae on pereopods 5 to 7. Within the parvorder two of the three families
(Paracalliopiidae and Exoedicerotidae) have an oblique setal row on the basal endite of maxilla 2 and coalesced
urosomites 2 and 3. The six families of the Eusiridira have pereopods 5 to7 progressively longer, uropods 1 and 2
with lanceolate rami and the telson is cleft. The Amphilochidira is a large weakly supported monophyletic
parvorder with three superfamilies and 24 families. It is defined by having the carpus of pereopod 4 shorter than the
propodus and pereopods 5 to 7 of similar length. In addition the molar is generally small, non-triturative or absent,
gnathopod 1 is usually simple or chelate and the uropod 2 endopod is generally longer than exopod. Within the
Lysianassida (Fig. 14) the Synopiidira have a greatly elongated second antenna in the adult males, often urosomite
1 is free and 2–3 are coalesced and the telson is always cleft. The Haustoriidira have dorsoventrally flattened
bodies and pereopods 5–7 with many rows of facial or marginal robust and slender setae. The Lysianassidira have
the head deeper than long, generally a well-developed callynophore, gnathopod 2 mitten-shaped (may be poorly
developed in basal taxa) and with a long ischium, uropods 1 and 2 often with embedded setae and uropod 3
endopod generally shorter than exopod and two-articulate.
Tree discussion
The amphipod tree has three major clades: one of which includes the basal pseudingolfiellideans, one of which
includes the hyperiideans and the third that contains the remainder of the Amphipoda (Fig. 11). The
pseudingolfiellidean clade contains only the Pseudingolfiellidae, a small freshwater family known only from South
America and some subantarctic islands. It is apparently the closest living relative of the Ingolfiellida. The
hyperiidean clade, contains the Physosomata and Physocephalata of the well-supported Hyperiidea. It is a sister to
all other amphipods except the Pseudingolfiellida. The pagetinidiran and colomastigidiran parvorders that form the
relatively well-supported Colomastigidea and are both sister taxa to the hyperiopsideans. The hyperiopsidiran and
podosiridiran parvorders form the not as well-supported Hyperiopsidea which are planktonic/pelagic. In the third
clade there are two suborders, the Senticaudata (see Lowry & Myers 2013) and the Amphilochidea. There are two
infraorders in the Amphilochidea, the Amphilochida and the Lysianassida both of which are monophyletic.
Current ingolfiellidans are highly derived interstitial crustaceans with a worm-like body, vestigial coxae, etc. It
is possible that extinct ingolfiellidans with a more generalised form, epigean habit, large coxae, pedunculate eyes,
etc., gave rise to true amphipods like the amphilochids.
There are six suborders, 13 infraorders and 23 parvorders among the Amphipoda. We consider the Ingolfiellida
to be a sister taxon to the Amphipoda. Amphipods are broadly differentiated from other peracaridans by the sessile
eyes, no carapace, coxal gills and a urosome of three segments with epimera and with three uropods. Ingolfiellida
are similar to Amphipoda in that they have no carapace and coxal gills, but they have vestigial pedunculate eyes, a
pleosome of six segments without epimera and with reduced pleopod/uropod appendages. These significant
characters justify the exclusion of the Ingolfiellida from the Amphipoda. Accordingly, we establish the new order
Ingolfiellida. Verheye et al. (2015), carried out an 18S rDNA analysis, using a very small selection of amphipod
species. Ingolfiella tabularis represented the Ingolfiellida and was basal in their maximum likelihood tree.
The six suborders represent the broadest evolutionary groups among the amphipods. Basal
pseudingolfiellideans are benthic free-living freshwater amphipods with reduced discontiguous coxae and vestigial
epimeral plates. Basal colomastigideans are benthic, with short coxae and subcylindrical bodies, free-living or

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commensals. Basal hyperiopsideans are pelagic or benthic. One group, the hyperiopsoids are at least superficially
similar to hyperiidean amphipods. At some stage, apparently relatively early in the history of the amphipods, a
basal benthic amphipod moved up into the pelagic realm and diversified into a large, morphologically diverse
group of predators (large-eyed Physocephalata) and larval parasites (small-eyed Physosomata) on jellyfish (Laval
1980, Madin & Harbison 1977 and Harbison, Biggs & Madin 1977). They retain entire telsons. Laval (1980)
suggested that they basically traded one benthic surface for another.
Another group, the senticaudates, successfully invaded the shallow-water marine environment and world-wide
freshwater and terrestrial environments. These taxa have completely lost the callynophore and developed apical
robust setae on the rami of uropods 1 and 2. The males developed large gnathopods for mate-guarding (Conlan
1991).
The Amphilochidea form a large diverse group of 88 families in seven parvorders. At the base of the
Amphilochidira are the Amphilochoidea which are near in form to the hypothesized original amphipod type. While
the Amphilochoidea are not basal in our cladogram, they have retained many plesiomorphic attributes while other
branches have diversified and evolved into the manifold forms extant today. We believe that the ancestral
amphipod was a clinger in anastomoses that used the reflexed urosome as a means of propulsion. The stiffened
uropods provided grip on the substrate for the propulsive force. Clinging to anastomoses is best attained if the
opposing limbs are close together. This explains the characteristic features of an amphipod—presence of uropods,
reflexed abdomen and laterally flattened form. In a laterally flattened animal, the coxae can increase in size without
it having a detrimental effect on locomotion. The enlarged coxae can then act as shields protecting body parts and
in females, eggs. We therefore consider enlarged coxae to be a plesiomorphic character state. Amphilochids have
no special reproductive modifications such as brush setae, elongate second antennae or plumose setae on the rami
of the third uropods, often found in more advanced amphilochideans. Strategic reproductive character states such
as brush setae and elongate second antennae first appear among the Eusiroidea and the Oedicerotoidea.
But within the Amphilochidea there is evidence of evolution into major habitats. Within the Synopiidira
dexaminoids have developed the swimming male reproductive strategy as have the soft bottom suspension feeding
ampeliscids in the Synopioidea; iphimediids live among corals and in anastomoses etc. and also swim actively in
both sexes; haustoriidirans live in shallow water soft bottom habitats or high energy sand habitats and also take
advantage of the swimming male reproductive strategy to enhance their gene mixing.
The majority of highly derived lysianassidiran amphipods moved into a new habitat for the Amphipoda as
scavengers on dead animals. They retain the swimming male reproductive strategy, but usually have a callynophore
in the male and female for detecting odours in the water column, they developed a smooth slicing incisor and a
tongue-like molar and the setal-teeth on maxilla 1 modified into a fork-like arrangement in highly derived
scavengers such as the uristids. The alicelloid amphipods are deep water scavengers which retain the lacinia
mobilis on both mandibles, a 7/4 setal-tooth arrangement and an entire telson. The gigantic alicellids have a
gammaroid-like calceolus uncharacteristic of lysianassidirans. At least two species of Valettidae occur in the
alimentary canal of deep sea holothurians (Thurston 1989b). Parargissids are uncharacteristic lysianassidirans
which share asymmetrical palps on maxilla 1 with hyperiopsids, but the legs are atypical of lysianassidirans, while
podoprionids have highly serrated incisor uncharacteristic of lysianassidirans. Stegocephaloids are the sister taxa to
the remaining lysianassidirans. According to Berge & Vader (2001) stegocephalids appear to be micro-predators
often in association with hydroids and bryozoans (Sars 1891) and with sponges and tunicates (Vader 1984), but
they are also reported as scavengers (Lowry & Stoddart 1995b, Berge & Vader 2003).
The remaining members of the Lysianassidira are characterised by a mitten-shaped second gnathopod and a
left lacinia mobilis only. One group, the Lysianassoidea, is made up almost entirely of scavengers often with a type
3 lysianassoid calceolus and a cleft telson. Several families, the Adeliellidae, Amaryllididae, Cyphocarididae
(pelagic predators) Cebocaridae and Opisidae are not scavengers. No Aristioidea are scavengers. Families such as
the thoriellids, wandinids, acidostomatids, trischizostomatids and aristiids appear to be associated with other
invertebrates, or in the case of the trischizostomatids, epiparasites on sharks.
Discussion
This is first ever attempt at a phylogenetic classification of the entire order Amphipoda. The order is well known
for its morphological convergence in unrelated lineages occupying similar environments (Barnard & Karaman

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AMPHIPODA PHYL OGENY AND CLASSIFICATION
1975; Bousfield 1977; Barnard & Barnard 1983). This may explain why no attempt has previously been made at a
complete phylogenetic classification of the Amphipoda.
In this study, many of the clades are supported by several strong synapomorphies, but some inevitably are
more weakly supported. Our aim in this work was to provide a complete classification, so we did not have the
option of ignoring weakly supported clades.
We do not stand over the exact relationship of each and every family one to another. Further work is necessary
to clarify some of these relationships. Our main aim was to provide a framework classification to superfamily level.
Most of the higher taxa (above family level) can be clearly diagnosed. Our classification of the Amphipoda is a
hypothesis for future testing by both morphological and molecular analyses.
While many morphological and molecular studies have produced conflicting results, some show good
agreement. Good conformity is shown between the present work and the molecular study of Hurt et al. (2013). The
physomatidiran part of our hyperiidean tree shows the lanceoloids and the scinoids as two distinct lines of
evolution and this is in agreement with Hurt et al. (2013). Like their study, the present work also shows a primarily
phronimid—phrosinid clade and a vibiliid—cyllopodid clade. Another example, the good conformity between the
results of a morphological study (Myers & Lowry 2003) and a molecular study (Ito et al. 2011) in the case of the
caprellid amphipods, was described earlier in this paper.
Classification
A full classification of the Ingolfiellida and the Amphipoda is provided for completeness and to indicate at a glance
changes in higher taxa (Table 1). Families are listed in alphabetical order for ease of use.
TABLE 1. Classification of the Ingolfiellida and the Amphipoda including the Senticaudata (Lowry & Myers 2013).
Order Ingolfiellida
Suborder Ingolfiellidea
Infraorder Ingolfiellidamorpha
Parvorder Ingolfiellidira
Superfamily Ingolfielloidea
Family Ingolfiellidae
Parvorder Metaingolfiellidira
Superfamily Metaingolfielloidea
Family Metaingolfiellidae
Order Amphipoda
Suborder Pseudingolfiellidea
Infraorder Pseudingolfiellida
Parvorder Pseudingolfiellidira
Superfamily Pseudingolfielloidea
Family Pseudingolfiellidae
Suborder Hyperiidea
Infraorder Physosomata
Parvorder Physosomatidira
Superfamily Lanceoloidea
Family Chuneolidae
Family Lanceolidae
Family Megalanceolidae
Family Metalanceolidae
Family Microphasmidae
Family Mimonecteolidae
Family Prolanceolidae
Superfamily Scinoidea
Family Archaeoscinidae
Family Microscinidae
Family Mimonectidae
Family Mimoscinidae
Family Scinidae
Infraorder Physocephalata
Parvorder Physocephalatidira
Superfamily Vibilioidea
Family Cyllopodidae
Family Paraphronimidae
Family Vibiliidae
Superfamily Cystisomatoidea
Family Cystisomatidae
Superfamily Phronimoidea
Family Bougisidae
Family Dairellidae
Family Hyperiidae
Family Iulopididae
Family Lestrigonidae
Family Phronimidae
Family Phrosinidae
Superfamily Platysceloidea
Family Amphithyridae
Family Anapronoidae
Family Brachyscelidae
Family Eupronoidae
Family Lycaeidae
Family Lycaeopsidae
Family Oxycephalidae
Family Parascelidae
Family Platyscelidae
Family Pronoidae
Family Thamneidae

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Family Tryphanidae
Suborder Colomastigidea
Infraorder Colomastigida
Parvorder Colomastigidira
Superfamily Colomastigoidea
Family Colomastigidae
Parvorder Pagetinidira
Superfamily Pagetinoidea
Family Pagetinidae
Suborder Hyperiopsidea
Infraorder Hyperiopsida
Parvorder Hyperiopsidira
Superfamily Hyperiopsoidea
Family Hyperiopsidae
Family Vitjazianidae
Parvorder Podosiridira
Superfamily Podosiroidea
Family Podosiridae
Suborder Senticaudata
Infraorder Carangoliopsida
Parvorder Carangoliopsidira
Superfamily Carangoliopsoidea
Family Carangoliopsidae
Family Kairosidae
Infraorder Talitrida
Parvorder Talitridira
Superfamily Biancolinoidea
Family Biancolinidae
Superfamily Caspicoloidea
Family Caspicolidae
Superfamily Kurioidea
Family Kuriidae
Family Tulearidae
Superfamily Talitroidea
Family Ceinidae
Family Chiltoniidae
Family Dogielinotidae
Family Eophliantidae
Family Hyalellidae
Family Hyalidae
Subfamily Hyacheliinae
Subfamily Hyalinae
Family Najnidae
Family Phliantidae
Family Plioplateidae
Family Talitridae
Family Temnophliantidae
Infraorder Corophiida
Parvorder Corophiidira
Superfamily Aoroidea
Family Aoridae
Family Unciolidae
Subfamily Acuminodeutopinae
Subfamily Unciolinae
Superfamily Cheluroidea
Family Cheluridae
Superfamily Chevalioidea
Family Chevaliidae
Superfamily Corophioidea
Family Ampithoidae
Subfamily Ampithoinae
Subfamily Exampithoinae
Family Corophiidae
Subfamily Corophiinae
Tri b e Corophiini
Tri b e Haplocheirini
Tri b e Paracorophiini
Subfamily Protomedeiinae
Parvorder Caprellidira
Superfamily Aetiopedesoidea
Family Aetiopedesidae
Family Paragammaropsidae
Superfamily Caprelloidea
Family Caprellidae
Subfamily Caprellinae
Subfamily Paracercopinae
Subfamily Phtisicinae
Family Caprogammaridae
Family Cyamidae
Family Dulichiidae
Family Podoceridae
Superfamily Isaeoidea
Family Isaeidae
Superfamily Microprotopoidea
Family Australomicroprotopidae
Family Microprotopidae
Superfamily Neomegamphopoidea
Family Neomegamphopidae
Family Priscomilitariidae
Superfamily Photoidea
Family Ischyroceridae
Subfamily Bonnierellinae
Subfamily Ischyrocerinae
Tri b e Ischyrocerini
Tri b e Siphonoecetini
Family Kamakidae
Subfamily Aorchinae
Subfamily Kamakinae
Family Photidae
Superfamily Rakirooidea
Family Rakiroidae
Infraorder Hadziida
Parvorder Hadziidira
Superfamily Hadzioidea
Family Crangoweckeliidae
Family Eriopisidae
Family Gammaroporeiidae
Family Hadziidae
Family Maeridae
Family Melitidae
Family Metacrangonyctidae
Family Nuuanuidae

