Centropagidae

Male Boeckella gracilipes, from here.

Belongs within: Copepoda.

The Centropagidae are a group of pelagic copepods found in both fresh and salt water.

Characters (from J. M. Bradford-Grieve): Female with rostrum with 2 filaments; rostral margin not extended. Cephalosome and pedigerous somite 1 separate; pedigerous somites 4 and 5 separate. Posterior corners of prosome in lateral view bluntly rounded, or with 1 acute point, or divided into 2 protrusions; dorsal view symmetrical, or asymmetrical. Urosome with 3 free somites, or with 4 free somites. Genital double-somite asymmetrical in dorsal view; gonopores close together and totally covered by single genital operculum; without seminal receptacles. Caudal rami symmetrical. Antenna 1 with 24-25 free segments; ancestral segments X and XI partly fused. Antenna 2 exopod with 6-7 free segments; 10-11 setae; basis and endopod separate. Mandible basis with 4 setae; endopod terminal segment with 7-9 setae. Maxilla 1 basal exite seta present; exopod extends short of endopod distal border. Maxilla 2 coxal epipodite seta absent; endopod setae normal, or claw-like. Maxilliped coxal endite 2 with 3 setae. Leg 1 basis inner distal seta inserted directly on surface of joint; outer seta absent; endopod with 3 segments; exopod with 3 segments; exopod segment 1 with outer distal spine, or without outer distal spine; exopod segment 3 with 4 inner setae. Leg 2 endopod with 3 segments; exopod with 3 segments; exopod segment 3 with 2 outer spines, or with 3 outer spines. Leg 3 basis outer distal spine absent; endopod segment 3 with 5-8 setae. Legs 3 and 4 endopod with 3 segments; exopod with 3 segments; exopod segment 3 with 2 outer spines, or 3 outer spines; exopod segment 3 with 5 inner setae. Leg 4 endopod segment 3 with 4-7 setae. Legs 2-4 surfaces naked; terminal spine with outer border serrated. Leg 5 present; similar to legs 2-4; coxa inner border without seta; exopod segment 2 inner distal border extended into large spine completely fused to segment. Leg 5 exopod segment 3 outer border with 2 spines; inner border with 2 setae, or 4 setae. Male with urosome symmetrical in dorsal view, or asymmetrical in dorsal view. Mouthparts well-developed. Antenna 1 geniculate on right. Maxilliped endopod segments 5 and 6 with outer setae normal. Right leg 5 uniramous, or biramous; endopod 2-segmented, or 3-segmented; exopod 1-3-segmented, may be similar to other legs or may be highly modified into grasping organ. Left leg 5 uniramous, or biramous; endopod 2-segmented, or 3-segmented; exopod 2-3-segmented, may be similar to other legs or may be highly modified.

<==Centropagidae
    |--Pseudoboeckella F45
    |--Hemiboeckella searli Sars 1912 F45
    |--Eurytemora D01
    |    |--E. affinus K-M02
    |    |--E. herdmani B26
    |    `--E. velox D01
    |--Gladioferens Henry 1919 N44
    |    |--G. brevicornis Henry 1919 N44
    |    |--G. gracilis Kiefer 1931 N44
    |    |--G. inermis Nicholls 1944 N44
    |    |--G. pectinatus (Brady 1899) [=Centropages pectinatus] N44
    |    |--G. spinosus Henry 1919 N44
    |    `--G. subsalaria Percival 1937 N44
    |--Centropages Kroyer 1849 F44
    |    |--C. australiensis Fairbridge 1944 F44
    |    |--C. bradyi (Wheeler 1901) B26
    |    |--C. dorsispinatus ZS10
    |    |--C. hamatus (Lilljeborg 1853) F44
    |    |--C. kroyeri (Giesbrecht 1892) F44
    |    |--C. longicornis (Mori 1932) F44
    |    |--C. ponticus (Karawiew 1895) F44
    |    `--C. typicus (Kroyer 1848) F44
    |--Brunella Smith 1909 N44
    |    |--*B. tasmanica Smith 1909 N44
    |    |--B. ampulla Searle 1911 N44
    |    |--B. australis Searle 1911 N44
    |    |--B. expansa Sars 1912 N44
    |    |--B. longicornis Searle 1912 N44
    |    |--B. salina Nicholls 1944 N44
    |    |--B. steeli Henry 1924 N44
    |    `--B. viridis Searle 1911 N44
    `--Boeckella Deguerne & Richard 1889 F45
         |--B. ambigua Percival 1937 F45
         |--B. asymmetrica Searle 1914 F45
         |--B. bergi Richard 1897 [incl. B. bergi var. cornuta Brehm 1937, B. bergi var. serrifera Brehm 1937] F45
         |--B. coronaria Henry 1922 F45
         |--B. delicata Percival 1937 F45
         |--B. dentifera Brehm 1935 F45
         |--B. fluvialis Henry 1922 F45
         |--B. gracilipes Daday 1901 F45
         |--B. gracilis Daday 1902 F45
         |--B. hamata Brehm 1928 F45
         |--B. insignis Smith 1909 F45
         |--B. lacuna Fairbridge 1945 F45
         |--B. longiseta Smith 1909 F45
         |--B. major Searle 1938 F45
         |--B. meteoris Kiefer 1927 F45
         |--B. michaelseni Mrazek 1901 F45
         |--B. minuta Sars 1896 F45
         |--B. nyoraensis Searle 1912 F45
         |--B. occidentalis Marsh 1906 F45
         |--B. opaqua Fairbridge 1945 F45
         |--B. orientalis Sars 1903 F45
         |--B. pellucida Fairbridge 1945 F45
         |--B. poopoensis Marsh 1906 F45
         |--B. propinqua Sars 1904 F45
         |--B. pseudocheles Searle 1912 F45
         |--B. rahmi Brehm 1935 F45
         |--B. robusta Sars 1896 F45
         |--B. rubra Smith 1909 F45
         |--B. saycei Sars 1908 F45
         |--B. symmetrica Sars 1908 F45
         |--B. tenera Sars 1912 F45
         `--B. triarticulata Thompson 1883 (see below for synonymy) F45

Boeckella triarticulata Thompson 1883 [incl. B. triarticulata var. oblonga Brehm 1939, B. triarticulata var. quarta Brehm 1939, B. triarticulata var. sarsi Brehm 1939] F45

*Type species of generic name indicated

REFERENCES

[B26] Bigelow, H. B. 1926. Plankton of the offshore waters of the Gulf of Maine. Bulletin of the Bureau of Fisheries 40 (2): 1–509.

[D01] Daday, E. 1901. Édesvizi mikroszkópi állatok [Mikroskopische Süsswasserthiere]. In: Horváth, G. (ed.) Zichy Jenő Gróf Harmadik Ázsiai Utazása [Dritte Asiatische Forschungsreise des Grafen Eugen Zichy] vol. 2. Zichy Jenő Gróf Harmadik Ázsiai Utazásának Állattani Eredményei [Zoologische Ergebnisse der Dritten Asiatischen Forschungsreise des Grafen Eugen Zichy] pp. 375–470. Victor Hornyánszky: Budapest, and Karl W. Hierseman: Leipzig.

[F44] Fairbridge, W. S. 1944. Marine copepods from Western Australia. Series II. Two pelagic copepods from Cockburn Sound. Journal of the Royal Society of Western Australia 28: 209–217.

[F45] Fairbridge, W. S. 1945. West Australian fresh-water calanoids (Copepoda). I.–Three new species of Boeckella, with an account of the developmental stages of B. opaqua n. sp. and a key to the genus. Journal of the Royal Society of Western Australia 29: 25–65.

[K-M02] Klein-MacPhee, G. 2002. Temperate basses. Family Moronidae. In: Collette, B. B., & G. Klein-MacPhee (eds) Bigelow and Schroeder’s Fishes of the Gulf of Maine 3^rd ed. pp. 374–389. Smithsonian Institute Press: Washington.

[N44] Nicholls, A. G. 1944. Littoral Copepoda from South Australia. (II) Calanoida, Cyclopoida, Notodelphyoida, Monstrilloida and Caligoida. Records of the South Australian Museum 8 (1): 1–62.

[ZS10] Zhang, G.-T., S. Sun, Z.-L. Xu & Q.-L. Zhang. 2010. Unexpected dominance of the subtropical copepod Temora turbinata in the temperate Changjiang River estuary and its possible causes. In: China-Russia Bilateral Symposium: Proceedings of the China-Russia Bilateral Symposium of "Comparison on Marine Biodiversity in the Northwest Pacific Ocean", 10–11 October 2010, Qingdao (China) pp. 100–102. Institute of Oceanology, Chinese Academy of Sciences; A. V. Zhirmunsky Institute of Marine Biology, Far East Branch of the Russian Academy of Sciences.

Diaptomidae

Diaptomus sp., from the United States Great Lakes Environmental Research Laboratory.

Belongs within: Copepoda.

The Diaptomidae are a group of pelagic freshwater copepods.