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AMPHIPODA PHYL OGENY AND CLASSIFICATION
Superfamily Calliopioidea
Family Calliopiidae
Family Cheirocratidae
Family Hornelliidae
Family Megaluropidae
Family Pontogeneiidae
Infraorder Bogidiellida
Parvorder Bogidiellidira
Superfamily Bogidielloidea
Family Artesiidae
Family Bogidiellidae
Family Salentinellidae
Infraorder Gammarida
Parvorder Crangonyctidira
Superfamily Allocrangonyctoidea
Family Allocrangonyctidae
Family Crymostygiidae
Family Dussartiellidae
Family Pseudoniphargidae
Family Kergueleniolidae
Superfamily Crangonyctoidea
Family Austroniphargidae
Family Chillagoeidae
Family Crangonyctidae
Family Giniphargidae
Family Kotumsaridae
Family Neoniphargidae
Family Niphargidae
Family Paracrangonyctidae
Family Paramelitidae
Family Perthiidae
Family Pseudocrangonyctidae
Family Sandroidae
Family Sternophysingidae
Family Uronyctidae
Parvorder Gammaridira
Superfamily Gammaroidea
Family Acanthogammaridae
Family Anisogammaridae
Family Baikalogammaridae
Family Bathyporeiidae
Family Behningiellidae
Family Falklandellidae
Family Gammaracanthidae
Family Gammarellidae
Family Gammaridae
Family Iphigenellidae
Family Luciobliviidae
Family Macrohectopidae
Family Mesogammaridae
Family Micruropodidae
Family Pachyschesidae
Family Pallaseidae
Family Paraleptamphopidae
Family Phreatogammaridae
Family Pontogammaridae
Family Sensonatoridae
Family Typhlogammaridae
Family Zaramillidae
Incertae Sedis
Family Iciliidae [Senticaudata]
Family Sanchoidae [Senticaudata]
Suborder Amphilochidea
Infraorder Amphilochida
Parvorder Maxillipiidira
Superfamily Maxillipioidea
Family Maxillipiidae
Parvorder Oedicerotidira
Superfamily Oedicerotoidea
Family Exoedicerotidae
Family Oedicerotidae
Family Paracalliopiidae
Parvorder Eusiridira
Superfamily Eusiroidea
Family Bateidae
Family Eusiridae
Family Miramarassidae
Family Thurstonellidae
Superfamily Liljeborgioidea
Family Liljeborgiidae
Subfamily Idunellinae
Subfamily Liljeborgiinae
Family Pseudamphilochidae
Parvorder Amphilochidira
Superfamily Amphilochoidea
Family Amphilochidae
Family Bolttsiidae
Family Cressidae
Family Cyproideidae
Family Didymocheliidae
Family Nihotungidae
Family Pleustidae
Subfamily Atylopsinae
Subfamily Austropleustinae
Subfamily Dactylopleustinae
Subfamily Eosymtinae
Subfamily Mesopleustinae
Subfamily Neopleustinae
Subfamily Parapleustinae
Subfamily Pleusirinae
Subfamily Pleustinae
Subfamily Pleustoidinae
Subfamily Pleusymtinae
Subfamily Stenopleustinae
Family Sebidae
Family Seborgiidae
Family Stenothoidae
Superfamily Leucothoidea
Family Leucothoidae
Superfamily Iphimedioidea
Family Acanthonotozomatidae
Family Acanthonotozomellidae

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Family Amathillopsidae
Subfamily Amathillopsinae
Subfamily Cleonardopsinae
Subfamily Parepimeriinae
Family Dikwidae
Family Epimeriidae
Family Iphimediidae
Family Lafystiidae
Family Laphystiopsidae
Family Ochlesidae
Family Odiidae
Family Sicafodiidae
Family Stilipedidae
Subfamily Astryinae
Subfamily Alexandrellinae
Subfamily Stilipedinae
Family Vicmusiidae
Infraorder Lysianassida
Parvorder Synopiidira
Superfamily Dexaminoidea
Family Atylidae
Subfamily Anatylinae
Subfamily Atylinae
Subfamily Lepechinellinae
Subfamily Nototropiinae
Family Dexaminidae
Subfamily Dexamininae
Subfamily Dexaminoculinae
Subfamily Polycheriinae
Subfamily Prophliantinae
Family Melphidippidae
Family Pardaliscidae
Superfamily Synopioidea
Family Ampeliscidae
Family Argissidae
Family Synopiidae
Parvorder Haustoriidira
Superfamily Haustorioidea
Family Cheidae
Family Condukiidae
Family Haustoriidae
Family Ipanemidae
Family Otagiidae
Family Phoxocephalidae
Subfamily Harpiniinae
Subfamily Phoxocephalinae
Family Phoxocephalopsidae
Family Platyischnopidae
Family Pontoporeiidae
Family Priscillinidae
Family Sinurothoidae
Family Urohaustoriidae
Family Urothoidae
Family Zobrachoidae
Parvorder Lysianassidira
Superfamily Alicelloidea
Family Alicellidae
Family Parargissidae
Family Podoprionidae
Family Valettidae
Family Valettiopsidae
Family Vemanidae
Superfamily Stegocephaloidea
Family Stegocephalidae
Subfamily Andaniexinae
Subfamily Andaniopsinae
Subfamily Bathystegocephalinae
Subfamily Parandaniinae
Subfamily Stegocephalinae
Superfamily Lysianassoidea
Family Adeliellidae
Family Amaryllididae
Subfamily Amaryllidinae
Subfamily Vijayiinae
Family Cebocaridae
Family Cyclocaridae
Family Cyphocarididae
Family Eurytheneidae
Family Hirondelleidae
Family Lysianassidae
Subfamily Lysianassinae
Subfamily Waldeckiinae
Family Opisidae
Family Scopelocheiridae
Subfamily Paracallisominae
Subfamily Scopleocheirinae
Family Tryphosidae
Family Uristidae
Superfamily Aristioidea
Family Acidostomatidae
Family Ambasiidae
Family Aristiidae
Family Conicostomatidae
Family Derjugianidae
Family Endevouridae
Family Izinkalidae
Family Kergueleniidae
Family Lepidepecreellidae
Family Pakynidae
Family Sophrosynidae
Family Thoriellidae
Family Trischizostomatidae
Family Wandinidae

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AMPHIPODA PHYL OGENY AND CLASSIFICATION
Superorder Peracarida
Order Ingolfiellida Hansen, 1903 stat. nov.
Diagnosis. Head with vestigial pedunculate eyes. Gnathopods 1–2 eucarpochelate. Pleosome with 6 relatively
undifferentiated segments, without epimera, with reduced pleopods and uropods.
Remarks. The important apomorphies that define the order are the eucarpochelate gnathopods and the six-
segmented pleosome. The eucarpochelate gnathopods occur when the propodus is reduced and combined with the
dactylus to form a dactylar complex, both close onto an expanded carpus. This situation is approached in
amphipods in some species of the ischyrocerid genus Ericthonius and the aorid genus Grandidierella, but this is
considered as convergent.
Suborder Ingolfiellidea Hansen, 1903 (Ruffo, 1970)
Diagnosis. As for order.
Infraorder Ingolfiellidamorpha Hansen, 1903 stat. nov.
Diagnosis. As for suborder.
Parvorder Ingolfiellidira Hansen, 1903 stat. nov.
Diagnosis. Body vermiform. Pleopods modified. Uropod 3 uniramous.
Included superfamilies. Ingolfielloidea Hansen, 1903 stat. nov.
Superfamily Ingolfielloidea Hansen, 1903 stat. nov.
Diagnosis. As for parvorder.
Included families. Ingolfiellidae Hansen, 1903.
Family Ingolfiellidae Hansen, 1903
Included genera. Ingolfiella Hansen, 1903; Proleleupia Vonk & Schram, 2003; Rapaleleupia Vonk & Schram,
2007; Stygobarnardia Ruffo, 1985; Trogloleleupia Ruffo, 1975.
Parvorder Metaingolfiellidira Ruffo, 1969 stat. nov.
Diagnosis. Mandible palp present. Body subcylindrical. Pleopods well-developed. Uropod 3 biramous.
Included superfamilies. Metangolfielloidea Ruffo, 1969 stat. nov.
Superfamily Metaingolfielloidea Ruffo, 1969 stat. nov.
Diagnosis. As for parvorder.
Included families. Metaingolfiellidae Ruffo, 1969.

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Family Metaingolfiellidae Ruffo, 1969
Included genera. Metaingolfiella Ruffo, 1969.
Order Amphipoda Latreille, 1816
Diagnosis. Head with sessile eyes. Carapace absent. Coxal gills present. Abdomen with 6 segments, differentiated
into a pleosome with 3 segments, with epimera (vestigial in Pseudingolfiella) and urosome with 3 segments. Last
three pairs of pleopods converted into uropods, pleopods and uropods usually well-developed.
Suborder Pseudingolfiellidea Lowry & Myers, 2012a stat. nov.
Diagnosis. Body vermiform. Maxilliped palps well developed; ischial endites small, not longer than palp article 2.
Coxae greatly reduced, discontiguous. Gnathopods subchelate. Pleopods reduced. Epimera vestigial. Urosome
poorly defined. Uropods 1–2 rami without apical robust setae.
Included infraorders. Pseudingolfiellida Lowry & Myers, 2012a stat. nov.
Remarks. The urosome is weakly differentiated but the vestigial uropods are notched into the urosome. Lowry
& Myers (2013) excluded Pseudingolfiellidae from the Senticaudata based on the lack of apical robust setae on
uropods 1 and 2.
Infraorder Pseudingolfiellida Lowry & Myers, 2012a stat. nov.
Diagnosis. As for Suborder.
Included parvorders. Pseudingolfiellidira Lowry & Myers, 2012a stat. nov.
Parvorder Pseudingolfiellidira Lowry & Myers, 2012a stat. nov.
Diagnosis. As for Suborder.
Included superfamilies. Pseudingolfielloidea Lowry & Myers, 2012a stat. nov.
Superfamily Pseudingolfielloidea Lowry & Myers, 2012a stat. nov.
Diagnosis. As for Suborder.
Included families. Pseudingolfiellidae Lowry & Myers, 2012a stat. nov.
Family Pseudingolfiellidae Lowry & Myers, 2012a
Included genera. Pseudingolfiella Noodt, 1965.
Suborder Hyperiidea H. Milne Edwards, 1830
Diagnosis. Maxilliped without palps. Urosomite 1 free, 2–3 coalesced. Uropods 1–2 rami without apical robust
setae.
Included infraorders. Physosomata Pirlot, 1929; Physocephalata Bowman & Gruner, 1973.

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AMPHIPODA PHYL OGENY AND CLASSIFICATION
Infraorder Physosomata Pirlot, 1929 (Bowman & Gruner, 1973)
Diagnosis. Eyes small or absent (except Vibiliidae). Urosomites 2–3 partially fused. Male genital papillae present.
Included parvorders. Physosomatidira Pirlot, 1929 stat. nov.
Remarks. Zeidler (2006) indicates two new characters for the Physosomata: ‘a partial ventral suture between
urosomites 2 & 3 in all specimens and the presence of male genital papillae’.
Parvorder Physosomatidira Pirlot, 1929 stat. nov.
Diagnosis. Molar with patch of fine setae. Maxilla 1 basal endite present. Maxilliped basal endite separate.
Included superfamilies. Lanceoloidea Bovallius, 1887 (Bowman & Gruner, 1973); Scinoidea Stebbing, 1888
(Bowman & Gruner, 1973).
Superfamily Lanceoloidea Bovallius, 1887 (Bowman & Gruner, 1973)
Diagnosis. Head deeper than long. Antenna 2 adult male flagellum not greatly elongate. Maxilla 1 basal endite
covered in setae. Pereonites separate. Pereopod 4 coxa subequal or smaller than coxa 3.
Included families. Chuneolidae Woltereck, 1909; Lanceolidae Bovallius, 1887; Megalanceolidae Zeidler,
2009; Metalanceolidae Zeidler, 2009; Microphasmidae Stephensen & Pirlot, 1931; Mimonecteolidae Zeidler, 2009;
Prolanceolidae Zeidler, 2009.
Family Chuneolidae Woltereck, 1909
Included genera. Chuneola Woltereck, 1909.
Family Lanceolidae Bovallius, 1887
Included genera. Lanceola Say, 1818b; Scypholanceola Woltereck, 1905.
Family Megalanceolidae Zeidler, 2009
Included genera. Megalanceola Pirlot, 1935; Megalanceoloides Zeidler, 2009.
Family Metalanceolidae Zeidler, 2009
Included genera. Metalanceola Pirlot, 1931.
Family Microphasmidae Stephensen & Pirlot, 1931
Included genera. Microphasma Woltereck, 1909; Microphasmoides Vinogradov, 1960.
Family Mimonecteolidae Zeidler, 2009
Included genera. Mimonecteola Woltereck, 1909.

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Family Prolanceolidae Zeidler, 2009
Included genera. Prolanceola Woltereck, 1907.
Superfamily Scinoidea Stebbing, 1888 (Bowman & Gruner, 1973)
Diagnosis. Mandible lacinia mobilis present only on left side; palp absent. Gnathopods 1–2 simple (except in some
Scinidae).
Included families. Archaeoscinidae Stebbing, 1904; Microscinidae Zeidler, 2012; Mimonectidae Bovallius,
1885; Mimoscinidae Zeidler, 2012; Scinidae Stebbing, 1888.
Remarks. Zeidler (2012) rejects the family status of Proscinidae Pirlot, 1933a and transfers species of the
genus Proscina to Mimonectes and Mimoscina (see Zeidler 2012).
Family Archaeoscinidae Stebbing, 1904
Included genera. Archaeoscina Stebbing, 1904; Paralanceola K.H. Barnard, 1930.
Remarks. The placement of this family in the Scinoidea is contentious. It differs from all other Scinoidea in
having mandibular palps. Vinogradov, Volkov & Semenova, 1982 placed it in the superfamily Archaeoscinoidea.
Zeidler (pres. com.) believes that this family should be placed in the Lanceoloidea.
Family Microscinidae Zeidler, 2012
Included genera. Microscina Zeidler, 2012.
Family Mimonectidae Bovallius, 1885
Included genera. Cheloscina Shih & Hendrycks, 1996; Mimonectes Bovallius, 1885; Pseudomimonectes
Vinogradov, 1960.
Family Mimoscinidae Zeidler, 2012
Included genera. Mimoscina Pirlot, 1933a.
Family Scinidae Stebbing, 1888
Included genera. Scina Prestandrea, 1833; Acanthoscina Vosseler, 1901; Ctenoscina Wagler, 1926; Spinoscina
Bowman & Gruner, 1973.
Infraorder Physocephalata Bowman & Gruner, 1973
Diagnosis (after Bowman & Gruner 1973). Head large, longer than pereonite 1. Eyes large occupying most of head
(small to moderately large in Vibiliidae and Bougisidae). Maxilla 1 without basal endite (inner lobe) (except
Cyllopodidae, Paraphronimidae). Maxilliped basal endites (inner lobes) completely fused (except Lycaeopsidae).
Male genital papillae absent or very small (Zeidler 2006).
Included parvorders. Physocephalatidira Bowman & Gruner, 1973 stat. nov.