Characters (from J. M. Bradford-Grieve): Female with rostrum without filaments; rostral margin extends into 2 points. Cephalosome and pedigerous somite 1 separate; pedigerous somites 4 and 5 separate. Posterior corners of prosome in lateral view with 1 acute point, or divided into 2 protrusions; dorsal view symmetrical. Urosome with 2 free somites, or with 3 free somites. Genital double-somite symmetrical in dorsal view; gonopores close together and partially covered by single cuticular flap; without seminal receptacles. Caudal rami symmetrical. Antenna 1 with 25 free segments. Antenna 2 exopod with 7 free segments; 12 setae; basis and endopod separate. Mandible basis with 3 setae; endopod terminal segment with 8 setae. Maxilla 1 basal exite seta present; exopod extends short of endopod distal border. Maxilla 2 coxal epipodite seta absent. Maxilliped coxal endite 2 with 2 setae. Leg 1 basis inner distal seta absent; outer seta absent; endopod with 2 segments; exopod with 3 segments; exopod segment 1 with outer distal spine; exopod segment 3 with 4 inner setae. Leg 2 endopod with 3 segments; exopod with 3 segments; exopod segment 3 with 1 outer spine. Leg 3 basis outer distal spine absent. Legs 3 and 4 endopod with 3 segments; exopod with 3 segments; exopod segment 3 with 1 outer spine; exopod segment 3 with 5 inner setae. Leg 4 endopod segment 3 with 7 setae. Legs 2-4 surfaces naked; terminal spine with outer border serrated. Leg 5 present; very dissimilar from legs 2-4; coxa inner border without seta; biramous but not natatory, may be slightly asymmetrical; coxae and intercoxal sclerite fused with outer distal tooth; basis with outer border seta; endopods 1-segmented, rod shaped; exopods 3-segmented, segment 1 cylindrical and unornamented, segment 2 tooth-like with inner border denticles and outer spinule, terminal segment very small and inserted subterminally bearing about 2 spinules. Male with pedigerous somites 4 and 5 fused or partly fused, or separate. Posterior corners of prosome in dorsal view symmetrical, or asymmetrical. Mouthparts well-developed. Antenna 1 geniculate on right. Maxilliped endopod segments 5 and 6 with outer setae normal. Right leg 5 biramous; endopod separate from basis and 1-segmented, or 2-segmented; exopod elongate, 3-segmented, segment 1 with outer distal spine, segment 2 with long spine on outer proximal border and sometimes additional spines and process, segment 3 curved, elongate and spine-like. Left leg 5 biramous; endopod 1-segmented, or 2-segmented; exopod short, 2-segmented, segment 1 naked, segment 2 very small with a few terminal spines.

<==Diaptomidae [Pseudodiaptomidae]
    |--Speodiaptominae B86
    |    |--Speodiaptomus Borutsky 1962 B86
    |    |    `--S. birsteini Borutsky 1962 B86
    |    `--Troglodiaptomus Petkovsky 1978 B86
    |         `--T. sketi Petkovsky 1978 B86
    `--Diaptominae B86
         |--Anadiaptomus Brehm 1952 B86
         |    `--A. poseidon [incl. A. poseidon var. spelaea Brehm 1953] B86
         |--Argyrodiaptomus Brehm 1933 B86
         |    `--A. cavernicolax Shen & Tai 1965 B86
         |--Microdiaptomus Osorio Tafall 1942 B86
         |    `--M. cokeri Osorio Tafall 1942 B86
         |--Notodiaptomus Kiefer 1936 B86
         |    `--N. caperatus Bowman 1979 B86
         |--Spelaeodiaptomus Dussart 1970 B86
         |    `--S. rouchi Dussart 1970 B86
         |--Stygodiaptomus Petkovski 1981 B86
         |    `--S. kieferi Petkovski 1981 B86
         `--Diaptomus D01
              |--D. asiaticus D01
              |--D. castor D37
              |--D. coeruleus D01
              |--D. denticornis D01
              |--D. gracilis D01
              |--D. minutus B68
              |--D. sanguineus R96
              |--D. spinosus D01
              `--D. zichyi Daday 1901 D01

Diaptomidae incertae sedis:
  Arctodiaptomus dorsalis (Marsh 1907) S-MR03
  Leptodiaptomus novamexicanus (Herrick 1895) S-MR03
  Mastigodiaptomus S-MR03
    |--M. albuquerquensis (Herrick 1895) S-MR03
    |--M. maya Suárez-Morales & Elías-Gutiérrez 2000 S-MR03
    |--M. nesus Bowman 1986 S-MR03
    |--M. reidae Suárez-Morales & Elías-Gutiérrez 2000 S-MR03
    `--M. texensis (Wilson 1953) S-MR03
  Poppella N44
  Pseudodiaptomus Herrick 1884 S-MR03, N44
    |--P. cornutus Nicholls 1944 N44
    |--P. marshi Wright 1936 S-MR03
    `--P. salinus N44

*Type species of generic name indicated

REFERENCES

[B86] Bowman, T. E. 1986. Copepoda: Calanoida. In: Botosaneanu, L. (ed.) Stygofauna Mundi: A Faunistic, Distributional, and Ecological Synthesis of the World Fauna inhabiting Subterranean Waters (including the Marine Interstitial) pp. 295–298. E. J. Brill/Dr W. Backhuys: Leiden.

[B68] Brooks, J. L. 1968. The effects of prey size selection by lake planktivores. Systematic Zoology 17 (3): 273–291.

[D01] Daday, E. 1901. Édesvizi mikroszkópi állatok [Mikroskopische Süsswasserthiere]. In: Horváth, G. (ed.) Zichy Jenő Gróf Harmadik Ázsiai Utazása [Dritte Asiatische Forschungsreise des Grafen Eugen Zichy] vol. 2. Zichy Jenő Gróf Harmadik Ázsiai Utazásának Állattani Eredményei [Zoologische Ergebnisse der Dritten Asiatischen Forschungsreise des Grafen Eugen Zichy] pp. 375–470. Victor Hornyánszky: Budapest, and Karl W. Hierseman: Leipzig.

[D37] Dobzhansky, T. 1937. Genetics and the Origin of Species. Columbia University Press: New York.

[N44] Nicholls, A. G. 1944. Littoral Copepoda from South Australia. (II) Calanoida, Cyclopoida, Notodelphyoida, Monstrilloida and Caligoida. Records of the South Australian Museum 8 (1): 1–62.

[R96] Roff, D. A. 1996. The evolution of threshold traits in animals. Quarterly Review of Biology 71 (1): 3–35.

[S-MR03] Suárez-Morales, E., & J. W. Reid. 2003. An updated checklist of the continental copepod fauna of the Yucatan Peninsula, Mexico, with notes on its regional associations. Crustaceana 76 (8): 977–991.

Cyclops

Cyclops sp., copyright David Nicholls.

Belongs within: Cyclopidae.

Cyclops is a cosmopolitan genus of freshwater copepods characterised by well-defined, angular epimeres on the anterior segments of the body, giving the sides a more or less jagged appearance, and a rudimentary last pair of legs (Sars 1913a).

Characters (from Sars 1913a): Anterior division of body moderately tumid, generally more tapered behind than in front; lateral parts of three anterior trunk segments well defined, sub-angular behind; last trunk segment more or less produced laterally. Tail slender and attenuated, genital segment in female more or less dilated in front. Anterior antennae of varying length, more generally composed of 17 well defined joints, though in some cases, by concrescence, their number may be considerably reduced; those in male strongly hinged, with terminal section distinctly biarticulate. Posterior antennae with all 4 joints well defined, the 1st carrying at the end posteriorly a long ciliated seta. Rudimentary palp of mandibles carrying 2 very long and densely plumose setae accompanied by a short bristle. Maxillary palp scarcely extending beyond masticatory lobe, provided outside near base with 4 ciliated setae, 3 of which issue from a slight expansion of the margin. Anterior maxillipeds moderately strong, with 1st basal joint generally subdivided in the middle, provided anteriorly with a small rounded lobe carrying 2 plumose setae; digitiform lobe issuing from end of this joint anteriorly comparatively small, with none of the setae unguiform. Posterior maxillipeds much more slender, composed of 4 well-defined joints. Natatory legs with both rami generally 3-articulate, though in some cases, by an imperfect separation of the 2 outer joints, one or other of them, or all, may be only biarticulate; inner ramus of 4th pair with 2 apical spines. Last pair of legs very small, generally not extended laterally, composed of 2 joints, the proximal of which in some cases may be coalesced with the corresponding segment, distal joint not expanded, and carrying a slender apical seta and a short lateral spine. Seminal receptacle in female of comparatively simple structure, being generally transversely oval in shape.

Cyclops Müller 1776 BR05
    |--C. abyssicola S13b
    |--C. abyssorum Sars 1863 E75 [=C. strenuus abyssorum E75; incl. C. strenuus var. tenuipes S13b]
    |    |--C. a. abyssorum E75
    |    |--C. a. apenninae Lindberg 1956 E75
    |    |--C. a. bodanus Kiefer 1954 [=C. rubens bodanus, C. strenuus bodanus] E75
    |    |--C. a. bohemicus (Sramek-Husek 1937) E75
    |    |--C. a. carinthicus Lindberg 1955 E75
    |    |--C. a. corsicanus (Lindberg 1955) [=C. rubens corsicanus] E75
    |    |--C. a. divergens Lindberg 1936 E75
    |    |--C. a. divulsus Lindberg 1956 E75
    |    |--C. a. gracilipes (Sars 1903) E75 [=C. strenuus var. gracilipes S13]
    |    |--C. a. laevis (Losito 1902) E75
    |    |--C. a. larianus (Stella 1934) E75
    |    |--C. a. maiorus Lindberg 1957 [incl. C. abyssorum f. resiae Stella 1934] E75
    |    |--C. a. mauritaniae (Lindberg 1950) E75
    |    |--C. a. medianus (Lindberg 1949) E75
    |    |--C. a. novarensis Einsle 1968 E75
    |    |--C. a. paternonis Lindberg 1956 [=C. rubens paternonis] E75
    |    |--C. a. prealpinus (Kiefer 1939) E75
    |    |--C. a. sevani (Mechkova 1947) E75
    |    |--C. a. sibiricus (Lindberg 1949) [=C. rubens sibiricus] E75
    |    |--C. a. tatricus (Kozminski 1927) [incl. C. tatricus tovelanus (Baldi 1941)] E75
    |    `--C. a. vranae (Kozminski 1927) E75
    |--C. arnaudi S13b
    |--C. attenuatus S13b
    |--C. bicolor S13b
    |--C. bisetosus S13b
    |--C. bohater Kozminski 1933 E75
    |--C. caeruleus [=Monoculus caeruleus] L02
    |--C. capillatus S13b
    |--C. captivus [=Monoculus captivus] L02
    |--C. chelifer [=Monoculus chelifer] L02
    |--C. claviger [=Monoculus claviger] L02
    |--C. columbianus Lindberg 1956 E75
    |--C. crassicaudis S13b
    |--C. cunningtoni S13b
    |--C. curticornis L02
    |--C. dengizica [=C. diaphanus var. dengizica] S13b
    |--C. diaphanus S13b [incl. Monoculus minutus L02, C. minutus S13b, C. nanus S13b]
    |--C. furcifer Claus 1857 E75
    |    |--C. f. furcifer E75
    |    |--C. f. borealis E75
    |    `--C. f. caspicus E75
    |--C. gigas [incl. C. ingens] S13b
    |--C. hutchinsoni Kiefer 1936 E75
    |--C. insignis Claus 1857 E75
    |--C. kolensis Lilljeborg 1901 E75
    |--C. lacinulatus L02
    |--C. lacustris Sars 1863 E75
    |--C. ladakanus Kiefer 1936 E75
    |--C. languidus S13b
    |--C. longicornis [=Monoculus longicornis; incl. C. finmarchius] L02
    |--C. lucidulus [incl. C. elongatus, C. parcus, C. vernalis] S13b
    |--C. minuticornis L02
    |--C. ochridanus Kiefer 1932 E75
    |--C. pachycomus S13b
    |--C. pictus S18
    |--C. pulchellus [incl. C. bicuspidatus, C. helgolandicus, C. lubbocki, C. odessanus] S13b
    |--C. robustus [incl. C. brevispinosus] S13b
    |--C. scmirnovi Rylov 1948 E75
    |--C. scutifer Sars 1863 E75
    |    |--C. s. scutifer E75
    |    |--C. s. laurenticus (Lindberg 1956) E75
    |    |--C. s. scandinavicus (Lindberg 1957) E75
    |    `--C. s. wigrensis Kozminski 1927 E75
    |--C. strenuus Fischer 1851 E75 (see below for synonymy)
    |    |--C. s. strenuus E75
    |    `--C. s. landei Kozminski 1933 E75
    |--C. thomasi S13b [=C. bicuspidatus thomasi K-M02]
    |--C. varicans [incl. C. orientalis, C. rubellus] S13b
    |--C. vicinus Uljanin 1875 [incl. C. v. var. glacialis] E75
    |    |--C. v. vicinus E75
    |    |--C. v. brevicornis E75
    |    |--C. v. kikuchii E75
    |    `--C. v. lobosus E75
    `--C. vulgaris [incl. C. viridis var. americana, Monoculus quadricornis viridis, C. viridis] S13b

Cyclops strenuus Fischer 1851 E75 [incl. C. brevicaudatus S13a, C. clausi S13a, Monoculus quadricornis Linnaeus 1758 L58, C. quadricornis S13a, M. quadricornis rubens S13a, C. rubens L02]

*Type species of generic name indicated

REFERENCES

[BR05] Bouchet, P., & J.-P. Rocroi. 2005. Classification and nomenclator of gastropod families. Malacologia 47 (1–2): 1–397.

[E75] Einsle, U. 1975. Revision der Gattuung Cyclops s. str. speziell der abyssorum-Gruppe. Memorie dell’Istituto Italiano di Idrobiologia 32: 57–219.

[K-M02] Klein-MacPhee, G. 2002. Temperate basses. Family Moronidae. In: Collette, B. B., & G. Klein-MacPhee (eds) Bigelow and Schroeder’s Fishes of the Gulf of Maine 3^rd ed. pp. 374–389. Smithsonian Institute Press: Washington.

[L02] Latreille, P. A. 1802b. Histoire Naturelle, générale et particulière des crustacés et des insectes vol. 4. Familles naturelles des genres. F. Dufart: Paris.

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

[S13a] Sars, G. O. 1913a. An Account of the Crustacea of Norway with short descriptions and figures of all the species vol. 6. Copepoda. Cyclopoida pts 1–2. Oithonidae, Cyclopinidae, Cyclopidae (part). Bergen Museum: Bergen.

[S13b] Sars, G. O. 1913b. An Account of the Crustacea of Norway with short descriptions and figures of all the species vol. 6. Copepoda. Cyclopoida pts 3–4. Cyclopidae (continued). Bergen Museum: Bergen.

[S18] Sars, G. O. 1918. An Account of the Crustacea of Norway with short descriptions and figures of all the species vol. 6. Copepoda. Cyclopoida pts 11–12. Lichomolgidae (concluded), Oncaeidae, Corycaeidae, Ergasilidae, Clausiidae, Eunicicolidae, supplement. Bergen Museum: Bergen.

Euchaetidae

Paraeuchaeta barbara, from Russ Hopcroft.

Belongs within: Copepoda.

The Euchaetidae are a group of bathypelagic copepods characterised by a prominent supralabrum bearing a rake formed from long stiff hairs.

Characters (from Park 1995): Supralabrum anterior to labrum, highly prominent with rows of long stiff hairs forming a 'rake'; antennule with segments 8 and 9 fused, extremely long seta on each of segments 3, 7, 9, 14, 18, 21, 24; second exopodal segment of male left P5 extending distally into serrated lamella and with a digitiform process; appendicular caudal setae highly developed, straight, smoothly curved, or geniculated, usually longer than marginal caudal setae.