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AMPHIPODA PHYL OGENY AND CLASSIFICATION
Parvorder Physocephalatidira Bowman & Gruner, 1973 stat. nov.
Diagnosis. As for Infraorder.
Included superfamilies. Phronimoidea Dana, 1852 (Bowman & Gruner, 1973); Platysceloidea Spence Bate,
1862 (Bowman & Gruner, 1973); Vibilioidea Dana, 1852 (Bowman & Gruner, 1973).
Remarks. Bowman & Gruner (1973) erected a monotypic superfamily (Lycaeopsoidea) for the family
Lycaeopsidae Chevreux, 1913. In our analysis the Lycaeopsidae is a sister taxon to the Tryphanidae and both
together form a sister group to the Pronoidae within the superfamily Platysceloidea. Zeidler (pers. com.) has
questioned this relationship.
Superfamily Vibilioidea Dana, 1852 (Bowman & Gruner, 1973)
Diagnosis. Antenna 1 inserted on anterior surface of head. Antenna 2 flagellum not elongated. Maxilliped basal
endite fused medially. Body subcylindrical or laterally compressed, coxae small. Pereopods 5, 6 similar in length,
pereopod 7 shorter.
Included families. Cyllopodidae Bovallius, 1887; Paraphronimidae Bovallius, 1887; Vibiliidae Dana, 1852.
Family Cyllopodidae Bovallius, 1887
Included genera. Cyllopus Dana, 1853.
Family Paraphronimidae Bovallius, 1887
Included genera. Paraphronima Claus, 1879.
Family Vibiliidae Dana, 1852
Included genera. Vibilia H. Milne Edwards, 1830; Vibilioides Chevreux, 1905b.
Superfamily Phronimoidea Dana, 1852 (Bowman & Gruner, 1973)
Diagnosis. Antenna 1 peduncular article 1 longer than article 2. Antenna 1 inserted on anterior surface of head.
Mandible lacinia mobilis present on both sides. Maxilla 1 basal endite absent. Pereopod 4 coxa fused to pereonite
or subequal to that of pereopod 3. Urosomite 1 subequal or longer than urosomite 2 or 2–3. Uropod 2 endopod
subequal to or shorter than exopod or absent.
Included families. Bougisidae Zeidler, 2004; Cystisomatidae Willemöes-Suhm, 1875; Dairellidae Bovallius,
1887; Hyperiidae Dana, 1852; Iulopididae Zeidler, 2004; Lestrigonidae Zeidler, 2004; Phronimidae Dana, 1852;
Phrosinidae Dana, 1852.
Remarks. A new superfamily, Cystisomatoidea, was proposed by Zeidler (2003) to accommodate the family
Cystisomatidae, primarily based on the unique method of brooding the young. We did not use this character in our
analysis. In our analysis the Cystisomatidae is a sister taxon to the Dairellidae and deeply embedded in the
Phronimoidea. It should be noted that Browne et al. (2007) and Hurt et al. (2013) found support for placing the
Cystisomatidae with the Physosomata.
Family Bougisidae Zeidler, 2004

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Included genera. Bougisia Laval, 1966.
Family Cystisomatidae Willemöes-Suhm, 1875
Included genera. Cystisoma Guérin-Méneville, 1842.
Family Dairellidae Bovallius, 1887
Included genera. Dairella Bovallius, 1887.
Family Hyperiidae Dana, 1852
Included genera. Hyperia Latreille, 1823; Hyperiella Bovallius, 1887; Hyperoche Bovallius, 1887; Laxohyperia
Vinogradov & Volkov, 1982; Pegohyperia K.H. Barnard, 1931; Prohyperia Zeidler, 2015; Themisto Guérin, 1825.
Remarks. According to Bowman et al. (1982) “The amphipod genus Themisto Guérin-Méneville, 1825, was
replaced with Euthemisto by Bovallius (1887) because it was a junior homonym of the nudibranch Themisto Oken
(1815). Parathemisto Boeck (1870) and Themisto Guérin-Méneville were combined under Themisto by Stephensen
(1924), corrected by K.H. Barnard (1930) to Parathemisto, the name currently used by most zoologists. Volume 3
of Okens Lehrbuch der Naturgeschichte (1815) was placed, however, on the Official List of Rejected Works by
Opinion 417 of the International Commission of Zoological Nomenclature (1956), which ruled that no name
published in Oken's volume 3 acquired the status of availability by reason of having been so published.
Consequently the name Themisto, first made available by Guérin and a senior synonym of Parathemisto, is the
valid name of the amphipod genus”.
Family Iulopididae Zeidler, 2004
Included genera. Iulopis Bovallius, 1887.
Family Lestrigonidae Zeidler, 2004
Included genera. Hyperietta Bowman, 1973; Hyperioides Chevreux, 1900; Hyperionyx Bowman, 1973;
Lestrigonus H. Milne Edwards, 1830; Phronimopsis Claus, 1879; Themistella Bovallius, 1887.
Family Phronimidae Dana, 1852
Included genera. Phronima Latreille, 1802; Phronimella Claus, 1871.
Family Phrosinidae Dana, 1852 (Stebbing, 1888)
Included genera. Anchylomera Milne Edwards, 1830; Phrosina Risso, 1822; Primno Guérin-Méneville, 1836.
Superfamily Platysceloidea Spence Bate, 1862 (Bowman & Gruner, 1973)
Diagnosis. Eyes occupying most of lateral surface of the head. Mandible molar vestigial or absent. Maxilla 1 basal
endite absent. Maxilliped palps absent. Urosomite 1 shorter than urosomites 2 or 3.

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Included families. Amphithyridae Zeidler, 2016; Anapronoidae Bowman & Gruner, 1973; Brachyscelidae
Stephensen, 1923; Eupronoidae Zeidler, 2016; Lycaeidae Claus, 1879; Lycaeopsidae Chevreux 1913;
Oxycephalidae Spence Bate, 1862; Parascelidae Bovallius, 1887; Platyscelidae Spence Bate, 1862; Pronoidae
Claus, 1879; Thamneidae Zeidler, 2016; Tryphanidae Boeck, 1871.
Remarks. The following character states occur in all family level taxa except the Lycaeopsidae: antenna 1
inserted on ventral surface of the head; antenna 2 articles folded on one another in zigzag fashion; pereopods 5–6
basis greatly enlarged.
Family Amphithyridae Zeidler, 2016
Included genera. Amphithyropsis Zeidler, 2016; Amphithyrus Claus, 1879; Paralycaea Claus, 1879.
Family Anapronoidae Bowman & Gruner, 1973
Included genera. Anapronoe Stephensen, 1925c.
Family Brachyscelidae Stephensen, 1923
Included genera. Brachyscelus Spence Bate, 1861.
Remarks. Zeidler (2016) reserved the Brachyscelidae for Brachyscelus and removed Thamneus to a new
monotypic family Thamneidae Zeidler, 2016.
Family Eupronoidae Zeidler, 2016
Included genera. Eupronoe Claus, 1879; Parapronoe Claus, 1879.
Family Lycaeidae Claus, 1879
Included genera. Lycaea Dana, 1852; Simorhynchotus Stebbing, 1888 (Fide Vinogradov et al. 1982).
Family Lycaeopsidae Chevreux, 1913
Included genera. Lycaeopsis Claus, 1879.
Family Oxycephalidae Spence Bate, 1862
Included genera. Calamorhynchus Streets, 1878; Cranocephalus Bovallius, 1890; Glossocephalus Bovallius,
1887; Leptocotis Streets, 1877; Oxycephalus H. Milne Edwards, 1830; Rhabdosoma White, 1847; Streetsia
Stebbing, 1888; Tullbergella Bovallius, 1887.
Family Parascelidae Bovallius, 1887
Included genera. Euscelus Claus, 1879; Parascelus Claus, 1879; Schizoscelus Claus, 1879; Thyropus Dana, 1852.
Family Platyscelidae Spence Bate, 1862

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Included genera. Hemityphis Claus, 1879; Paratyphis Claus, 1879; Platyscelus Spence Bate, 1862; Tetrathyrus
Claus, 1879.
Remarks. Zeidler (2016) removed Amphithyrus Claus, 1879 to Amphithyridae Zeidler, 2016.
Family Pronoidae Claus, 1879
Included genera. Pronoe Guérin-Méneville, 1836.
Remarks. Zeidler (2016) moved Eupronoe Claus, 1879 and Parapronoe Claus, 1879 to the Eupronoidae
Zeidler, 2016 and Paralycaea Claus, 1879 to Amphithyridae Zeidler, 2016.
Family Thamneidae Zeidler, 2016
Included genera. Thamneus Bovallius, 1887.
Family Tryphanidae Boeck, 1871
Included genera. Tryphana Boeck, 1871.
Suborder Colomastigidea Stebbing, 1899 stat. nov.
Diagnosis. Body subcylindrical. Maxilliped palps present. Urosomite 1 free, 2–3 coalesced. Uropods 1–2 rami
without apical robust setae.
Included infraorders. Colomastigida stat. nov.
Infraorder Colomastigida Stebbing, 1899 stat. nov.
Diagnosis. As for suborder.
Included parvorders. Colomastigidira Stebbing, 1899 stat. nov.; Pagetinidira K.H. Barnard, 1931 stat. nov.
Parvorder Colomastigidira Stebbing, 1899 stat. nov.
Diagnosis. Antenna 1 callynophore present. Mandible palp absent. Maxilla 1 palp absent. Gnathopod 1 simple.
Uropod 3 biramous.
Included superfamilies. Colomastigoidea Stebbing, 1899 stat. nov.
Superfamily Colomastigoidea Stebbing, 1899 stat. nov.
Diagnosis. As for parvorder.
Included families. Colomastigidae Stebbing, 1899
Family Colomastigidae Stebbing, 1899
Included genera. Colomastix Grube, 1861; Yulu m a r a J.L. Barnard, 1972a.

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Parvorder Pagetinidira K.H. Barnard, 1931 stat. nov.
Diagnosis. Antenna 1 callynophore absent. Mandible palp present. Maxilla 1 palp present. Gnathopod 1
subchelate. Uropod 3 uniramous.
Included superfamilies. Pagetinoidea K.H. Barnard, 1931 stat. nov.
Superfamily Pagetinoidea K.H. Barnard, 1931 stat. nov.
Diagnosis. As for parvorder.
Included families. Pagetinidae K.H. Barnard, 1931.
Family Pagetinidae K.H. Barnard, 1931
Included genera. Pagetina K.H. Barnard, 1931.
Suborder Hyperiopsidea Bovallius, 1886, stat. nov.
Diagnosis. Body laterally compressed with small coxae. Mandible molar triturative. Maxilliped palps well
developed. Pereopod 4 coxa without posteroventral lobe. Uropods 1–2 rami without apical robust setae.
Included infraorders. Hyperiopsida Bovallius, 1866 stat. nov.
Remarks. Basal hyperiopsideans are similar to physocephalatan hyperiideans, particularly vibilioids and
phronimoids.
Infraorder Hyperiopsida Bovallius, 1886, stat. nov.
Diagnosis. As for suborder.
Included parvorders. Hyperiopsidira Bovallius, 1866 stat. nov.; Podosiridira Lowry & Myers, 2012b stat.
nov.
Parvorder Hyperiopsidira Bovallius, 1886, stat. nov.
Diagnosis. Antenna 1 accessory flagellum present; with callynophore, weakly to well developed. Gnathopod 1
simple. Pereopod 3 coxa subequal to that of 4. Pereopod 5 coxa smaller than that of 4.
Included superfamilies. Hyperiopsoidea Bovallius, 1866 stat. nov.
Superfamily Hyperiopsoidea Bovallius, 1886, stat. nov.
Diagnosis. As for parvorder.
Included families. Hyperiopsidae Bovallius, 1886; Vitjazianidae Birstein & Vinogradov, 1955.
Family Hyperiopsidae Bovallius, 1886
Included genera. Hyperiopsis Sars, 1885; Protohyperiopsis Birstein & Vinogradov, 1955.

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Family Vitjazianidae Birstein & Vinogradov, 1955
Included genera. Vitjaziana Birstein & Vinogradov, 1955.
Parvorder Podosiridira Lowry & Myers, 2012b stat. nov.
Diagnosis. Antenna 1 accessory flagellum absent; without callynophore. Gnathopod 1 subchelate. Pereopod 3 coxa
larger than that of 4. Pereopod 5 coxa subequal to that of 4.
Included superfamilies. Podosiroidea Lowry & Myers, 2012b stat. nov.
Superfamily Podosiroidea Lowry & Myers, 2012b stat. nov.
Diagnosis. As for parvorder.
Included families. Podosiridae Lowry & Myers, 2012b
Family Podosiridae Lowry & Myers, 2012b
Included genera. Podosirus Bellan-Santini, 2007.
Suborder Senticaudata Lowry & Myers, 2013
(see Lowry & Myers, 2013)
Suborder Amphilochidea Boeck, 1871 stat. nov.
Diagnosis. Body laterally compressed or dorsoventrally flattened with large or occasionally small coxae. Antenna
2 peduncle with brush setae (weakly developed). Maxilliped palps well-developed. Pereopod 4 coxa smaller or
larger than that of 3. Uropods 1–2 rami without apical robust setae.
Included infraorders. Amphilochida Boeck, 1871 stat. nov.; Lysianassida Dana, 1849 stat. nov.
Remarks. The Amphilochidea is a monophyletic clade, although the defining character state (brush setae
present) is weak. All Amphilochidea have laterally compressed or dorsoventrally flattened bodies with large coxae
except the Pardaliscidae and Synopiidae which have laterally compressed bodies with relatively shortened coxae,
apparently a homoplasious character.
Infraorder Amphilochida Boeck, 1871 stat. nov.
Diagnosis. Body laterally compressed or dorsoventrally flattened. Mandible incisors dentate [except,
Sicafodiidae]. Coxae 1–4 overlapping [except Miramarassidae]. Pereopod 4 coxa subequal or larger than that of 3.
Pereopod 6 subequal or shorter than pereopod 7 or immensely elongate [except Dikwidae]; urosomites 1–3 free
[except Paracalliopiidae, Sebidae]. Uropod 3 exopod 1 articulate [except Didymocheliidae, Nihotungidae, Sebidae,
Stenothoidae].
Included parvorders. Amphilochidira Boeck, 1871 stat. nov.; Eusiridira Stebbing 1888 stat. nov.;
Maxillipiidira Ledoyer, 1973 stat. nov.; Oedicerotidira Lilljeborg, 1865b (Bousfield, 1979) stat. nov.
Remarks. The Amphilochida are monophyletic, but they are defined by rather weak apomorphic character
states: coxae 1–4 overlapping and uropod 3 exopod longer than peduncle. They can be diagnosed by the above
diagnostic characters with several exceptions. A number of small highly derived taxa (Nihotungidae,
Didymocheliidae, Dikwidae, Miramarassidae, Paracalliopiidae, Sebidae and Sicafodiidae) are anomalous. The
large family Stenothoidae has coxa 4 enlarged without a posteroventral lobe and uropod 3 with a 2-articulate
exopod. As a consequence it does not conform to the diagnosis.