<==Euchaetidae
    |--Euchaeta Philippi 1843 P95
    |    |  i. s.: E. ovata Sato 1913 P95
    |    |         E. wrighti Park 1968 P95
    |    |--E. spinosa Giesbrecht 1892 P95
    |    `--+--+--E. acuta Giesbrecht 1892 P95
    |       |  |--E. magniloba Park 1978 P95
    |       |  |--E. media Giesbrecht 1888 [incl. E. diegensis Esterly 1911, E. acuta var. pacifica Esterly 1911] P95
    |       |  |--E. paracuta Tanaka 1973 P95
    |       |  |--E. pubera Sars 1907 P95
    |       |  `--E. tenuis Esterly 1906 [incl. E. solida Esterly 1911] P95
    |       `--+--+--*E. marina (Prestandrea 1833) [=Cyclops marinus] P95
    |          |  |--E. indica Wolfenden 1905 [incl. E. wolfendeni Scott 1909] P95
    |          |  |--E. marinella Bradford 1974 P95
    |          |  `--E. rimana Bradford 1974 P95
    |          `--+--E. concinna Dana 1849 [incl. E. consimilis Farran 1936] P95
    |             |--E. longicornis Giesbrecht 1888 P95
    |             |--E. paraconcinna Fleminger 1957 [incl. E. gladiofera Gaudy 1963] P95
    |             `--E. plana Mori 1937 [incl. E. murrayi Sewell 1947] P95
    `--Paraeuchaeta Scott 1909 P95
         |  i. s.: P. abyssaloides Heptner 1987 P95
         |         P. affinis (Cleve 1904) [=Euchaeta affinis] P95
         |         P. bulbirostris Heptner 1987 P95
         |         P. flava (Giesbrecht 1888) [=Euchaeta flava] P95
         |         P. perplexa Heptner 1987 P95
         |         P. rotundirostris Heptner 1987 P95
         |         P. tridentata Heptner 1987 P95
         |--+--P. bisinuata (Sars 1907) [=Euchaeta bisinuata] P95
         |  `--+--P. antarctica (Giesbrecht 1902) [=Euchaeta antarctica] P95
         |     |--P. austrina (Giesbrecht 1902) [=Euchaeta austrina] P95
         |     |--P. erebi Farran 1929 P95
         |     |--P. similis (Wolfenden 1908) [=Euchaeta similis] P95
         |     `--P. tycodesma (Park 1978) [=Euchaeta tycodesma] P95
         `--+--+--P. elongata (Esterly 1913) [=Euchaeta elongata; incl. E. japonica Marukawa 1921] P95
            |  |--P. hebes (Giesbrecht 1888) [=Euchaeta hebes] P95
            |  |--P. russelli (Farran 1936) [=Euchaeta russelli; incl. E. daitomarui Mori 1937] P95
            |  `--P. simplex Tanaka 1958 P95
            `--+--+--P. glacialis (Hansen 1887) [=Euchaeta glacialis] P95
               |  `--P. tumidula (Sars 1905) (see below for synonymy) P95
               `--+--P. grandiremis (Giesbrecht 1888) [=Euchaeta grandiremis] P95
                  `--+--+--P. biloba Farran 1929 P95
                     |  `--+--*P. norvegica (Boeck 1872) [=Euchaeta norvegica] P95
                     |     |--P. exigua (Wolfenden 1911) [=Euchaeta exigua] P95
                     |     |--P. gracilis (Sars 1905) [=Euchaeta gracilis; incl. E. quadrata Farran 1908] P95
                     |     |--P. incisa (Sars 1905) [=Euchaeta incisa] P95
                     |     |--P. pseudotonsa (Fontaine 1967) [=Euchaeta pseudotonsa] P95
                     |     |--P. tonsa (Giesbrecht 1895) [=Euchaeta tonsa; incl. E. spinifera Esterly 1906] P95
                     |     |--P. tuberculata Scott 1909 [incl. Euchaeta scaphula Fontaine 1967] P95
                     |     `--P. weberi Scott 1909 P95
                     `--+--+--P. dactylifera (Park 1978) [=Euchaeta dactylifera] P95
                        |  |--P. pavlovskii Brodsky 1955 P95
                        |  |--P. scopaeorhina Park 1995 P95
                        |  |--P. sesquipedalis Park 1995 P95
                        |  `--P. vervoorti (Park 1978) [=Euchaeta vervoorti] P95
                        `--+--P. abbreviata (Park 1978) [=Euchaeta abbreviata] P95
                           |--P. abrikosovi Heptner 1971 P95
                           |--P. abyssalis Brodsky 1950 P95
                           |--P. aequatorialis Tanaka 1958 P95
                           |--P. alaminae (Park 1975) [=Euchaeta alaminae] P95
                           |--P. altibulla Park 1995 P95
                           |--P. anfracta Park 1995 P95
                           |--P. barbata (Brady 1883) (see below for synonymy) P95
                           |--P. birostrata Brodsky 1950 P95
                           |--P. bradyi (With 1915) [=Euchaeta bradyi] P95
                           |--P. brevirostris Brodsky 1950 [incl. P. laudabilis Tanaka & Omori 1968] P95
                           |--P. californica (Esterly 1906) [=Euchaeta californica; incl. E. dubia Esterly 1906] P95
                           |--P. calva Tanaka 1958 (see below for synonymy) P95
                           |--P. comosa Tanaka 1958 P95
                           |--P. confusa Tanaka 1958 P95
                           |--P. copleyae Park 1995 P95
                           |--P. eltaninae (Park 1978) [=Euchaeta eltaninae] P95
                           |--P. eminens Tanaka & Omori 1968 P95
                           |--P. euryrhina Park 1995 P95
                           |--P. gracilicauda Scott 1909 P95
                           |--P. guttata Heptner 1971 P95
                           |--P. hansenii (With 1915) [=Euchaeta hansenii; incl. P. withi Sewell 1947] P95
                           |--P. hastata Heptner 1987 P95
                           |--P. implicata Heptner 1987 P95
                           |--P. investigatoris Sewell 1929 P95
                           |--P. kurilensis Heptner 1971 [incl. Euchaeta longissima Park 1978] P95
                           |--P. longisetosa Heptner 1971 P95
                           |--P. malayensis Sewell 1929 P95
                           |--P. megaloba Park 1995 P95
                           |--P. mexicana Park 1995 P95
                           |--P. modesta Brodsky 1950 P95
                           |--P. oculata Heptner 1971 P95
                           |--P. orientalis Brodsky 1950 P95
                           |--P. papilliger Park 1995 P95
                           |--P. parabbreviata Park 1995 P95
                           |--P. paraprudens Park 1995 P95
                           |--P. parvula (Park 1978) [=Euchaeta parvula] P95
                           |--P. plaxiphora Park 1995 P95
                           |--P. plicata Heptner 1971 P95
                           |--P. polaris Brodsky 1950 P95
                           |--P. prima Heptner 1971 P95
                           |--P. propinqua (Esterly 1906) [=Euchaeta propinqua] P95
                           |--P. prudens Tanaka & Omori 1968 P95
                           |--P. rasa Farran 1929 P95
                           |--P. regalis (Grice & Hulsemann 1968) [=Euchaeta regalis] P95
                           |--P. robusta (Wolfenden 1911) [=Euchaeta robusta] P95
                           |--P. rubicunda (Farran 1908) [=Euchaeta rubicunda] P95
                           |--P. rubra Brodsky 1950 [incl. P. crassa Tanaka 1958] P95
                           |--P. sarsi (Farran 1908) [=Euchaeta sarsi; incl. P. dentata Scott 1909] P95
                           |--P. scotti (Farran 1908) [=Euchaeta scotti] P95
                           |--P. sibogae Scott 1909 P95
                           |--P. subtilirostris Heptner 1971 P95
                           |--P. triloba Park 1995 P95
                           `--P. vorax (Grise & Hulsemann 1968) (see below for synonymy) P95

Paraeuchaeta barbata (Brady 1883) [=Euchaeta barbata; incl. E. farrani With 1915, E. porrecta Sars 1905] P95

Paraeuchaeta calva Tanaka 1958 [incl. P. polita Tanaka & Omori 1968, P. simulantis Tanaka & Omori 1968] P95

Paraeuchaeta tumidula (Sars 1905) [=Euchaeta tumidula; incl. E. biconvexa Park 1978, Paraeuchaeta pseudotumidula Brodsky 1950] P95

Paraeuchaeta vorax (Grise & Hulsemann 1968) [=Euchaeta vorax; incl. P. striata Tanaka & Omori 1968] P95

*Type species of generic name indicated

REFERENCES

[P95] Park, T. 1995. Taxonomy and distribution of the marine calanoid copepod family Euchaetidae. Bulletin of the Scripps Institution of Oceanography, University of California, San Diego 29: i-xi, 1-203.

Siphonostomatoida

Specimens of Cancerilla on the disc of the brittle star Ophiacantha antarctica, photographed by Rafael Martín-Ledo.

Belongs within: Podoplea.
Contains: Asterocheridae, Nicothoidae, Dyspontiidae, Caligoida.

The Siphonostomatoida are a group of parasitic marine copepods with sucking mouthparts.

Characters (from Nicholls 1944): Second antenna usually with an exopod; mouth-parts suctorial; maxillae and maxillipeds sub-chelate.

<==Siphonostomatoida [Siphonostoma, Siphonostomata]
    |--Brychiopontius [Brychiopontiidae] BO01
    |    `--B. falcatus BO01
    `--+--+--Calverocheres [Calverocheridae] BO01
       |  |    `--C. globosus BO01
       |  `--+--Codoba Heegaard 1951 [Codobidae] BO01
       |     |    `--*C. discoveryi Heegaard 1951 B88
       |     `--Cancerillidae BO01
       |          |--Ophiopsyllus reductus BO01
       |          |--Parophiopsyllus ligatus BO01
       |          |--Parartotrogus Scott & Scott 1893 BO01
       |          |    |--P. arcticus BO01
       |          |    `--P. richardi BK77
       |          `--Cancerilla IFS01
       |               |--C. durbanensis BK77
       |               |--C. neozelanica BK77
       |               `--C. tubulata Dalyell 1851 IFS01 [incl. Caligidium vagabundum Claus 1889 BK77]
       `--+--Micropontius [Micropontiidae] BO01
          |    |--M. glaber BO01
          |    `--M. ovoides BO01
          `--+--Stellicomitidae BO01
             |    |--Stellicomes tumidulus BO01
             |    `--Onychopygus impavidus BO01
             `--+--Dichelina Stephensen 1933 [Dichelinidae] BO01
                |    |--D. phormosomae Stephensen 1933 BO01
                |    `--D. seticauda BO01
                `--Nanaspididae [Nanaspidae] BO01
                     |--Allantogynus Changeux 1958 BO01
                     |    `--A. delamarei BO01
                     |--Stephopontius Thompson & Scott 1903 BO01
                     |    `--S. typicus BO01
                     `--Nanaspis Humes & Cressey 1959 BO01
                          |--N. mixta BO01
                          `--N. ninae BK77

Siphonostomatoida incertae sedis:
  Scottomyzon Giesbrecht 1897 [Scottomyzontidae] IFS01
    `--*S. gibberum (Scott & Scott 1894) [=Dermatomyzon gibberum] IFS01
  Asterocheridae IFS01
  Dioithona oculata (Farran 1913) IFS01
  Bariaka IFS01
  Jushyus IFS01
  Nicothoidae BH88
  Artotrogidae MD01
    |--Dystrogus uncinatus Brady 1910 N44
    `--Artotrogus Boeck 1859 N44
         |--*A. orbicularis N44
         |--A. australis Wilson 1923 N44
         |--A. gigas Brady 1910 N44
         |--A. latifurcatus Nicholls 1944 N44
         `--A. sphaericus Brady 1910 N44
  Dyspontiidae N44
  Myzopontius Giesbrecht 1895 [Myzopontiidae] N44
    |--*M. pungens N44
    `--M. australis Nicholls 1944 N44
  Acontiophorus Brady 1880 [=Solenostoma Brady & Robertson 1873 (preoc.); Acontiophoridae] N44
    |--A. attenuatus Hansen 1923 N44
    |--A. ornatus (Brady & Robertson 1875) [incl. A. armatus Brady 1880] N44
    |--A. scutatus (Brady & Robertson 1873) [incl. A. angulatus Thompson 1888] N44
    `--A. zealandicus Nicholls 1944 N44
  Archidactylinidae MD01
  Caligoida MD01
  Cecrops K-M02d [Cecropidae MD01]
    `--C. latereilli K-M02d
  Coralliomyzontidae MD01
  Dichelesthium M02 [Dichelesthiidae MD01]
    `--D. oblongum M02
  Dinopontiidae MD01
  Dirivultidae MD01
  Dissonidae MD01
  Ecbathyriontidae MD01
  Entomolepididae MD01
  Eudactylinidae MD01
  Euryphoridae MD01
  Hatschekia K-M02c [Hatschekiidae MD01]
    `--H. reinhardtii K-M02c
  Hyponeoidae MD01
  Kroyeriidae MD01
  Lernaeopoda B02 [Lernaeopodidae MD01]
    `--L. centroscyllii B02
  Lernanthropus D83 [Lernanthropidae MD01]
    |--L. gisleri D83
    |--L. pomatomi K-M02a
    |--L. trachuri K-M02b
    `--L. tylosuri C02
  Megapontiidae MD01
  Pandaridae MD01
  Pennellidae MD01
    |--Cardiodectes H01
    |--Haemobaphaes cyclopterina K-MC02b
    `--Penella W81
         |--P. balaenopterae W81
         `--P. filosa K-M02d
  Pontoeciellidae MD01
  Pseudocycnidae MD01
  Rataniidae MD01
  Sphyrion [Sphyriidae MD01]
    `--S. lumpi K-MC02a
  Sponginticolidae MD01
  Spongiocnizontidae MD01
  Tanypleuridae MD01
  Trebiidae MD01

*Type species of generic name indicated

REFERENCES

[BK77] Barel, C. D. N., & P. G. N. Kramers. 1977. A survey of the echinoderm associates of the north-east Atlantic area. Zoologische Verhandelingen 156: 1-159.

[B88] Boxshall, G. A. 1988. A review of the copepod endoparasites of brittle stars. Bulletin of the British Museum (Natural History), Zoology series 54 (6): 261-270.

[BH88] Boxshall, G. A., & K. Harrison. 1988. New nicothoid copepods (Copepoda: Siphonostomatoida) from an amphipod and from deep-sea isopods. Bulletin of the British Museum (Natural History), Zoology series 54 (6): 285-299.

[BO01] Boxshall, G. A., & S. Ohtsuka. 2001. Two new families of copepods (Copepoda: Siphonostomatoida) parasitic on echinoderms. Journal of Crustacean Biology 21 (1): 96-105.

[B02] Burgess, G. H. 2002. Spiny dogfishes. Family Squalidae. In Bigelow and Schroeder’s Fishes of the Gulf of Maine (B. B. Collette & G. Klein-MacPhee, eds) 3^rd ed. pp. 48-57. Smithsonian Institute Press: Washington.

[C02] Collette, B. B. 2002. Needlefishes. Family Belonidae. In Bigelow and Schroeder’s Fishes of the Gulf of Maine (B. B. Collette & G. Klein-MacPhee, eds) 3^rd ed. pp. 287-289. Smithsonian Institute Press: Washington.

[D83] Darovec, J. E. 1983. Sciaenid fishes (Osteichthyes: Perciformes) of western penisular Florida. Memoirs of the Hourglass Cruises 6 (3): 1-73.

[H01] Huys, R. 2001. Splanchnotrophid systematics: A case of polyphyly and taxonomic myopia. Journal of Crustacean Biology 21 (1): 106-156.

[IFS01] Ivanenko, V. N., F. D. Ferrari & A. V. Smurov. 2001. Nauplii and copepodids of Scottomyzon gibberum (Copepoda: Siphonostomatoida: Scottomyzontidae, a new family), a symbiont of Asterias rubens (Asteroidea). Proceedings of the Biological Society of Washington 114 (1): 237-261.

[K-M02a] Klein-MacPhee, G. 2002a. Bluefish. Family Pomatomidae. In Bigelow and Schroeder’s Fishes of the Gulf of Maine (B. B. Collette & G. Klein-MacPhee, eds) 3^rd ed. pp. 400-406. Smithsonian Institute Press: Washington.

[K-M02b] Klein-MacPhee, G. 2002b. Jacks. Family Carangidae. In Bigelow and Schroeder’s Fishes of the Gulf of Maine (B. B. Collette & G. Klein-MacPhee, eds) 3^rd ed. pp. 411-427. Smithsonian Institute Press: Washington.

[K-M02c] Klein-MacPhee, G. 2002c. Righteye flounders. Family Pleuronectidae. In Bigelow and Schroeder’s Fishes of the Gulf of Maine (B. B. Collette & G. Klein-MacPhee, eds) 3^rd ed. pp. 560-587. Smithsonian Institute Press: Washington.

[K-M02d] Klein-MacPhee, G. 2002d. Ocean sunfishes. Family Molidae. In Bigelow and Schroeder’s Fishes of the Gulf of Maine (B. B. Collette & G. Klein-MacPhee, eds) 3^rd ed. pp. 603-607. Smithsonian Institute Press: Washington.

[K-MC02a] Klein-MacPhee, G., & B. B. Collette. 2002a. Scorpionfishes. Family Scorpaenidae. In Bigelow and Schroeder’s Fishes of the Gulf of Maine (B. B. Collette & G. Klein-MacPhee, eds) 3^rd ed. pp. 331-338. Smithsonian Institute Press: Washington.

[K-MC02b] Klein-MacPhee, G., & B. B. Collette. 2002b. Eelpouts. Family Zoarcidae. In Bigelow and Schroeder’s Fishes of the Gulf of Maine (B. B. Collette & G. Klein-MacPhee, eds) 3^rd ed. pp. 466-474. Smithsonian Institute Press: Washington.

[MD01] Martin, J. W., & G. E. Davis. 2001. An updated classification of the Recent Crustacea. Natural History Museum Los Angeles County, Science Series 39: 1-124.

[M02] Musick, J. A. 2002. Sturgeons. Family Acipenseridae. In Bigelow and Schroeder’s Fishes of the Gulf of Maine (B. B. Collette & G. Klein-MacPhee, eds) 3^rd ed. pp. 83-88. Smithsonian Institute Press: Washington.

[N44] Nicholls, A. G. 1944. Littoral Copepoda from South Australia. (II) Calanoida, Cyclopoida, Notodelphyoida, Monstrilloida and Caligoida. Records of the South Australian Museum 8 (1): 1-62.

[W81] Watson, L. 1981. Sea Guide to Whales of the World. Hutchinson: London.

Pseudanthessius

Diagnostic figures of Pseudanthessius tenuis, from Nicholls (1944).

Belongs within: Poecilostomatoida.

Pseudanthessius is a genus of marine parasitic copepods associated with a wide range of invertebrate hosts of numerous phyla.

Characters (from Humes & Stock 1973): Body cyclopiform. Urosome in female 5-segmented (occasionally 4-segmented), in male 6-segmented (occasionally 5-segmented). Caudal ramus with six setae. Rostrum not well developed. First antenna 7-segmented, in female with armature 4, 13, 6, 3, 4 + 1 aesthete, 2 + 1 aesthete, and 7 + 1 aesthete; in male with same armature or in some species with additional aesthete on second segment. Second antenna 4-segmented, with formula 1,1,3 and variable terminal armature (1-4 claws or clawlike spines and a few setae; or, in P. tortuosus, two lamelliform elements, one seta, and three spines; or, in P. mucronatus, all elements setiform). Labrum deeply incised medially. Mandible with basal region having its convex edge smooth or with a weak scalelike area or a process and on concave edge a row of spinules; lash long or short. Paragnath a small lobe. First maxilla with three or four elements. Second maxilla of usual lichomolgid type. Maxilliped in female 3-segmented, with pointed tip; in male 4-segmented (assuming that proximal part of the claw represents a fourth segment). Legs 1-4 with 3-segmented rami, except leg 4 endopod which is 1-segmented with two distal elements. Leg 1 endopod of male same armature as in female, or with same formula but with slight differences in form of processes or elements, or in P. deficiens 1,1,4 or in both P. tortuosus and P. aestheticus 1,1,1,4. Leg 5 represented by three elements (subterminal seta and terminal spine and seta); no free segment. Leg 6 represented by two setae near genital openings.