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We place the family Eusiridae in the Amphilochida, but taxa such as Rhachotropis confound the diagnosis. We
believe that the family Eusiridae is too broadly defined.
Parvorder Maxillipiidira Ledoyer, 1973 stat. nov.
Diagnosis. Body dorsoventrally flattened. Pereopod 6 immensely elongate, flagellate.
Included superfamilies. Maxillipioidea Ledoyer, 1973 stat. nov.
Superfamily Maxillipioidea Ledoyer, 1973 stat. nov.
Diagnosis. As for parvorder.
Included families. Maxillipiidae Ledoyer, 1973.
Family Maxillipiidae Ledoyer, 1973
Included genera. Maxillipides Ledoyer, 1984; Maxillipius Ledoyer, 1973.
Parvorder Oedicerotidira Ledoyer, 1973, stat. nov.
Diagnosis. Pereopod 4 coxa with well-developed posteroventral lobe. Pereopod 5 coxa equilobate. Pereopod 7
immensely elongate. Pereopod 7 different in structure to pereopod 6. Telson entire.
Included superfamilies. Oedicerotoidea Ledoyer, 1973 stat. nov.
Superfamily Oedicerotoidea Lilljeborg, 1865b (Bousfield, 1979)
Diagnosis. As for parvorder.
Included families. Exoedicerotidae Barnard & Drummond, 1982a; Oedicerotidae Lilljeborg, 1865b;
Paracalliopiidae Barnard & Karaman, 1982.
Family Exoedicerotidae Barnard & Drummond, 1982a
Included genera. Bathyporeiapus Schellenberg, 1931; Exoediceroides Bousfield, 1983; Exoediceropsis
Schellenberg, 1931; Exoediceros Stebbing, 1899; Kanaloa J.L. Barnard, 1970; Methalimedon Schellenberg, 1931;
Metoediceropsis Dang, 1968; Metoediceros Schellenberg, 1931; Notoediceros Bousfield, 1983; Parhalimedon
Chevreux, 1906b; Patuki Cooper & Fincham, 1974; Vadosiapus Barnard & Thomas, 1988a.
Family Oedicerotidae Lilljeborg, 1865b
Included genera. Aborolobatea Ledoyer, 1984; Acanthostepheia Boeck, 1871; Aceroides G.O. Sars, 1892;
Americhelidium Bousfield & Chevrier, 1996; Ameroculodes Bousfield & Chevrier, 1996; Anoediceros Pirlot, 1932;
Arrhinopsis Stappers, 1911; Arrhis Stebbing, 1906; Bathymedon G.O. Sars, 1892; Carolobatea Stebbing, 1899;
Caviphaxus Ren, 1992; Chitonomandibulum Jo, 1990; Cornudilla Barnard & Karaman, 1991; Deflexilodes
Bousfield & Chevrier, 1996; Eochelidium Bousfield & Chevrier, 1996; Finoculodes J.L. Barnard, 1971;
Gulbarentsia Stebbing, 1894; Halicreion Boeck, 1871; Hartmanodes Bousfield & Chevrier, 1996; Hongkongvena
Hirayama, 1992; Imbachoculodes Kim, Hendrycks & Lee, 2012; Kroyera Spence Bate, 1857; Limnoculodes
Bousfield & Chevrier, 1996; Lopiceros J.L. Barnard, 1961; Machaironyx Coyle, 1980; Monoculodes Stimpson,

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1853; Monoculodopsis Ledoyer, 1973; Monoculopsis G.O. Sars, 1892; Oedicerina Stephensen, 1931; Oediceroides
Stebbing, 1888; Oediceropsis Lilljeborg, 1865a; Oediceros Krøyer, 1842; Orthomanus Kim, Hendrycks & Lee,
2012; Pacifoculodes Bousfield & Chevrier, 1996; Paramonoculopsis Alonso de Pina, 1997; Paraperioculodes
K.H. Barnard, 1931; Parexoediceros Bousfield, 1983; Paroediceroides Schellenberg, 1931; Paroediceros G.O .
Sars, 1892; Perioculodes G.O. Sars, 1892; Perioculopsis Schellenberg, 1925; Pontocrates Boeck, 1871;
Rostroculodes Bousfield & Chevrier, 1996; Sinoediceros Shen, 1955; Synchelidium G.O. Sars, 1892; Westwoodilla
Spence Bate, 1857.
Family Paracalliopiidae Barnard & Karaman, 1982
Included genera. Doowia Barnard & Drummond, 1987; Indocalliope Barnard & Karaman, 1982; Katocalliope
Barnard & Drummond, 1984; Paracalliope Stebbing, 1899; Yhi Barnard & Thomas, 1991.
Parvorder Eusiridira Stebbing, 1888 stat. nov.
Diagnosis. Body laterally compressed with large coxae. Antenna 1 callynophore weakly developed (1-field) or
absent. Mandible incisor dentate. Pereopods 5–7 progressively longer.
Included superfamilies. Eusiroidea Stebbing, 1888 (Bousfield, 1979); Liljeborgioidea Stebbing, 1899
(Bousfield, 1979).
Superfamily Eusiroidea Stebbing, 1888 (Bousfield, 1979)
Diagnosis. As for parvorder plus antenna 2 flagellum shorter than peduncle; mandible molar triturative.
Included families. Bateidae Stebbing, 1906; Eusiridae Stebbing, 1888; Miramarassidae Lowry, 2006;
Thurstonellidae Lowry & Zeidler, 2008.
Family Bateidae Stebbing, 1906
Included genera. Batea Müller, 1865.
Family Eusiridae Stebbing, 1888
Included genera. Cleonardo Stebbing, 1888; Eusirella Chevreux, 1908b; Eusirogenes Stebbing, 1904; Eusiropsis
Stebbing, 1897; Eusirus Krøyer, 1845; Harcledo J.L. Barnard, 1964c; Meteusiroides Pirlot, 1934; Pareusirogenes
Birstein & Vinogradov, 1955; Rhachotropis S.I. Smith, 1883; Sennaia Bellan-Santini, 1997; Triquetramana
Hendrycks & Conlan, 2003.
Remarks. Sennaia is listed in the World Amphipoda Database (Horton et al. 2017) in the Pontogeneiidae, but
the lack of apical robust setae on uropods 1 and 2 exclude it from the Senticaudata and it is here placed in the
Eusiridae as originally specified by Bellan-Santini 1997.
Family Miramarassidae Lowry, 2006
Included genera. Miramarassa Ortiz, Lalana & Lio, 1999.
Family Thurstonellidae Lowry & Zeidler, 2008

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Included genera. Thurstonella Lowry & Zeidler, 2008.
Superfamily Liljeborgioidea Stebbing, 1899
Diagnosis. As for parvorder plus gnathopod 1 present and uropod 3 endopod shorter than exopod.
Included families. Liljeborgiidae Stebbing, 1899; Pseudamphilochidae fam. nov.
Family Liljeborgiidae Stebbing, 1899
Included subfamilies. Idunellinae d’Udekem d’Acoz, 2010; Liljeborgiinae Stebbing, 1899.
Remarks. d’Udekem d’Acoz, 2010 established the subfamilies Idunellinae and Liljeborgiinae. Species of
Idunellinae have apical robust setae on the rami of uropods 1 and 2, species of Liljeborgiinae do not. This suggests
that the apical robust setae of Idunellinae are homoplasious.
Subfamily Idunellinae d’Udekem d’Acoz, 2010
Included genera. Idunella G.O. Sars, 1894; Listriella J.L. Barnard, 1959a; Sextonia Chevreux, 1920.
Subfamily Liljeborgiinae Stebbing, 1899 (d’Udekem d’Acoz, 2010)
Included genera. Isipingus Barnard & Karaman, 1987; Liljeborgia Spence Bate, 1862.
Remarks. d’Udekem d’Acoz (2010) provisionally considered Isipingus to be a subgenus of Liljeborgia.
Family Pseudamphilochidae fam. nov.
Pseudamphilochidae Barnard & Karman, 1982: 184 (invalid attribution to Schellenberg, 1931: not considered as new).—
Barnard & Karaman, 1991: 667.—De Broyer & Jażdżewski, 1993: 88.—Gutt, Sirenko, Arntz, Smirnov & De Broyer,
2000: 67.—De Broyer et al. 2007: 199.
Type genus. Pseudamphilochus Schellenberg, 1931.
Diagnosis (based on world family database). Body laterally compressed with large coxae. Head, eyes round.
Antenna 1 shorter than antenna 2; antenna 1 accessory flagellum absent, callynophore absent. Antennae 1–2
calceoli absent. Mandible molar vestigial or absent. Maxilla 1 basal endite with apical setae. Maxilliped outer plate
longer than palp article 1, not longer than palp article 2. Gnathopods 1–2 subchelate, carpus not produced along
posterior margin of propodus; propodus palms with row of robust setae along margin. Gnathopod 1 coxa large, not
hidden by 2. Uropods 1–2 rami without apical robust or embedded setae. Uropod 3 biramous; endopod shorter than
exopod. Telson laminar, moderately cleft.
Included genera. Pseudamphilochus Schellenberg, 1931.
Habitat. Marine, epigean.
Distribution. South Georgia (Schellenberg 1931), Weddell Sea (Gutt et al. 2000), Southern Ocean.
Remarks. According to Barnard & Karaman (1982: 184) ‘This family has not been properly heralded in the
literature before and is brought to light from a provisional proposal (Schellenberg, 1931: 92)’. Schellenberg (1931)
placed his new genus Pseudamphilochus in the family Amphilochidae, with the suggestion that there could be a
family Pseudamphilochidae. Barnard & Karaman (1982) diagnosed the family but attributed authorship to
Schellenberg (1931). In accordance with Article 13 of the International Code of Zoological Nomenclature (ICZN
1999, fourth edition) they needed to state that they were making a new family for the name to be considered valid.
The original and only illustrations of Pseudamphilochus are of a female and do not include some of the important
morphology. For instance pereopods 3, 5 and 6 are not illustrated. New collections are available but the taxon has

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not yet been redescribed (Gutt et al. 2000).
Parvorder Amphilochidira Boeck, 1871 stat. nov.
Diagnosis. Body laterally compressed or dorsoventrally flattened. Antenna 1 callynophore weakly developed (1-
field) or absent [except Stilipedidae]. Antenna 2 without many robust setae. Gnathopod 1 well developed.
Gnathopod 2 simple, subchelate [except Didymocheliidae], not mitten-shaped. Pereopod 6 and 7 similar in
structure, not immensely elongate. Urosomites 1–3 free, not coalesced [except Sebidae]. Telson entire.
Included superfamilies. Amphilochoidea Boeck, 1871 stat. nov.; Iphimedioidea Boeck, 1871 stat. nov.
Remarks. Amphilochidirans are a well-defined group based on the current set of characters. Three families are
slightly aberrant. The Didymocheliidae have gnathopod 2 chelate, the Stilipedidae maintain a well-developed
callynophore and the Sebidae have coalesced urosomites.
Superfamily Amphilochoidea Boeck, 1871 stat. nov.
Diagnosis. Body laterally compressed. Antenna 1 callynophore weakly-developed or absent. Mouthpart bundle
subquadrate (except Didymocheliidae). Gnathopod 1 simple or subchelate. Gnathopod 2 carpus subequal or shorter
than propodus.
Included families. Amphilochidae Boeck, 1871; Bolttsiidae Barnard & Karaman, 1987; Cressidae Stebbing,
1899; Cyproideidae J.L. Barnard, 1974; Didymocheliidae Bellan-Santini & Ledoyer, 1986; Nihotungidae J.L.
Barnard, 1972a; Pleustidae Buchholz, 1874; Sebidae Walker, 1907; Seborgiidae Holsinger, 1980; Stenothoidae
Boeck, 1871.
Remarks. Didymocheliids are difficult to allocate because they align with the Amphilochoidea in the chelate
second gnathopods and the non-distally acuminate coxae 1–4, but they share the conical mouthpart bundle with the
Iphimedioidea.
Pleustids are also difficult to allocate because they share many iphimedioid character states but they do not
have the conical mouthpart bundle nor the distally acuminate coxae 1–4 of the Iphimedioidea.
Family Amphilochidae Boeck, 1871
Included genera. Afrogitanopsis G. Karaman, 1980b; Amphilochella Schellenberg, 1926a; Amphilochoides G. O .
Sars, 1892; Amphilochopsis Stephensen, 1925b; Amphilochus Spence Bate, 1862; Apolochus Hoover & Bousfield,
2001; Cyclotelson Potts, 1915; Frigora Ren, 1991; Gitana Boeck, 1871; Gitanogeiton Stebbing, 1910; Gitanopsilis
Rauschert, 1994; Gitanopsis G.O. Sars, 1892; Hourstonius Hoover & Bousfield, 2001; Paramphilochoides
Lincoln, 1979; Paramphilochus Ishimaru & Ikehara, 1986; Rostrogitanopsis G. Karaman, 1980b.
Remarks. Frigora Ren, 1991 is here transferred from the Eusiridae to the Amphilochidae.
Family Bolttsiidae Barnard & Karaman, 1987
Included Genera. Bolttsia Griffiths, 1976.
Remarks. Bolttsia has an extraordinarily disjunct distribution from Lake Sebaya, a coastal lake previously
connected to the sea in western South Africa (Griffiths 1976) and estuarine mangroves on the Great Barrier Reef,
Australia (Azman 2009). From the middle Jurassic (170 Ma) to late Jurassic (152 Ma) a continuous coastline ran
from south-western Africa along the ‘northern’ coast of India and the northern coast of Australia. By the early
Cretaceous (~125 Ma) this link was no longer in place. The last time these regions were in contact was in the late
Jurassic, a presumed marker for the minimum age of this genus. A similar situation occurs in the deep water
aristioid genus Izinkala Griffiths, 1977 which is only known from off the south-western African coast and the east
coast of Australia.