<==Pseudanthessius Claus 1889 [Pseudanthessiidae] N44
    |--P. assimilis Sars 1917 N44
    |--P. concinnus Thompson & Scott 1903 N44
    |--P. dubius Sars 1918 N44
    |--P. gracilis Claus 1889 N44
    |--P. liber (Brady & Robertson 1875) N44 [=Lichomolgus liber BK77]
    |--P. mucronatus Gurney 1927 N44
    |--P. nemertophilus Gallien 1935 N44
    |--P. obscurus Scott 1909 N44
    |--P. sauvagei Canu 1891 N44
    |--P. tenuis Nicholls 1944 N44
    |--P. thorelli (Brady & Robertson 1875) N44 [=Lichomolgus thorelli BK77]
    `--P. weberi Scott 1909 N44

*Type species of generic name indicated

REFERENCES

[BK77] Barel, C. D. N., & P. G. N. Kramers. 1977. A survey of the echinoderm associates of the north-east Atlantic area. Zoologische Verhandelingen 156: 1-159.

Humes, A. G., & J. H. Stock. 1973. A revision of the family Lichomolgidae Kossmann, 1877, cyclopoid copepods mainly associated with marine invertebrates. Smithsonian Contributions to Zoology 127.

[N44] Nicholls, A. G. 1944. Littoral Copepoda from South Australia. (II) Calanoida, Cyclopoida, Notodelphyoida, Monstrilloida and Caligoida. Records of the South Australian Museum 8 (1): 1-62.

Poecilostomatoida

Drawing of Paranthessius anemoniae, from here.

Belongs within: Podoplea.
Contains: Splanchnotrophidae, Clausiidae, Clausidiidae, Lichomolgidae, Pseudanthessius.

The Poecilostomatoida are a group of copepods that are generally parasitic on other marine animals.

Characters (from Nicholls 1944): First antenna not hinged in male; second antenna without an exopod; mouth parts non-suctorial, non-raptorial, mandible lacking palp.

<==Poecilostomatoida [Poecilostoma]
    `--Lichomolgoidea [Lichomolginae] K03
         |--Lichomolgidae K03
         |--Pseudanthessius K03
         `--Rhynchomolgidae K03
              |--Critomolgus anthopleurus Kim 1996 K03
              |--Doridicola longicauda K03
              `--Paranthessius Claus 1889 K03, N44 [Sabelliphilidae, Sabelliphilinae]
                   |--P. anemoniae K03
                   |--‘Heteranthessius’ dubius (Scott 1903) N44
                   |--‘Pestalichomolgus’ pectinis (Pesta 1908) N44
                   `--P. propinquus Nicholls 1944 N44

Poecilostomatoida incertae sedis:
  Pionodesmatidae BO01
  Synapticolidae MD01
  Antheacheridae MD01
  Echiurophilus [Echiurophilidae] H01
  Splanchnotrophidae H01
  Gastrodelphyidae H01
  Clausiidae HK03
  Micrallecto Stock 1971 [incl. Nannallecto Stock 1973; Micrallectidae] H09
    |--*M. uncinata Stock 1971 H01
    `--M. fusii Stock 1973 [=*Nannallecto fusii] H01
  Synaptiphilus Canu & Cuénot 1892 [Synaptiphilidae] HK03
    |--S. cantacuzenei BK77
    |--S. luteus [incl. Colaceutes mulleri Hartmann 1856] BK77
    `--S. tridens (Scott & Scott 1893) [=Remigulus tridens] BK77
  Chondracanthidae H01
    |  i. s.: Diocus lycenchelus K-MC02
    |--Pharodinae [Pharodidae] H01
    |--Lernentominae H01
    `--Chondracanthus K-M02c [Chondracanthinae H01]
         |--C. cottunculi K-M02c
         `--C. merluccii K-M02b
  Philoblennidae H01
    |--Philoblenna arabici K03
    |--Chondrocarpus Bassett-Smith 1903 H01
    |    `--*C. reticulosus H01
    `--Briarella Bergh 1876 H01
         |--*B. microcephala Bergh 1876 H01
         |--B. disphaerocephala Monod & Dollfus 1932 H01
         `--B. risbeci Monod 1928 H01
  Clausidiidae MD01
  Lernaeosoleidae [Lernaeosoleinae] MD01
  Ergasilidae [Amazonicopeidae] MD01
    |--Neoergasilus japonicus K03
    `--Ergasilus K03
         |--E. briani K03
         |--E. funduli K-M02a
         |--E. labracis K-M02d
         `--E. sieboldi K03
  Anchimolgidae MD01
  Anomoclausiidae MD01
  Anthessiidae MD01
  Bomolochidae MD01
  Chitonophilidae MD01
  Corallovexiidae MD01
  Corycaeus B26 [Corycaeidae MD01]
    |--C. carinatus B26
    |--C. elongatus B26
    |--C. ovalis B26
    `--C. speciosus B26
  Entobiidae MD01
  Erebonasteridae MD01
  Eunicicolidae MD01
  Herpyllobiidae MD01
  Intramolgidae MD01
  Lamippidae MD01
  Lubbockiidae MD01
  Mesoglicolidae MD01
  Pseudomyicola K03 [Myicolidae MD01]
    `--P. spinosus K03
  Trochicola H01 [Mytilicolidae MD01]
    `--T. entericus Dollfus 1914 H01
  Nereicolidae [Anomopsyllidae] MD01
    |--Anomopsyllus pranizoides N44
    `--Dysgenopsyllus Nicholls 1944 N44
         `--*D. reevesbyensis Nicholls 1944 N44
  Nucellicolidae MD01
  Oncaea B26 [Oncaeidae MD01]
    |--O. conifera B26
    |--O. minuta B26
    `--O. venusta B26
  Paralubbockiidae MD01
  Philichthyidae MD01
  Phyllodicolidae MD01
  Polyankylidae MD01
  Saccopsidae MD01
  Sapphirinidae MD01
  Serpulidicolidae MD01
  Shiinoidae MD01
  Spiophanicolidae MD01
  Taeniacanthus K03 [Taeniacanthidae MD01]
    |--T. balistae K-M02e
    `--T. lagocephali K03
  Tegobomolochidae MD01
  Telsidae MD01
  Thamnomolgidae MD01
  Tucca K-M02f [Tuccidae MD01]
    `--T. impressus K-M02f
  Urocopiidae MD01
  Vahiniidae MD01
  Ventriculinidae MD01
  Xarifia H94 [Xarifiidae MD01]
    |--X. diminuta Humes & Ho 1967 H94
    |--X. finitima Humes 1985 H94
    `--X. longipes Humes 1962 H94
  Xenocoelomatidae MD01

*Type species of generic name indicated

REFERENCES

[BK77] Barel, C. D. N., & P. G. N. Kramers. 1977. A survey of the echinoderm associates of the north-east Atlantic area. Zoologische Verhandelingen 156: 1-159.

[B26] Bigelow, H. B. 1926. Plankton of the offshore waters of the Gulf of Maine. Bulletin of the Bureau of Fisheries 40 (2): 1-509.

[BH88] Boxshall, G. A., & A. G. Humes. 1988. A new genus of Lichomolgidae (Copepoda: Poecilostomatoida) associated with a phoronid in Hong Kong. Bulletin of the British Museum (Natural History), Zoology Series 54 (6): 301-307.

[BO01] Boxshall, G. A., & S. Ohtsuka. 2001. Two new families of copepods (Copepoda: Siphonostomatoida) parasitic on echinoderms. Journal of Crustacean Biology 21 (1): 96-105.

[HK03] Ho, J.-S., & I.-H. Kim. 2003. New clausiid copepods (Poecilostomatoida) associated with polychaetes of Korea, with cladistic analysis of the family Clausiidae. Journal of Crustacean Biology 23 (3): 568-581.

[H94] Humes, A. G. 1994. Two species of Paramolgus (Copepoda: Poecilostomatoida: Lichomolgidae) associated with the scleractinian Pavona in New Caledonia with a key to the females of Paramolgus. Beaufortia 44 (1): 1-9.

[H01] Huys, R. 2001. Splanchnotrophid systematics: A case of polyphyly and taxonomic myopia. Journal of Crustacean Biology 21 (1): 106-156.

[K03] Kim, I.-H. 2003. Copepodid stages of Critomolgus anthopleurus (Copepoda, Poecilostomatoida, Rhynchomolgidae). Journal of Crustacean Biology 23 (3): 558-567.

[K-M02a] Klein-MacPhee, G. 2002a. Smelts. Family Osmeridae. In Bigelow and Schroeder’s Fishes of the Gulf of Maine (B. B. Collette & G. Klein-MacPhee, eds) 3^rd ed. pp. 162-170. Smithsonian Institute Press: Washington.

[K-M02b] Klein-MacPhee, G. 2002b. Silver hakes. Family Merlucciidae. In Bigelow and Schroeder’s Fishes of the Gulf of Maine (B. B. Collette & G. Klein-MacPhee, eds) 3^rd ed. pp. 217-222. Smithsonian Institute Press: Washington.

[K-M02c] Klein-MacPhee, G. 2002c. Fathead sculpins. Family Psychrolutidae. In Bigelow and Schroeder’s Fishes of the Gulf of Maine (B. B. Collette & G. Klein-MacPhee, eds) 3^rd ed. pp. 357-358. Smithsonian Institute Press: Washington.

[K-M02d] Klein-MacPhee, G. 2002d. Temperate basses. Family Moronidae. In Bigelow and Schroeder’s Fishes of the Gulf of Maine (B. B. Collette & G. Klein-MacPhee, eds) 3^rd ed. pp. 374-389. Smithsonian Institute Press: Washington.