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Family Cressidae Stebbing, 1899
Included Genera. Cressa Boeck, 1871; Cressina Stephensen, 1931.
Family Cyproideidae J.L. Barnard, 1974
Included genera. Austropheonoides J.L. Barnard, 1972a; Cyproidea Haswell, 1879b; Gbroidea Lowry & Azman,
2008; Hoplopheonoides Shoemaker, 1956; Hoplopleon K.H. Barnard, 1932; Metacyproidea Ariyama, 2016;
Mokuoloe J.L. Barnard, 1970; Moolapheonoides J.L. Barnard, 1974; Narapheonoides J.L. Barnard, 1972a;
Neocyproidea Hurley, 1955; Paracyproidea Stebbing, 1899; Peltocoxa Catta, 1875; Peltopes K.H. Barnard, 1930;
Pseudopeltocoxa Schiecke, 1977; Sisalia Ortiz & Winfield, 2014; Stegoplax G.O. Sars, 1883; Terepeltopes
Hirayama, 1983; Unguja Griffiths, 1976; Unyapheonoides J.L. Barnard, 1972a; Victorhensenoides Rauschert,
1996.
Family Didymocheliidae Bellan-Santini & Ledoyer, 1986
Included genera. Aidamochelia Thomas & Watling, 2012; Apodidymochelia Thurston, 1997; Didymochelia K.H.
Barnard, 1931.
Remarks. The family exhibits some character states (antenna 1 callynophore weak (1-field) and gnathopod 2
chelate with a short ischium) of the Iphimedioidea.
Family Nihotungidae J.L. Barnard, 1972a
Included genera. Nihotunga J.L. Barnard, 1972a.
Family Pleustidae Buchholz, 1874
Included subfamilies. Atylopsinae Bousfield & Hendrycks, 1994a; Austropleustinae Bousfield & Hendrycks,
1994a; Dactylopleustinae Bousfield & Hendrycks, 1994a; Eosymtinae Bousfield & Hendrycks, 1994a;
Mesopleustinae Bousfield & Hendrycks, 1994a; Neopleustinae Bousfield & Hendrycks, 1994a; Parapleustinae
Bousfield & Hendrycks, 1994a; Pleusirinae Bousfield & Hendrycks, 1994a; Pleustinae Buchholz, 1874;
Pleustoidinae Bousfield & Hendrycks, 1994a; Pleusymtinae Bousfield & Hendrycks, 1994a; Stenopleustinae
Bousfield & Hendrycks, 1994a.
Subfamily Atylopsinae Bousfield & Hendrycks, 1994a
Included genera. Atylopsis Stebbing, 1888; Myzotarsa Cadien & Martin, 1999.
Subfamily Austropleustinae Bousfield & Hendrycks, 1994a
Included genera. Austropleustes K.H. Barnard, 1931; Tepidopleustes Karaman & Barnard, 1979.
Subfamily Dactylopleustinae Bousfield & Hendrycks, 1994a

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Included genera. Dactylopleustes Barnard & Karaman, 1979.
Subfamily Eosymtinae Bousfield & Hendrycks, 1994a
Included genera. Eosymtes Bousfield & Hendrycks, 1994a.
Subfamily Mesopleustinae Bousfield & Hendrycks, 1994a
Included genera. Mesopleustes Stebbing, 1899.
Subfamily Neopleustinae Bousfield & Hendrycks, 1994a
Included genera. Neopleustes Stebbing, 1906; Pleustostenus Gurjanova, 1972; Shoemakeroides Hendrycks &
Bousfield, 2004.
Subfamily Parapleustinae Bousfield & Hendrycks, 1994a
Included genera. Chromopleustes Bousfield & Hendrycks, 1995a; Commensipleustes Bousfield & Hendrycks,
1995a; Gnathopleustes Bousfield & Hendrycks, 1995a; Incisocalliope J.L. Barnard, 1959b; Micropleustes
Bousfield & Hendrycks, 1995a; Parapleustes Buchholz, 1874; Trachypleustes Bousfield & Hendrycks, 1995a.
Subfamily Pleusirinae Bousfield & Hendrycks, 1994a
Included genera. Pleusirus J.L. Barnard, 1969a.
Subfamily Pleustinae Buchholz, 1874
Included genera. Pleustes (Catapleustes) Bousfield & Hendrycks, 1994b; Pleustes (Pleustes) Spence Bate, 1858
(Bousfield & Hendrycks, 1994b); Thorlaksonius Bousfield & Hendrycks, 1994b.
Subfamily Pleustoidinae Bousfield & Hendrycks, 1994a
Included genera. Pleustoides Gurjanova, 1972.
Subfamily Pleusymtinae Bousfield & Hendrycks, 1994a
Included genera. Anomalosymtes Hendrycks & Bousfield, 2004; Budnikopleustes Hendrycks & Bousfield, 2004;
Heteropleustes Hendrycks & Bousfield, 2004; Holopleustes Hendrycks & Bousfield, 2004; Kamptopleustes
Hendrycks & Bousfield, 2004; Pleustomesus Gurjanova, 1972; Pleusymtes J.L. Barnard, 1969b; Rhinopleustes
Hendrycks & Bousfield, 2004.
Subfamily Stenopleustinae Bousfield & Hendrycks, 1994a
Included genera. Arctopleustes Gurjanova, 1972; Domicola Pretus & Abello 1993; Gracilipleustes Hendrycks &

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Bousfield, 2004; Stenopleustes G.O. Sars, 1893.
Remarks. Domicola Pretus & Abello, 1993 is apparently not considered in the monographs of Bousfield &
Hendrycks (1994a, b; 1995a) or Hendrycks & Bousfield (2004). Based on the key to subfamilies in Bousfield &
Hendrycks (1994a) it appears to fit best in the subfamily Stenopleustinae.
Family Sebidae Walker, 1907
Included genera. Seba Spence Bate, 1862.
Family Seborgiidae Holsinger, 1980 stat. nov.
Included genera. Relictoseborgia G. Karaman, 1982; Seborgia Bousfield, 1970.
Family Stenothoidae Boeck, 1871
Included genera. Antatelson J.L. Barnard, 1972a; Aurometopa Barnard & Karaman, 1987; Ausatelson J.L.
Barnard, 1972a; Chuculba J.L. Barnard, 1974; Goratelson J.L. Barnard, 1972a; Hardametopa Barnard & Karaman,
1991; Knysmetopa Barnard & Karaman, 1987; Kyphometopa Krapp-Schickel, 2013; Ligulodactylus Krapp-
Schickel, 2013; Malvinometopa Krapp-Schickel, 2011; Mesometopa Gurjanova, 1938; Mesoproboloides
Gurjanova, 1938; Metopa Boeck, 1871; Metopella G.O. Sars, 1892; Metopelloides Gurjanova, 1938; Metopoides
Della Valle, 1893; Parametopa Chevreux, 1901; Parametopella Gurjanova, 1938; Paraprobolisca Ren, 1991;
Parathaumatelson Gurjanova, 1938; Probolisca Gurjanova, 1938; Proboloides Della Valle, 1893; Prometopa
Schellenberg, 1926a; Prostenothoe Gurjanova, 1938; Prothaumatelson Schellenberg, 1931; Pseudothaumatelson
Schellenberg, 1931; Ptychotelson Krapp-Schickel, 2000; Pycnopyge Krapp-Schickel, 2000; Raukamara Krapp-
Schickel, 2000; Raumahara J.L. Barnard, 1972a; Sandrothoe Krapp-Schickel, 2006a; Scaphodactylus Rauschert &
Andres, 1993; Stenothoe Dana, 1852; Stenothoides Chevreux, 1900; Stenula J.L. Barnard, 1962b; Sudanea Krapp-
Schickel, 2015; Synkope Krapp-Schickel, 1999; Thaumatelson Walker, 1906b; Thaumatelsonella Rauschert &
Andres, 1991; Torometopa Barnard & Karaman, 1987; Verticotelson Krapp-Schickel, 2006b;Victometopa Krapp-
Schickel, 2011; Vonimetopa Barnard & Karaman, 1987; Wallametopa J.L. Barnard, 1974; Yarra Krapp-Schickel,
2000; Zaikometopa Barnard & Karaman, 1987.
Remarks. Historically there have been two subfamilies, Stenothoinae Boeck, 1871 (J.L. Barnard, 1972b) and
Thaumatelsoninae Gurjanova, 1938 (J.L. Barnard, 1972a). However we were not able to confidently place some
genera in either subfamily. Krapp-Schickel (pers comm) thinks that the subfamily concept no longer works for this
group and that several key characters have been independently derived several times.
Superfamily Leucothoidea Dana, 1852 (Bousfield, 1979)
Diagnosis. Maxilliped palps well developed. Gnathopods 1–2 carpochelate. Pereopod 4 coxa larger than that of
pereopod 3.
Included families. Leucothoidae Dana, 1852.
Remarks. The Leucothoidea is the only taxon in the Amphilochida with both gnathopods carpochelate.
Family Leucothoidae Dana, 1852
Included genera. Anamixis Stebbing, 1897; Leucothoe Leach, 1814; Nepanamixis Thomas, 1997; Paraleucothoe
Stebbing, 1899; Paranamixis Schellenberg, 1938.

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Superfamily Iphimedioidea Boeck, 1871 stat. nov.
Diagnosis. Antenna 1 accessory flagellum vestigial or absent. Gnathopod 1 coxa not vestigial. Gnathopod 2 coxa
subequal or smaller than that of coxa 3. Pereopod 4 coxal lobe well-developed or absent.
Included families. Acanthonotozomatidae Stebbing, 1906; Acanthonotozomellidae Coleman & Barnard,
1991a; Amathillopsidae Pirlot, 1934; Dikwidae Coleman & Barnard, 1991a; Epimeriidae Boeck, 1871;
Iphimediidae Boeck, 1871; Lafystiidae Sars, 1893; Laphystiopsidae Stebbing, 1899; Ochlesidae Stebbing, 1910;
Odiidae Coleman & Barnard, 1991a; Sicafodiidae Just, 2004; Stilipedidae Holmes, 1908; Vicmusiidae Just, 1990.
Family Acanthonotozomatidae Stebbing, 1906
Included genera. Acanthonotozoma Boeck, 1876.
Family Acanthonotozomellidae Coleman & Barnard, 1991a
Included genera. Acanthonotozomella Schellenberg, 1926a; Acantonotozomoides Schellenberg, 1931;
Amatiquakius Coleman & Barnard, 1991b.
Family Amathillopsidae Pirlot, 1934
Included subfamilies. Amathillopsinae Pirlot, 1934; Cleonardopsinae Lowry, 2006; Parepimeriinae Lowry, 2006.
Subfamily Amathillopsinae Pirlot, 1934
Included genera. Amathillopsis Heller, 1875.
Subfamily Cleonardopsinae Lowry, 2006
Included genera. Cleonardopsis K.H. Barnard, 1916.
Subfamily Parepimeriinae Lowry, 2006
Included genera. Parepimeria Chevreux, 1911b.
Family Dikwidae Coleman & Barnard, 1991a
Included genera. Dikwa Griffiths, 1974.
Family Epimeriidae Boeck, 1871
Included genera. Actinacanthus Stebbing, 1888; Epimeria Costa, 1851; Metepimeria Schellenberg, 1931;
Paramphithoe Bruzelius, 1859; Uschakoviella Gurjanova, 1955.
Family Iphimediidae Boeck, 1871

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Included genera. Anchiphimedia K.H. Barnard, 1930; Anisoiphimedia G. Karaman, 1980c; Coboldus Krapp-
Schickel, 1974; Echiniphimedia K.H. Barnard, 1930; Gnathiphimedia K.H. Barnard, 1930; Iphimedia Rathke,
1843; Iphimediella Chevreux, 1911b; Labriphimedia K.H. Barnard, 1931; Maxilliphimedia K.H. Barnard, 1930;
Nodotergum Bellan-Santini, 1972; Paranchiphimedia Ruffo, 1949; Parapanoploea Nicholls, 1938; Pariphimedia
Chevreux 1906a; Pseudiphimediella Schellenberg, 1931; Stegopanoploea G. Karaman, 1980c.
Family Lafystiidae Sars, 1893
Included genera. Lafystius Krøyer, 1842; Paralafystius Bousfield, 1987; Protolafystius Bousfield, 1987.
Family Laphystiopsidae Stebbing, 1899
Included genera. Laphystiopsis G.O. Sars, 1893; Prolaphystiopsis Schellenberg, 1931; Prolaphystius K.H.
Barnard, 1930.
Family Ochlesidae Stebbing, 1910
Included genera. Ochlesis Stebbing, 1910; Curidia Thomas, 1983; Meraldia Barnard & Karaman, 1987;
Ochlesodius Ledoyer, 1982.
Family Odiidae Coleman & Barnard, 1991a
Included genera. Antarctodius Berge, Vader & Coleman, 1999; Cryptodius Moore, 1992a; Gordonodius Ariyama,
2011; Imbrexodius Moore, 1992a; Odius Lilljeborg, 1865a; Postodius Hirayama, 1983.
Family Sicafodiidae Just, 2004
Included genera. Sicafodia Just, 2004.
Family Stilipedidae Holmes, 1908
Included subfamilies. Astryinae Pirlot, 1934; Alexandrellinae Holman & Watling, 1983; Stilipedinae Holmes,
1908.
Subfamily Astryinae Pirlot, 1934
Included genera. Astyra Boeck, 1871; Eclysis K.H. Barnard, 1932.
Subfamily Alexandrellinae Holman & Watling, 1983
Included genera. Alexandrella Chevreux, 1911b; Astyroides Birstein & Vinogradov, 1960; Bathypanoplea
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Subfamily Stilipedinae Holmes, 1908
Included genera. Stilipes Holmes, 1908.
Family Vicmusiidae Just, 1990
Included genera. Acanthonotozomopsis Watling & Holman, 1980.
Infraorder Lysianassida Dana, 1849 stat. nov.
Diagnosis. Antenna 1 primary flagellum 5 or more articulate (except Izinkalidae and Vemanidae). Antenna 1 and 2
without eusiroid calceolus. Gnathopod 2 carpus long (except Melphidippidae, Haustoriidae), longer than propodus
(except Priscillinidae and Vemanidae). Coxae 1–4 not acuminate [except Haustoriidae, Ipanemidae, Priscillinidae,
Stegocephalidae]. Pereopod 7 not immensely elongate. Pleonites without dorsal spines (except Priscillinidae).
Included parvorders. Synopiidira Dana, 1852 stat. nov.; Haustoriidira Stebbing, 1906 stat. nov.;
Lysianassidira Dana, 1849 stat. nov..
Remarks. The Lysianassida are defined by the above diagnostic characters, but a number of small highly
derived taxa (Haustoriidae, Izinkalidae, Melphidippidae, Priscillinidae and Vemanidae) are anomalous. The large
family Haustoriidae has a short carpus on gnathopod 2. In this respect it does not conform to the diagnosis.
Parvorder Synopiidira Dana, 1852 stat. nov.
Diagnosis. Antennae 1–2 calceoli absent. Antenna 2 peduncle with brush setae; flagellum in adult male greatly
elongate. Pereopods 5–7 with few or no robust setae. Telson cleft (except in some Synopia).
Included superfamilies. Dexaminoidea Leach, 1814; Synopioidea Dana, 1852 (Bousfield, 1979).
Remarks. Bousfield & Kendall (1994) indicate that male atylids have elongate second antennae, but they
illustrate some species with a short second antenna.
Superfamily Dexaminoidea Leach, 1814 (Bousfield, 1979)
Diagnosis. Antenna 2 brush setae present in adult males, Mandible lacinia mobilis present on both sides. Pereopods
5–7 similar in length or 5 shorter and 6–7 similar length, with few or no robust or slender setae. Pereopod 5 basis
linear.
Included families. Atylidae Lilljeborg, 1865a; Dexaminidae Leach, 1814; Melphidippidae Stebbing, 1899;
Pardaliscidae Boeck, 1871.
Family Atylidae Lilljeborg, 1865a
Included subfamilies. Anatylinae Bulycheva, 1955; Atylinae Lilljeborg, 1865a; Lepechinellinae Schellenberg,
1926a; Nototropiinae Bousfield & Kendall, 1994.
Subfamily Anatylinae Bulycheva, 1955
Included genera. Anatylus Bulycheva, 1955; Kamehatylus J.L. Barnard, 1970.