[K-M02e] Klein-MacPhee, G. 2002e. Triggerfishes. Family Balistidae. In Bigelow and Schroeder’s Fishes of the Gulf of Maine (B. B. Collette & G. Klein-MacPhee, eds) 3^rd ed. pp. 590-592. Smithsonian Institute Press: Washington.

[K-M02f] Klein-MacPhee, G. 2002f. Porcupinefishes. Family Diodontidae. In Bigelow and Schroeder’s Fishes of the Gulf of Maine (B. B. Collette & G. Klein-MacPhee, eds) 3^rd ed. pp. 601-602. Smithsonian Institute Press: Washington.

[K-MC02] Klein-MacPhee, G., & B. B. Collette. 2002. Eelpouts. Family Zoarcidae. In Bigelow and Schroeder’s Fishes of the Gulf of Maine (B. B. Collette & G. Klein-MacPhee, eds) 3^rd ed. pp. 466-474. Smithsonian Institute Press: Washington.

[MD01] Martin, J. W., & G. E. Davis. 2001. An updated classification of the Recent Crustacea. Natural History Museum Los Angeles County, Science Series 39: 1-124.

[N44] Nicholls, A. G. 1944. Littoral Copepoda from South Australia. (II) Calanoida, Cyclopoida, Notodelphyoida, Monstrilloida and Caligoida. Records of the South Australian Museum 8 (1): 1-62.

Podogennonta

Diagnostic features of Enhydrosoma curticaudatum, from Sars (1909).

Belongs within: Oligoarthra.
Contains: Leptastacus, Paraleptastacus, Psammastacus, Leptocaris, Amphiascopsis, Amphiascus, Mesamphiascus, Amphiascoides, Robertsonia, Schizopera, Arenopontia, Psammopsyllinae, Stenocaris, Phyllopodopsyllus, Parastenocaris, Forficatocaris, Remaneicaris, Cletodes, Diosaccinae, Stenhelia, Bryocamptus, Attheyella, Elaphoidella, Moraria, Nitocrella, Parapseudoleptomesochra, Nitokra, Stygonitocrella, Laophonte.

The Podogennonta is a major clade of harpacticoid copepods, many of which are interstitial in habitat. Members of the clade are divided between numerous families, some of which (such as Parastenocarididae and Canthocamptidae) are primarily freshwater whereas others (such as Ameiridae and Laophontidae) are predominantly marine. The Laophontidae have a distinctive first leg structure with the two-segmented endopod longer than the one- to three-segmented exopod and ending in a protruding claw.

Synapomorphies (from Seifried 2003): Females with endopod of mandible with 3 proximal lateral setae, 3 more distal lateral setae and 3 + 2 + 2 apical setae; endopod of maxillule with 4 setae; syncoxa of maxilla with (3 + 3), 3, 3 setae, the two proximal endites fused, distal praecoxal endite with 2 flagellate setae; both coxal endites with setae of characteristic shape: anteriorly 1 claw-like seta, posteriorly 2 strong, mostly unilaterally spinulose setae with subterminal flagellum; acessory armature of fused endopod segment: 1 seta on posterior surface, seta 9 displaced to proximal segment of endopod, seta 10 lacking; endopod 3-segmented with armature formula: 2, 1, 3; setae 4 and six lost. Maxilliped 4-segmented with praecoxa and coxa separated, basis, and 1-segmented endopod; endopodal setae 1 to 4 only represented by a group of 4 small setae on anterior surface of endopod, no setae geniculated. Leg 1 of characteristic shape: endopod-1 elongate, endopod-2 and endopod-3 short; endopod-3 with 1 innter seta, terminally with anterior claw, 1 middle geniculate seta and 1 posterior miniaturised seta; 1 inner seta lacking; exopod-3 terminally with 2 geniculated setae, 2 inner setae lacking. Formula of armature of leg 1: coxa 0-0, basis I-I, exopod I-0, I-1, III-2-0, endopod 0-1, 0-1, I-2-1. Leg 5 with endopodal lobe with six setae: 3 inner setae, 2 terminal setae, 1 outer seta; exopod foliated with 8 setae: 3 inner, 2 terminal, 3 outer setae.