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Subfamily Atylinae Lilljeborg, 1865a
Included genera. Atylus Leach, 1815.
Subfamily Lepechinellinae Schellenberg, 1926a (Bousfield & Kendall, 1994)
Included genera. Lepechinella Stebbing, 1908; Lepechinelloides Thurston, 1980; Lepechinellopsis Ledoyer, 1982;
Lepesubchela Johansen & Vader, 2015; Paralepechinella Pirlot, 1933b.
Subfamily Nototropiinae Costa 1853 (Bousfield & Kendall, 1994)
Included genera. Aberratylus Bousfield & Kendall, 1994; Nototropis Costa, 1853.
Family Dexaminidae Leach, 1814
Included subfamilies. Dexamininae Leach, 1814; Dexaminoculinae Bousfield & Kendall, 1994; Polycheriinae
Bousfield & Kendall, 1994; Prophliantinae Nicholls, 1939.
Subfamily Dexamininae Leach, 1814
Included genera. Delkarlye J.L. Barnard, 1972a; Dexamine Leach, 1814; Dexaminella Schellenberg, 1928b;
Paradexamine Stebbing, 1899; Sebadexius Ledoyer, 1984; Syndexamine Chilton, 1914.
Subfamily Dexaminoculinae Bousfield & Kendall, 1994
Included genera. Dexaminoculus Lowry, 1981.
Subfamily Polycheriinae Bousfield & Kendall, 1994
Included genera. Polycheria Haswell, 1879b; Tritaeta Boeck, 1876.
Subfamily Prophliantinae Nicholls, 1939
Included genera. Guernea Chevreux, 1887; Haustoriopsis Schellenberg, 1938; Prophlias Nicholls, 1939.
Family Melphidippidae Stebbing, 1899
Included genera. Melphidippa Boeck, 1871; Melphidipella G.O. Sars, 1894; Melphisana J.L. Barnard, 1962a;
Melphisubchela Andres, 1981b.
Family Pardaliscidae Boeck, 1871
Included genera. Andeepia Biswas, Coleman & Hendrycks, 2009; Antronicippe Stock & Iliffe, 1990; Arculfia J.L.
Barnard, 1961; Caleidoscopsis G. Karaman, 1974; Epereopus Mills, 1967; Halice Boeck, 1871; Halicella

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Schellenberg, 1926a; Halicoides Walker, 1896; Macroarthrus Hendrycks & Conlan, 2003; Necochea Barnard,
1962c; Nicippe Bruzelius, 1859; Octomana Hendrycks & Conlan, 2003; Parahalice Birstein & Vinogradov, 1962;
Pardalisca Krøyer, 1842; Pardaliscella G.O. Sars, 1893; Pardaliscoides Stebbing, 1888; Pardaliscopsis Chevreux,
1911a; Parpano J.L. Barnard, 1964a; Princaxelia Dahl, 1959; Rhynohalicella G. Karaman, 1974; Spelaeonicippe
Stock & Vermeulen, 1982; To s i l us J.L. Barnard, 1966.
Superfamily Synopioidea Dana, 1852 (Bousfield, 1979)
Diagnosis. Gnathopod 2 simple (also some pardaliscids). Pereopod 4 coxa with well-developed posteroventral
lobe. Pereopods 5–7 with few or no robust or slender setae.
Included families. Ampeliscidae Krøyer, 1842; Argissidae Walker, 1904; Synopiidae Dana, 1852.
Family Ampeliscidae Krøyer, 1842
Included genera. Ampelisca Krøyer, 1842; Byblis Boeck, 1871; Byblisoides K.H. Barnard, 1931; Haploops
Liljeborg, 1856.
Family Argissidae Walker, 1904
Included genera. Argissa Boeck, 1871.
Remarks. Argissa is in need of revision. The distribution, depth range and morphological variation attributed
to A. hamatipes are implausible when attributed to a single species.
Argissa hamatipes (Norman, 1869) was originally described from shallow water in St Magnus Bay, Shetland
Islands, Scotland. Argissa stebbingi Bonnier, 1896 was described from 940 m depth in the Bay of Biscay, France.
These two species are currently in synonymy and many more records have been added so that based on current
taxonomy the climatic distribution of Argissa hamatipes extends from Iceland (Sub-Arctic) to Sri Lanka (high
tropics), the depth distribution extends from the shallow subtidal (14 m depth) to the lower bathyal (nearly 1000 m
depth) and the geographic distribution is one of the widest of any non-hyperiidean amphipod species, from
California to the Sea of Japan in the Pacific Ocean, from southern India and Sri Lanka to Madagascar and southern
Africa in the Indian Ocean, and in the Atlantic Ocean from West Africa and the Azores, to the Mediterranean Sea
and along the European coast to Norway, and across the northern North Atlantic to Iceland and Greenland and from
the Gulf of St Lawrence to New England in North America.
A number of records have been published without illustrations (Walker, 1904 (Sri Lanka); Stephensen, 1940
(Iceland); Schellenberg, 1942; Reid, 1951 (West Africa)). More were published with illustrations: Chevreux &
Fage, 1925 (Bay of Biscay, the Azores and West Africa); Shoemaker, 1930 (Gulf of St Lawrence); Gurjanova,
1951; Nagata, 1965 (Sea of Japan); J.L. Barnard, 1967 (southern California); Bousfield, 1973 (New England);
Griffiths, 1976 (Cape Province, South Africa); Lincoln, 1979 (British Isles) and Ledoyer, 1982 (Madagascar)).
These illustrations often indicate species level morphological differences. For instance, based on morphology, the
well-illustrated material of Rabindranath (1972) from Trivandrum, Kerala, India should be considered as a new
species.
Most authors have accepted the synonymy without question. Ruffo (1982) maintained two species, but the
problem is that the taxon is so confused that trying to maintain the two names does not help. There has to be a
revision based on all available material.
Family Synopiidae Dana, 1852
Included genera. Austrosyrroe K.H. Barnard, 1925; Bruzelia Boeck, 1871; Bruzeliopsis Chevreux, 1911a;
Cardenio Stebbing, 1888; Garosyrrhoe J.L. Barnard, 1964a; Ileraustroe J.L. Barnard, 1969b; Jeddo J.L. Barnard,
1962c; Latacunga J.L. Barnard, 1972c; Metatiron Rabindranath, 1972; Priscosyrrhoe J.L. Barnard, 1972c;

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Pseudotiron Chevreux, 1895; Stephobruzelia J.L. Barnard, 1969b; Synopia Dana, 1852; Syrrhoe Goës, 1866;
Syrrhoites G.O. Sars, 1893; Telsosynopia G. Karaman, 1986; Tiron Lilljeborg, 1865b.
Remarks. J.L. Barnard (1969b: 460) described the new genus Ileraustroe and then later described the genus as
new for a second time (J.L. Barnard 1972c: 34).
Parvorder Haustoriidira Stebbing, 1906 stat. nov.
Diagnosis. Head rostrum present. Antenna 1 and 2 peduncles and pereopods 5–7 with many robust and slender
setae (except Pontoporeiidae and Priscillinidae). Pereopod 3 merus longer than propodus. Epimeron 2 with
plumose setae (except Pontoporeiidae, Priscillinidae and Platyischnopidae).
Included superfamilies. Haustorioidea Stebbing, 1906.
Remarks. The basal families Pontoporeiidae and Priscillinidae do not have the many robust and slender setae
on the peduncles of antenna 1 and 2 and on pereopods 5–7, nor the plumose setae on epimeron 2 that are typical of
most Haustorioidea. However, both families share with phoxocephalids, the characteristically dissimilar pereopods
6 and 7. Pontoporeiids share embedded setae on the rami of uropods 1 and 2 with the Phoxocephalidae. Also the
Phoxocephalidae, Pontoporeiidae and Priscillinidae are the only families in the parvorder with reniform eyes.
Superfamily Haustorioidea Stebbing, 1906 (Barnard & Drummond, 1982b)
Diagnosis. As for parvorder.
Included families. Cheidae Thurston, 1982; Condukiidae Barnard & Drummond, 1982b; Haustoriidae
Stebbing, 1906; Ipanemidae Barnard & Thomas, 1988b; Otagiidae Hughes & Lörz, 2013; Phoxocephalidae G.O.
Sars, 1891; Phoxocephalopsidae Barnard & Drummond, 1982b; Platyischnopidae Barnard & Drummond, 1979;
Pontoporeiidae Dana, 1853; Priscillinidae d’Udekem d’Acoz, 2007; Sinurothoidae Ren, 1999; Urohaustoriidae
Barnard & Drummond, 1982b; Urothoidae Bousfield, 1979; Zobrachoidae Barnard & Drummond, 1982b.
Family Cheidae Thurston, 1982
Included genera. Cheus Thurston, 1982; Microcheus Souza-Fihlo, 2011; Ruffosius Souza-Fihlo, 2011.
Family Condukiidae Barnard & Drummond, 1982b
Included genera. Condukius Barnard & Drummond, 1982b.
Family Haustoriidae Stebbing, 1906
Included genera. Acanthohaustorius Bousfield, 1965; Eohaustorius J.L. Barnard, 1957b; Haustorius Müller,
1775; Lepidactylus Say, 1818a; Neohaustorius Bousfield, 1965; Parahaustorius Bousfield, 1965; Protohaustorius
Bousfield, 1965; Pseudohaustorius Bousfield, 1965.
Family Ipanemidae Barnard & Thomas, 1988b
Included genera. Ipanema Barnard & Thomas, 1988b.
Family Otagiidae Hughes & Lörz, 2013

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Included genera. Otagia Barnard & Karaman, 1991.
Family Phoxocephalidae G.O. Sars, 1891
Included subfamilies (based on De Broyer et al. 2007).
Remarks. Barnard & Drummond (1998) established nine subfamilies for the world phoxocephalid species
(Birubiinae Barnard & Drummond, 1978; Brolginae Barnard & Drummond, 1978; Harpiniinae Barnard &
Drummond, 1978; Joubinellinae Barnard & Drummond, 1978; Leongathinae Barnard & Drummond, 1978;
Palabriaphoxinae Gurjanova, 1977; Phoxocephalinae G.O. Sars, 1891; Pontharpiniinae Barnard & Drummond,
1978; Tipimeginae Barnard & Drummond, 1978). Barnard & Karaman (1991) partially revised this system but
were unable to unequivocally place a number of genera into specific subfamilies.
Jarrett & Bousfield (1994a) established the Metharpiniinae for several new genera and incorporated some
difficult genera. De Broyer et al. (2007) recognised only two subfamilies Harpiniinae and Phoxocephalinae in the
catalogue of Antarctic Amphipoda. Alonso de Pina et al. (2008) was critical of the scheme and excluded
subfamilies when cataloguing the Antarctic taxa.
Because of the confusion in the classification of this family, we follow the classification of De Broyer et al.
2007.
Subfamily Harpiniinae Barnard & Drummond, 1978
Included genera. Basuto Barnard & Drummond, 1978; Cocoharpinia G. Karaman, 1980a; Coxophoxus J.L.
Barnard, 1966; Feriharpinia Barnard & Karaman, 1982; Harpinia Boeck, 1876; Harpiniopsis Stephensen, 1925b;
Heterophoxus Shoemaker, 1925; Proharpinia Schellenberg, 1931; Pseudharpinia Schellenberg, 1931;
Torridoharpinia Barnard & Karaman, 1982.
Subfamily Phoxocephalinae G.O. Sars, 1891 (Barnard & Drummond, 1978)
Included genera. Baliphoxus Ortiz & Lalana, 1999; Bathybirubius Senna, 2010; Beringiaphoxus Jarrett &
Bousfield, 1994a; Birubius Barnard & Drummond, 1978; Booranus Barnard & Drummond, 1978; Brolgus Barnard
& Drummond, 1978; ephalophoxoides Gurjanova, 1977; Cephalophoxus Gurjanova, 1977; Cunmurra Barnard &
Drummond, 1978; Diogodias Barnard & Drummond, 1978; Elpeddo Barnard & Drummond, 1978; Eobrolgus J.L.
Barnard, 1979; Eusyrophoxus Gurjanova, 1977; Eyakia J.L. Barnard, 1979; Foxiphalus J.L. Barnard, 1979;
Fuegiphoxus Barnard & Barnard, 1980; Ganba Barnard & Drummond, 1978; Grandiphoxus J.L. Barnard, 1979;
Griffithsius Jarrett & Bousfield, 1994b; Hopiphoxus Barnard & Drummond, 1978; Indophoxus Dang & Le, 2005;
Japara Barnard & Drummond, 1978; Jerildaria Barnard & Drummond, 1978; Joubinella Chevreux, 1908a;
Kondoleus Barnard & Drummond, 1978; Kotla Barnard & Drummond, 1978; Kulgaphoxus Barnard & Drummond,
1978; Kuritus Barnard & Drummond, 1978; Leongathus Barnard & Drummond, 1978; Leptophoxoides J.L.
Barnard, 1962c; Leptophoxus G.O. Sars, 1891; Limnoporeia Fearn-Wannan, 1968; Linca Alonso de Pina, 1993;
Majoxiphalus Jarrett & Bousfield, 1994a; Mandibulophoxus J.L. Barnard, 1957a; Matong Barnard & Drummond,
1978; Mesophoxus Gurjanova, 1977; Metaphoxoides Ledoyer, 1968; Metaphoxus Bonnier, 1896; Metharpinia
Schellenberg, 1931; Microphoxus J.L. Barnard, 1960; Palabriaphoxus Gurjanova, 1977; Parafoxiphalus Alonso de
Pina, 2001; Parajoubinella Gurjanova, 1977; Paramesophoxus Gurjanova, 1977; Parametaphoxus Gurjanova,
1977; Paraphoxus G.O. Sars 1891; Parharpinia Stebbing, 1899; Phoxocephalus Stebbing, 1888; Phoxorgia
Barnard & Barnard, 1980; Pontharpinia Stebbing, 1897; Protophoxus K.H. Barnard, 1930; Pseudfoxiphalus
Andres, 1991; Rhepoxynius J.L. Barnard, 1979; Rikkarus Barnard & Drummond, 1978; Ringaringa Barnard &
Karaman, 1991; Synphoxus Gurjanova, 1980a; Tickalerus Barnard & Drummond, 1978; Tipimegus Barnard &
Drummond, 1978; Trichophoxus K.H. Barnard, 1930; Uldanamia Barnard & Drummond, 1978; Urophoxus
Gurjanova, 1977 (=Pontharpinia according to Barnard & Karman, 1991); Va sc o Barnard & Drummond, 1978;

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Vietophoxus Dang & Le, 2005; Waipirophoxus Gurjanova, 1980b; Waitangi Fincham, 1977; Wildus Barnard &
Drummond, 1978; Yammacoona Barnard & Drummond, 1978; Ya n Barnard & Drummond, 1978.
Family Phoxocephalopsidae Barnard & Drummond, 1982b
Included genera. Eophoxocephalopsis Thurston, 1989a; Phoxocephalopsis Schellenberg, 1931; Pseudurothoe
Ledoyer, 1986; Puelche Barnard & Clark, 1982a; Urothopsis Ledoyer, 1968.
Remarks. Sittrop et al. (2014) removed Pseudurothoe and Urothopsis from the Urothoidae and placed them in
the Phoxocephalopsidae.
Family Platyischnopidae Barnard & Drummond, 1979
Included genera. Eudevenopus Thomas & Barnard, 1983; Indischnopus Barnard & Drummond, 1979; Platyisao
Chiesa & Alonso, 2014; Platyischnopus Stebbing, 1888; Skaptopus Thomas & Barnard, 1983; Tiburonella Thomas
& Barnard, 1983; Tittakunara Barnard & Drummond, 1979; Tomituka Barnard & Drummond, 1979; Yu r r o k u s
Barnard & Drummond, 1979.
Family Pontoporeiidae Dana, 1853
Included genera. Diporeia Bousfield, 1979; Monoporeia Bousfield, 1979; Pontoporeia Krøyer, 1842.
Remarks. The genus Zaramilla Stebbing, 1888, historically placed in the Pontoporeiidae, has recently been
moved to the new monotypic family Zaramillidae Lowry & Myers, 2016 in the Senticaudata.
Family Priscillinidae d’Udekem d’Acoz, 2007
Included genera. Priscillina Stebbing, l888.
Remarks. In this study the Priscillinidae sits isolated at the base of the haustoriidiran clade next to the
Pontoporeiidae. There is no support to maintain the superfamily Pontoporeiidoidea for the Priscillinidae and
Pontoporeiidae. d’Udekem d’Acoz (2007) maintained that the Pontoporeiidae and the Priscillinidae were most
similar to the Gammaroidea, but there is little support for this view. Priscillinids, like pontoporeiids have important
haustoriidiran features, such as setose antennae and pereopods and pereopods 6 and 7 different in structure and
uropods 1 and 2 without apical robust setae.
Family Sinurothoidae Ren, 1999
Included genera. Sinurothoe Ren, 1999.
Family Urohaustoriidae Barnard & Drummond, 1982b
Included genera. Dirimus Barnard & Drummond, 1982b; Gheegerus Barnard & Drummond, 1982b; Huarpe
Barnard & Clark, 1982b; Narunius Barnard & Drummond, 1982b; Nepelle Barnard & Drummond, 1991; Tottungus
Barnard & Drummond, 1982b; Tuldarus Barnard & Drummond, 1982b; Urohaustorius Sheard, 1936; Warragaia
Berents, 1985.
Family Urothoidae Bousfield, 1979

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Included genera. Carangolia J.L. Barnard, 1961; Carangolioides Sittrop, Serejo, Souza-Filho & Senna, 2014;
Coronaurothoe Sittrop, Serejo, Souza-Filho & Senna, 2014; Cunicus Griffiths, 1974; Urothoe Dana, 1852;
Urothoides Stebbing, 1891.
Family Zobrachoidae Barnard & Drummond, 1982b
Included genera. Bumeralius Barnard & Drummond, 1982b; Chono Clark & Barnard, 1987; Prantinus Barnard &
Drummond, 1982b; Tono c o t e Clark & Barnard, 1986; Zobracho J.L. Barnard, 1961.
Parvorder Lysianassidira Dana, 1849 stat. nov.
Diagnosis. Body laterally compressed with large coxae. Antenna 1 callynophore well-developed. Mandible incisor
smooth (except Alicelloidea); lacinia mobilis on left side only or vestigial on right [except Valettiopsidae].
Gnathopod 1 ischium long (except Parargissidae, Vemanidae and Valettidae).
Included superfamilies. Alicelloidea Lowry & De Broyer, 2008; Stegocephaloidea Dana, 1852 (Bousfield,
1979); Lysianassoidea Dana, 1849 (Bousfield, 1979); Aristioidea Lowry & Stoddart, 1997.
Remarks. Members of the distinctive Alicelloidea do not fit well into the diagnostic framework of the
Lysianassidira, for example lysianassoids and aristioids have the lacinia mobilis present on the left side only. In the
Alicelloidea by contrast the Alicellidae and the Valettiopsidae have the lacinia mobilis on both sides. Other
alicelloids such as Vemanidae and Parargissidae are too poorly described to determine.
Superfamily Alicelloidea Lowry & De Broyer, 2008 stat. nov.
Diagnosis. Mandible molar and palp present. Maxilla 1 basal endite strongly setose along medial margin.
Gnathopod 2 not mitten-shaped.
Included families. Alicellidae Lowry & De Broyer, 2008; Parargissidae fam. nov.; Podoprionidae Lowry &
Stoddart, 1996; Valettidae Stebbing, 1888; Valettiopsidae Lowry & De Broyer, 2008; Vemanidae fam. nov.
Family Alicellidae Lowry & De Broyer, 2008
Included genera. Alicella Chevreux, 1899; Apotectonia Barnard & Ingram, 1990; Diatectonia Barnard & Ingram,
1990; Paralicella Chevreux, 1908a; Tectovalopsis Barnard & Ingram, 1990; Transtectonia Barnard & Ingram,
1990.
Family Parargissidae fam. nov.
Type genus. Parargissa Chevreux, 1908c.
Diagnosis (based on world family database). Antenna 1 article 2 much longer than article 3, forming a dorsal
flange. Maxilla 1 palps asymmetrical. Gnathopod 1 coxa smaller than 2. Gnathopods 1–2 merus linear, much loner
that propodus. Pereopod 3 prehensile, merus much longer than propodus. Uropods 1–2 without apical robust setae.
Telson moderately cleft.
Habitat. Marine, epigean, deep sea.
Included genera. Parargissa Chevreux, 1908c.
Distribution. Off Durban, south-western Indian Ocean, 4360 m; off Acapulco-Panama, eastern Central Pacific
Ocean, 3570 m; off the Azores, North Atlantic Ocean.
Remarks. Parargissa was formally assigned to the Hyperiopsidae. It forms a new monotypic family here

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which appears to be the sister group of vemanids. Both have sparse or no slender setae on the peduncle of antenna,
flagellum longer than peduncle; maxilla 1 strongly setose on medial margin; gnathopod 1 smaller than gnathopod
2; pereopod 5 basis subrectangular; pereopod 7 different in structure from pereopod 6 and telson, long, laminar and
cleft.
Family Podoprionidae Lowry & Stoddart, 1996
Included genera. Podoprion Chevreux, 1891.
Remarks. The Podoprionidae, formerly assigned to the Lysianassoidea is reassigned to the Alicelloidea.
Family Valettidae Stebbing, 1888
Included genera. Valettia Stebbing, 1888.
Family Valettiopsidae Lowry & De Broyer, 2008
Included genera. Valettietta Lincoln & Thurston, 1983; Valettiopsis Holmes, 1908.
Vemanidae fam. nov.
Type genus. Vemana J.L. Barnard, 1964a.
Diagnosis (based on world family database). Antenna 1 shorter than antenna 2; callynophore well-developed;
accessory flagellum short or minute. Antennae 1–2 calceoli absent. Mandible incisor dentate; molar large,
triturative. Maxilla 1 basal endite setose along medial margin. Maxilla 2 basal endite with oblique setal row.
Maxilliped palps well-developed. Gnathopods 1–2 subchelate. Pereopods 1–4 coxae not ventrally acute. Pereopod
4 coxa subequal or smaller than pereopod 3 coxa. Uropods 1–2 without apical robust setae. Telson laminar,
moderately cleft.
Included genera. Vemana J.L. Barnard, 1964a.
Habitat. Marine, deep sea.
Distribution. Madagascar, Western Indian Ocean, western North America, north-eastern Pacific Ocean,
Caribbean basin, tropical western Atlantic.
Remarks. Vemana was thought to be most similar to a hyperiopsid or a vitjazianid by J.L. Barnard (1964a). He
placed it in the Vitjazianidae based on the morphology of maxilla 1. Ledoyer (1986) contrasted Ve m a n a and
Vitjaziana and questioned whether they should be in the same family. He showed similarities between Vemana and
the lysianassids. In our analyses the Vemanidae aligns with the Parargissidae in the Lysianassidira.
Superfamily Stegocephaloidea Dana, 1852 (Bousfield, 1979)
Diagnosis. Body laterally compressed with large coxae. Mandible molar and palp absent. Gnathopod 2 not mitten-
shaped; ischium long.
Included families. Stegocephalidae Dana, 1852.
Family Stegocephalidae Dana, 1852
Included subfamilies. Andaniexinae Berge & Vader, 2001; Andaniopsinae Berge & Vader, 2001;
Bathystegocephalinae Berge & Vader, 2001; Parandaniinae Berge & Vader, 2001; Stegocephalinae Dana, 1852.

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Subfamily Andaniexinae Berge & Vader, 2001
Included genera. Andaniexis Stebbing, 1906; Andaniotes Stebbing, 1897; Glorandaniotes Ledoyer, 1986;
Mediterexis Berge & Vader, 2001; Metandania Stephensen, 1925b; Parandaniexis Schellenberg, 1929;
Stegosoladidus Barnard & Karaman, 1987.
Subfamily Andaniopsinae Berge & Vader, 2001
Included genera. Andaniopsis G.O. Sars, 1891; Sinoandaniopsis Ren, 2012; Steleuthera J.L. Barnard, 1964a.
Subfamily Bathystegocephalinae Berge & Vader, 2001
Included genera. Bathystegocephalus Schellenberg, 1926b.
Subfamily Parandaniinae Berge & Vader, 2001
Included genera. Parandania Stebbing, 1899.
Subfamily Stegocephalinae Dana, 1852 (Berge & Vader, 2001)
Included genera. Alania Berge & Vader, 2001; Austrocephaloides Berge & Vader, 2001; Austrophippsia Berge &
Vader, 2001; Bouscephalus Berge & Vader, 2001; Gordania Berge & Vader, 2001; Phippsia Stebbing, 1906 ;
Pseudo Berge & Vader, 2001 (=Schellenbergia Berge & Vader, 2001); Stegocephalexia Moore, 1992a;
Stegocephalina Stephensen, 1925b (=Stegophippsiella Bellan-Santini & Ledoyer, 1974); Stegocephaloides G.O .
Sars, 1891; Stegocephalus Krøyer, 1842 (=Phippsiella Schellenberg, 1925) (=Stegocephalopsis Schellenberg,
1925); Stegomorphia Berge & Vader, 2001; Stegonomadia Berge & Vader, 2001; Tetradeion Stebbing, 1899.
Superfamily Lysianassoidea Dana, 1849 (Bousfield, 1979)
Diagnosis. Antennae 1–2 type 3 lysianassoid calceolus present (except Acidostomatidae and Conicostomatidae).
Gnathopod 2 mitten-shaped (except Amaryllididae). Pereopod 5 propodus posterodistal spur absent.
Included families. Adeliellidae fam. nov.; Amaryllididae Lowry & Stoddart, 2002; Cebocaridae Lowry &
Stoddart, 2011a; Cyclocaridae Lowry & Stoddart, 2011a; Cyphocarididae Lowry & Stoddart, 1997; Eurytheneidae
Stoddart & Lowry, 2004; Hirondelleidae Lowry & Stoddart, 2010a; Lysianassidae Dana, 1849; Opisidae Lowry &
Stoddart, 1995b; Scopelocheiridae Lowry & Stoddart, 1997; Tryphosidae Lowry & Stoddart, 1997 stat. nov.;
Uristidae Hurley, 1963.
Family Adeliellidae fam. nov.
Type genus. Adeliella Nicholls, 1938.
Diagnosis (based on world family database). Antenna 1 subequal in length to antenna 2. Maxilla 1 ischial
endite with 6 setal-teeth. Gnathopod 1 subchelate. Gnathopod 2 minutely parachelate (mitten-shaped). Pereopod 4
coxa with small posteroventral lobe or without posteroventral lobe. Telson moderately or weakly cleft.
Habitat. Marine, epigean.
Included genera. Adeliella Nicholls, 1938.
Remarks. It is difficult to see similarities between Adeliellidae and other lysianassoid families. In our tree it

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sits in the eurytheneid clade between the scavenging hirondelleids and the predatory cyphocaridids.
Distribution. Southern Ocean. Western Indian Ocean. Eastern tropical Atlantic Ocean.
Family Amaryllididae Lowry & Stoddart, 2002
Included subfamilies. Amaryllidinae Lowry & Stoddart, 2002; Vijayiinae Lowry & Stoddart, 2002.
Subfamily Amaryllidinae Lowry & Stoddart, 2002
Included genera. Amaryllis Haswell, 1879a; Bamarooka Lowry & Stoddart, 2002; Erikus Lowry & Stoddart,
1987; Wonga Lowry & Stoddart, 2002.
Subfamily Vijayiinae Lowry & Stoddart, 2002
Included genera. Bathyamaryllis Pirlot, 1933b; Devo Lowry & Stoddart, 2002; Pseudamaryllis Andres, 1981a;
Vija ya Walker, 1904.
Family Cebocaridae Lowry & Stoddart, 2011a
Included genera. Cebocaris J.L. Barnard, 1964a; Crybelocephalus Tattersall, 1906; Crybelocyphocaris
Shoemaker, 1945; Cyphocarioides Birstein & Vinogradov, 1970; Mesocyclocaris Birstein & Vinogradov, 1964;
Mesocyphocaris Birstein & Vinogradov, 1960; Metacyclocaris Birstein & Vinogradov, 1955; Metacyphocaris
Tattersall, 1906; Paracyphocaris Chevreux, 1905a.
Family Cyclocaridae Lowry & Stoddart, 2011a
Included genera. Cyclocaris Stebbing, 1888
Family Cyphocarididae Lowry & Stoddart, 1997
Included genera. Cyphocaris Boeck, 1871; Procyphocaris J.L. Barnard, 1961.
Family Eurytheneidae Stoddart & Lowry, 2004
Included genera. Eurythenes S.I. Smith, 1882.
Family Hirondelleidae Lowry & Stoddart, 2010a
Included genera. Hirondellea Chevreux, 1889.
Family Lysianassidae Dana, 1849
Included subfamilies. Lysianassinae Dana, 1849; Waldeckiinae Lowry & Kilgallen, 2014a.

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Subfamily Lysianassinae Dana, 1849
Included genera. Acosta Özdikmen, 2012; Aruga Holmes, 1908; Arugella Pirlot, 1936; Azotostoma J.L. Barnard,
1965; Bonassa Barnard & Karaman, 1991; Concarnes Barnard & Karaman, 1991; Dartenassa Barnard &
Karaman, 1991; Dissiminassa Barnard & Karaman, 1991; Kakanui Lowry & Stoddart, 1983; Lysianassa H. Milne
Edwards, 1830; Lysianassina Costa, 1867; Lysianopsis Holmes, 1905; Macronassa Barnard & Karaman, 1991;
Nannonyx G.O. Sars, 1891; Parawaldeckia Stebbing, 1910; Phoxostoma K.H. Barnard, 1925; Pronannonyx
Schellenberg, 1953; Pseudambasia Stephensen, 1927; Riwo Lowry & Stoddart, 1995b; Shoemakerella Pirlot, 1936;
Socarnella Walker, 1904; Socarnes Boeck, 1871; Socarnoides Stebbing, 1888; Socarnopsis Chevreux, 1911c;
Tant ena Ortiz, Lalana & Varela, 2007; Thaumodon Lowry & Stoddart, 1995a.
Subfamily Waldeckiinae Lowry & Kilgallen, 2014a
Included genera. Waldeckia Chevreux, 1906a.
Family Opisidae Lowry & Stoddart, 1995b
Included genera. Normanion Bonnier, 1893; Opisa Boeck, 1876; Podoprionella G.O. Sars, 1895; Podoprionides
Walker, 1906a.
Family Scopelocheiridae Lowry & Stoddart, 1997
Included subfamilies. Scopelocheirinae Lowry & Stoddart, 1997 (Kilgallen & Lowry, 2015a); Paracallisominae
Kilgallen & Lowry, 2015a.
Subfamily Paracallisominae Kilgallen & Lowry, 2015a
Included genera. Anisocallisoma Hendrycks & Conlan, 2003; Austrocallisoma Kilgallen & Lowry, 2015a;
Bathycallisoma Dahl, 1959; Eucallisoma J.L. Barnard, 1961; Haptocallisoma Horton & Thurston, 2015;
Paracallisoma Chevreux, 1903; Pseudocallisoma Horton & Thurston, 2015; Scopelocheiropsis Schellenberg,
1926a; Tayabasa Kilgallen & Lowry, 2015a.
Subfamily Scopelocheirinae Lowry & Stoddart, 1997 (Kilgallen & Lowry, 2015a)
Included genera. Aroui Chevreux, 1911c; Paracallisomopsis Gurjanova, 1962; Scopelocheirus Spence Bate,
1857.
Family Tryphosidae Lowry & Stoddart, 1997 stat. nov.
Included genera. Allogaussia Schellenberg, 1926a; Bruunosa Barnard & Karaman, 1987; Cedrosella Barnard &
Karaman, 1987; Cheirimedon Stebbing, 1888; Coximedon Barnard & Karaman, 1991; Elimedon J.L. Barnard,
1962c; Falklandia De Broyer, 1985; Glorieusella Kilgallen & Lowry, 2014; Gronella Barnard & Karaman, 1991;
Hippomedon Boeck, 1871; Lepidepecreoides K.H. Barnard, 1931; Lepidepecreum Spence Bate & Westwood,
1868; Lepiduristes Barnard & Karaman, 1987; Lysianella G.O. Sars, 1883; Metambasia Stephensen, 1923;
Microlysias Stebbing, 1918; Onesimoides Stebbing, 1888; Orchomene Boeck, 1871; Orchomenella G.O. Sars,
1890; Orchomenopsis G.O. Sars, 1890; Orchomenyx De Broyer, 1984; Orenoquia Bellan-Santini, 1997;

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Ottenwalderia Jaume & Wagner, 1998; Paracentromedon Chevreux & Fage, 1925; Paralysianopsis Schellenberg,
1931; Paratryphosites Stebbing, 1899; Paronesimoides Pirlot, 1933b; Patonga Lowry & Kilgallen, 2014b;
Photosella Lowry & Stoddart, 2011b; Psammonyx Bousfield, 1973; Pseudokoroga Schellenberg, 1931;
Pseudonesimoides Bellan-Santini & Ledoyer, 1974; Pseudonesimus Chevreux, 1926a; Pseudorchomene
Schellenberg, 1926a; Rifcus Kudrjaschov, 1965; Rimakoroga Barnard & Karaman, 1987; Schisturella Norman,
1900; Stephensenia Schellenberg, 1928a; Tasmanosa Lowry & Kilgallen, 2014b; Thrombasia J.L. Barnard, 1966;
Tryphosa Boeck, 1871; Trypho sella Bonnier, 1893; Tryphosi tes G.O. Sars, 1891; Ulladulla Lowry & Kilgallen,
2015; Wecomedon Jarrett & Bousfield, 1982.
Remarks. Tryphosoides Schellenberg, 1931 is considered to be a junior synonym of Cheirimedon Stebbing,
1888 by Kilgallen & Lowry (2015b).
The families Lysianassidae, Tryphosidae and Uristidae are based on morphological characters. Corrigan et al.
(2014), using molecular techniques, considered these family groups to be polyphyletic. Together these families
contain nearly 100 genera. For their analysis Corrigan et al. (2014) analysed selected species from only five genera
in the complex and four genera from other lysianassoid family groups, based on an unusual choice of outgroups.
This is not a basis for a reclassification of this large set of families.
By raising the Tryphosinae to family level we have considerably reduced the amount of morphological
diversity in the Lysianassidae. There may be further taxonomic groupings within the family based around genera
such as Hippomedon.
Family Uristidae Hurley, 1963
Included genera. Abyssorchomene De Broyer, 1984; Anonyx Krøyer, 1838; Caeconyx Barnard & Karaman, 1991;
Centromedon G.O. Sars, 1891; Cicadosa Barnard & Karaman, 1991; Debroyerella Lowry & Kilgallen, 2015; Des
Lowry & Kilgallen, 2014c; Eclecticus Lowry & Stoddart, 1997; Euonyx Norman, 1867; Galathella Barnard &
Karaman, 1987; Gippsia Lowry & Stoddart, 1995a; Ichnopus Costa, 1853; Koroga Holmes, 1908; Kyska
Shoemaker, 1964; Menigrates Boeck, 1871; Menigratopsis Dahl, 1945; Nagada Lowry & Stoddart, 1995b;
Onisimus Boeck, 1871; Paralibrotus Stephensen, 1923; Parschisturella Andres, 1983; Stephonyx Lowry &
Stoddart, 1989; Tmetonyx Stebbing, 1906; Uristes Dana, 1849.
Remarks. Based on the current concept of Abyssorchomene, it would form three different genera according to
the molecular study of Havermans et al. 2010.
Superfamily Aristioidea Lowry & Stoddart, 1997 stat. nov.
Diagnosis. Antenna 1–2 without calceoli (except Thoriellidae). Mouthpart bundle subquadrate. Gnathopod 2
mitten-shaped.
Included families. Acidostomatidae Stoddart & Lowry, 2012; Ambasiidae fam. nov.; Aristiidae Lowry &
Stoddart, 1997; Conicostomatidae Lowry & Stoddart, 2012b stat. nov.; Derjugianidae fam. nov.; Endevouridae
Lowry & Stoddart, 1997; Izinkalidae Lowry & Stoddart, 2010c; Kergueleniidae Lowry & Stoddart, 2010d;
Lepidepecreellidae Stoddart & Lowry, 2010b; Pakynidae nom. nov.; Sophrosynidae Lowry & Stoddart, 2010b;
Thoriellidae Lowry & Stoddart, 2011a; Trischizostomatidae Lilljeborg, 1865b; Wandinidae Lowry & Stoddart,
1990.
Remarks. The Aristioidea have ovoid or rounded eyes and non-triturating molars, they have entire telsons
(except ambasiids) and lack calceoli (except thoriellids).
Family Acidostomatidae Stoddart & Lowry, 2012
Included genera. Acidostoma Liljeborg, l865a; Shackletonia K.H. Barnard, 1931.

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Family Ambasiidae fam. nov.
Type genus. Ambasia Boeck, 1871.
Diagnosis (based on world family database). Antennae 1–2 calceoli absent. Mandible lacinia mobilis absent;
accessory setal row absent; molar absent. Maxilla 1 basal endite with apical setae. Maxilla 2 basal endite without
oblique setal row. Maxilliped ischial endite longer than palp article 2, not longer than palp article 3. Gnathopod 1
simple; coxa reduced, tapering. Gnathopod 2 minutely subchelate (mitten-shaped). Urosomite 1 dorsally carinate.
Uropods 1–2 rami without apical robust setae. Telson deeply cleft.
Habitat. Marine, epigean.
Included genera. Ambasia Boeck, 1871; Ambasiella Schellenberg, 1935.
Distribution. Arctic Ocean; North Atlantic, Greenland, Iceland, Norway, off south-west Ireland.
Remarks. The sister taxon of ambasiids appears to be the Izinkalidae. They share the absence of a lacinia
mobilis, a reduced coxa 1, expanded merus on pereopods 5 and 6, a carinate urosome and apical robust setae on the
telson.
Family Aristiidae Lowry & Stoddart, 1997
Included genera. Aristias Boeck, 1871; Boca Lowry & Stoddart, 1997; Memana Stoddart & Lowry, 2010a;
Perrierella Chevreux & Bouvier, 1892; Pratinas Stoddart & Lowry, 2010a.
Family Conicostomatidae Lowry & Stoddart, 2012b stat. nov.
Included genera. Acontiostoma Stebbing, 1888; Amphorites Lowry & Stoddart, 2012b; Conicostoma Lowry &
Stoddart, 1983; Ocosingo J.L. Barnard, 1964b; Scolopostoma Lowry & Stoddart, 1983; Stomacontion Stebbing,
1899.
Family Derjugianidae fam. nov.
Type genus. Derjugiana Gurjanova, 1962.
Diagnosis (based on world family database). Antenna 1 with well-developed callynophore. Antennae 1–2
calceoli apparently absent. Mandible incisors minutely dentate; accessory setal row absent; molar absent. Maxilla 2
basal endite without oblique setal row. Maxilliped ischial endites longer than palp article 3. Gnathopod 1 chelate.
Gnathopod 2 minutely chelate (mitten-shaped). Pleonite 3 and urosomite 1 each with dorsal carina. Uropod 3
present, rami absent. Telson entire.
Habitat. Marine, epigean.
Included genera. Derjugiana Gurjanova, 1962.
Distribution. Russia. Okhotsk Sea: Sakhalin Island (Gurjanova, 1962).
Remarks. Derjugianidae are in a clade with Thoriellidae and Wandinidae. They share a vertical anterior
margin on the head, 6/5 setal-tooth arrangement on the ischial endite of maxilla 1 (Wandinidae), maxilliped ischial
endite very large and uropod 3 endopod absent (Thoriellidae).
Family Endevouridae Lowry & Stoddart, 1997
Included genera. Endevoura Chilton, 1921; Ensayara J.L. Barnard, 1964d.
Family Izinkalidae Lowry & Stoddart, 2010c

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Included genera. Izinkala Griffiths, 1977.
Remarks. See remarks for Bolttsiidae.
Family Kergueleniidae Lowry & Stoddart, 2010d
Included genera. Clepidecrella J.L. Barnard, 1962c; Kerguelenia Stebbing, 1888.
Family Lepidepecreellidae Stoddart & Lowry, 2010b
Included genera. Lepidepecreella Schellenberg, 1926a.
Pakynidae nom. nov.
Pachynidae Lowry & Stoddart, 2012a: 5 (homonym).—De Broyer et al., 2007: 157 (nomen nudum).
Included genera. Acheronia Lowry, 1984; Coriolisa Lowry & Stoddart, 1994; Drummondia Lowry, 1984;
Ekelofia Lowry, 1984; Figorella J.L. Barnard, 1962c; Pachychelium Stephensen, 1925b; Pakynus nom. nov.;
Prachynella Barnard, 1964b; Renella Lowry & Stoddart, 2012; Sheardella Lowry, 1984; Smaraldia Lowry &
Stoddart, 2012; Ultimachelium Lowry & Stoddart, 2012.
Habitat. Marine, epigean.
Distribution. Cosmopolitan.
Remarks. It was recently brought to our attention by Tony Rees (CSIRO Marine and Atmospheric Research)
that the generic name Pachynus was a nomen nudum, originally proposed by Rafinesque (1815) for a genus of
cephalopod (preoccupied by Hippurites Lamarck, 1801), by Reichenow (1881) and for a psittaciforme bird
(preoccupied by Graydidascalus Bonaparte, 1854). Even though Pachynus Rafinesque, 1815 and Pachynus
Reichenow, 1881 are unjustified emendations, they are still available names (ICZN art. 33). Thus Pachynus
Rafinesque 1815 is the senior available name and Pachynus Bulycheva, 1955 must be considered as preoccupied.
We propose the new name Pakynus to replace Pachynus Bulycheva, 1955 in accordance with Article 39 of the
International Code of Zoological Nomenclature (1999, fourth edition).
Family Sophrosynidae Lowry & Stoddart, 2010b
Included genera. Sophrosyne Stebbing, 1888.
Family Thoriellidae Lowry & Stoddart, 2011a
Included genera. Chevreuxiella Stephensen, 1915; Danaella Stephensen, 1925a; Parachevreuxiella Andres, 1987;
Thoriella Stephensen, 1915.
Family Trischizostomatidae Lilljeborg, 1865b
Included genera. Trischizostoma Boeck, 1861.
Family Wandinidae Lowry & Stoddart, 1990
Included genera. Pseudocyphocaris Ledoyer, 1986; Wandin Lowry & Stoddart, 1990.

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Acknowledgements
We are grateful for the World Amphipod Database (Horton et al. 2017) and Wim Vader’s Amphipod Newsletter for
keeping us up to date on recent new taxa. We appreciate the generous help of Wolfgang Zeidler in solving
hyperiidean problems and the advice of Traudl Krapp-Schickel on the family Stenothoidae. We thank Roger
Springthorpe who was a major contributor to the original world family DELTA database and for producing the
cladogram graphics. We also thank Gillian Whitney who scanned many of the papers used in this study and other
studies we have published over the years. We are deeply appreciative for the constructive comments of Tammy
Horton, Sarah LeCroy, Mike Thurston, Wim Vader and Wolfgang Zeidler that greatly improved the paper.
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