Podogennonta
    |--Argestidae S03
    |--Adenopleurellidae S03
    |--Cancrincolidae S03
    |--Cletopsyllus Willey 1935 N41b [Cletopsyllidae S03, Cletopsyllinae]
    |--Cristacoxidae S03
    |--Normanella Brady 1880 N41b [Normanellidae S03, Normanellinae]
    |--‘Stenhelia’ glacialis Brady 1918 N41b [Pseudotachidiidae S03, Pseudotachidiinae]
    |--Ismardis Leigh-Sharpe 1936 [Ismardiidae] S03
    |--Thompsonula N41b [Thompsonulidae S03]
    |    `--T. hyaenae (Thompson 1889) [incl. Robertsonia aculeifera Klie 1913] N41b
    |--Balaenophilus W81 [Balaenophilidae S03]
    |    `--B. unisetus W81
    |--Lourinia Wilson 1924 N41a (see below for synonymy)
    |    `--L. armata (Claus 1866) W67 (see below for synonymy)
    |--Laophontopsis Sars 1908 N41b [=Cleta Claus 1863 (preoc.) N41b; Laophontopsidae S03]
    |    |--*L. lamellifera (Claus 1863) [=*Cleta lamellifera] N41b
    |    `--L. secunda (Sewell 1924) N41b
    |--Parastenhelia Thompson & Scott 1903 N41b [incl. Microthalestris N41b; Parastenheliidae S03]
    |    |--‘Microthalestris’ forficula B26
    |    |--P. hornelli Thompson & Scott 1903 W67
    |    `--P. spinosa (Fischer 1860) W67
    |--Rhynchothalestris [Rhynchothalestridae] S03
    |    |--R. helgolandica (Claus 1863) S03
    |    |--R. rufocincta N44
    |    `--R. vanhoeffeni Brady 1910 N44
    |--Laophontodes Scott 1894 N41b [Ancorabolidae S03]
    |    |--L. armatus (Lang 1936) W67 [incl. Laophonte echinata Willey 1930 N41b]
    |    |--L. bicornis Scott 1896 N44
    |    |--L. brevis Nicholls 1944 N44
    |    `--L. typicus N44
    |--Huntemanniidae S03
    |    |--Huntemannia Poppe 1884 S03
    |    `--Nannopus S03
    |         |--N. abyssi Sars 1921 S03
    |         `--N. palustris Brady 1880 S03
    |--Dactylopusia N41b [Dactylopusiidae S03]
    |    |--D. micronyx N41b
    |    |--D. oculata Gurney 1927 N41b
    |    |--D. platysoma Thompson & Scott 1903 N41a
    |    |--D. tisboides N41b
    |    `--D. vulgaris N44
    |--Delamarella Chappuis 1954 W86 [Latiremidae S03]
    |    |--D. arenicola Chappuis 1954 W86
    |    |--D. eximia (Božić 1969) W86
    |    |--D. galateae Cottarelli 1971 W86
    |    |--D. karamani Petkovski 1957 W86
    |    `--D. phyllosetosa Kunz 1984 W86
    |--Phyllognathopodidae S03
    |    |--Allophyllognathopus Kiefer 1967 R86
    |    |    `--A. brasiliensis Kiefer 1967 R86
    |    `--Phyllognathopus Mrazek 1893 R86
    |         |--P. bassoti Rouch 1972 R86
    |         `--P. viguieri (Maupas 1892) B01
    |--Leptastacidae [Leptastacinae] S03
    |    |--Leptastacus W86
    |    |--Paraleptastacus W86
    |    |--Psammastacus W86
    |    `--Arenocaris Nicholls 1935 W86
    |         `--A. bifida Nicholls 1935 W86
    |--Orthopsyllus Brady & Robertson 1873 N41a [Orthopsyllidae S03]
    |    |--O. linearis (Claus 1866) W67
    |    |--O. littoralis Nicholls 1942 N42b
    |    |--O. major Klie 1939 N41a
    |    |--O. propinquus Monard 1926 N42b
    |    |--O. rugosus Nicholls 1941 N42b
    |    |--O. similis Nicholls 1942 N42b
    |    `--O. wallini Lang 1934 N42b
    |--Metidae S03
    |    |--Laubieria W67
    |    |    |--*L. corallicola Soyer 1966 W67
    |    |    `--L. secunda Wells 1967 W67
    |    `--Metis Philippi 1843 N41a
    |         |--M. holothuriae (Edwards 1891) W67 [=Abacola holothuriae B26]
    |         |--M. ignea (see below for synonymy) B26
    |         `--M. jousseaumei (Richard 1892) [=Ilyopsyllus jousseaumei] N42b
    |--Darcythompsoniidae S03
    |    |--Kristensenia Por 1983 S03
    |    |--Leptocaris S03
    |    |--Darcythompsonia W86
    |    `--Horsiella N44
    |         |--H. brevicornis N44
    |         |--H. gurneyi Nicholls 1944 N44
    |         `--H. trisetosa Kunz 1935 N44
    |--Miraciidae KM13
    |    |  i. s.: Eoschizopera Wells & Rao 1976 KM13
    |    `--Amphiascinae N41b
    |         |--Amphiascopsis N41b
    |         |--Amphiascus N41b
    |         |--Mesamphiascus N41b
    |         |--Amphiascoides N41b
    |         |--Robertsonia N41b
    |         `--Schizopera KM13
    |--Leptopontiidae [Leptopontiinae] S03
    |    |  i. s.: Arenopontia W86
    |    |         Notopontia Bodiou 1977 W86
    |    |           |--N. galapagoensis Mielke 1982 W86
    |    |           `--N. stephanieae Bodiou 1977 W86
    |    |         Ichnusella Cottarelli 1971 GL04
    |    |           |--I. eione Cottarelli 1971 R86
    |    |           `--I. pasquinii (Cottarelli 1969) R86
    |    |         Parasewellina Cottarelli, Saporito & Puccetti 1986 GL04
    |    |--Psammopsyllinae GL04
    |    `--Leptopontia Scott 1902 [Leptopontiinae] S03
    |         |--L. curvicauda Scott 1902 W86
    |         `--L. punctata Huys & Conroy-Dalton 1996 S03
    |--Cylindropsyllidae [Cylindropsyllinae] S03
    |    |--Stenocaris W86
    |    |--Boreopontia Willems 1981 W86
    |    |    `--B. heipi Willems 1981 W86
    |    |--Cylindropsyllus Brady 1880 W86
    |    |    |--C. laevis Brady 1880 W86
    |    |    `--C. remanei Kunz 1949 W86
    |    |--Syrticola Willems & Claeys 1982 W86
    |    |    |--S. flandricus Willems & Claeys 1982 W86
    |    |    `--S. galapagoensis (Mielke 1982) W86
    |    `--Evansula Scott 1906 W86
    |         |--E. arenicola Nicholls 1939 W86
    |         |--E. incerta (Scott 1892) W86
    |         `--E. pygmaea (Scott 1903) W86
    |--Rhizothrix Brady & Robertson 1875 W86 [Rhizothricidae S03]
    |    |  i. s.: R. curvata Brady & Robertson 1880 W86
    |    |         R. gracilis (Scott 1903) W86
    |    |         R. minuta (Scott 1903) W86
    |    |         R. reducta Noodt 1952 W86
    |    |           |--R. r. reducta W86
    |    |           `--R. r. noodti Galhano 1970 W86
    |    |         R. scotti Lang 1936 W86
    |    |         R. spinosa Coull 1971 W86
    |    |         R. tenella (Wilson 1932) W86
    |    |         R. wilsoni Bodin 1977 W86
    |    `--R. (Rhizothrix) W67
    |         |--R. (R.) pubescens Por 1959 W67
    |         `--R. (R.) quadriseta Wells 1967 W67
    |--Tetragonicipitidae [Pteropsyllidae, Tetragonicepsidae] S03
    |    |--Phyllopodopsyllus PH13
    |    |--Diagoniceps N44
    |    |--Laophontella Thompson & Scott 1903 W67, N41b
    |    |    `--L. armata (Willey 1935) [=Phyllopodopsyllus armatus] N44
    |    |--Tetragoniceps W67
    |    |    |--T. bergensis W67
    |    |    |--T. brownei Wells 1967 W67
    |    |    `--T. scotti W67
    |    `--Pteropsyllus W67
    |         |--P. consimilis (Scott 1894) W67
    |         `--P. plebius Monard 1935 W67
    |              |--P. p. plebius W67
    |              `--P. p. furcatus Kunz 1938 W67
    |--Parastenocarididae [Fontinalicaridinae, Parastenocaridinae] S03
    |    |--Parastenocaris CMA05
    |    |--Forficatocaris CMA05
    |    |--Remaneicaris CMA05
    |    |--Cafferocaris Jakobi 1972 GL04
    |    |--Paraforficatocaris Jakobi 1972 R86
    |    |    `--P. paranaensis Jakobi 1972 R86
    |    |--Murunducaris Reid 1994 CMA05
    |    |    `--M. juneae Reid 1994 CMA05
    |    |--Potamocaris Dussart 1970 CMA05
    |    |    |--P. bidens (Noodt 1955) GL04
    |    |    |--P. bifida Dussart 1979 R86
    |    |    `--P. estevesi Reid 1991 GL04
    |    `--Simplicaris Galassi & De Laurentiis 2004 CMA05
    |         |--*S. lethaea Galassi & De Laurentiis 2004 FS14
    |         |--‘Parastenocaris’ aedes Hertzog 1938 [=Pannonicaris aedes] GL04
    |         |--‘Parastenocaris’ hippuris Hertzog 1938 GL04
    |         `--S. veneris (Cottarelli & Maiolini 1980) [=Parastenocaris veneris] GL04
    |--Cletodidae [Cletodoidea] S03
    |    |--Cletodes N41b
    |    |--Parepactophanes Kunz 1935 W86
    |    |    |--P. dubia Noodt 1958 W86
    |    |    `--P. minuta Kunz 1935 W86
    |    |--Cletocamptus Schmankevitch 1875 R86
    |    |    |--C. albuquerquensis G88
    |    |    |--C. deitersi (Richard 1897) S-MR03
    |    |    `--C. xenuus Por 1968 R86
    |    |--Tryphoema Monard 1926 W86
    |    |    |--T. bocqueti (Božić 1954) W86
    |    |    |--T. lusitanica (Wells & Clark 1965) W86
    |    |    |--T. porca Monard 1926 W86
    |    |    |--T. ramabula (Pennak 1942) W86
    |    |    |--T. riedli Coull 1971 W86
    |    |    `--T. scilloniensis (Wells 1968) W86
    |    |--Paranannopus W56
    |    `--Enhydrosoma W86
    |         |--E. barnishi Wells 1967 W67
    |         |--E. buchholtzi (Boeck 1872) W67
    |         |--E. curticaudatum N41b
    |         |--E. curvirostre W67
    |         |--E. latipes (Scott 1909) W67
    |         |--E. littorale Wells 1967 W67
    |         |--E. longifurcatum Sars 1909 W67
    |         `--E. parapropinquum Gomez 2003 S03
    |--Harpacticidae S03
    |    |--Harpacticella SS03
    |    |    |--H. inopinata B50
    |    |    |--H. lacustris B50
    |    |    `--H. paradoxa B50
    |    |--Perissocope litoralis Lang 1934 S03
    |    |--Zausopsis mirabilis Lang 1934 S03
    |    |--Zausodes S03
    |    |    |--Z. septimus Lang 1965 S03
    |    |    `--Z. sextus Lang 1965 S03
    |    |--Tigriopus SS03
    |    |    |--T. brevicornis CCG01
    |    |    |--T. californicus SC17
    |    |    |--T. fulvus W67 [=Harpacticus fulvus PP64]
    |    |    `--T. japonicus WS10
    |    |--Harpacticus S03
    |    |    |--H. chelifer (Müller 1776) W67
    |    |    |--H. compsonyx Monard 1926 W67
    |    |    |--H. gracilis Claus 1863 W67
    |    |    |--H. littoralis B26
    |    |    |--H. trisetosus Lang 1948 S03
    |    |    `--H. uniremis B26
    |    `--Zaus Goodsir 1845 S03, N42a
    |         |--Z. abbreviatus Sars 1911 N42a
    |         |--Z. aurelii Poppe 1884 [incl. Z. caeruleus Campbell 1929, Z. intermedius Nicholls 1939] N42a
    |         |--Z. contractus Thomson 1883 N42a
    |         |--Z. goodsiri Brady 1880 N42a
    |         |--Z. sarsi Nicholls 1942 N42a
    |         `--Z. spinatus Goodsir 1845 N42a
    |--Thalestridae S03
    |    |  i. s.: Amenophia Boeck 1865 S03
    |    |         Paradactylopodia brevicornis (Claus 1866) W67
    |    |         Diarthrodes W67
    |    |           |--D. dissimilis W67
    |    |           |--D. glaber Wells 1967 W67
    |    |           |--D. minutus (Claus 1863) W67
    |    |           |--D. nanus (Scott 1914) W67
    |    |           `--D. sarsi (Scott 1909) W67
    |    |         Idomene [incl. Xouthous] W67
    |    |           |--I. aberrans Por 1964 W67
    |    |           |--I. aemula Thompson & Scott 1903 W67
    |    |           |--I. laticaudata (Thompson & Scott 1903) W67
    |    |           |--‘Xouthous’ maldivae Sewell 1940 W67
    |    |           `--I. simulans (Brady 1910) W67
    |    |         Dactylopodella flava N41b
    |    |--Dactylopodiinae N41a
    |    |    |--Dactylopodia tisboides (Claus 1863) W67
    |    |    `--Eudactylopus Scott 1909 N41a
    |    |         |--E. australis Nicholls 1941 N42b
    |    |         |--E. latipes (Scott 1894) N41a
    |    |         |--E. robustus (Claus 1863) W67
    |    |         `--E. spectabilis (Brian 1923) N41a
    |    `--Thalestrinae N41a
    |         |--Thalestris longimana BK77
    |         `--Phyllothalestris Sars 1905 N41a
    |              |--P. lata Nicholls 1942 N42b
    |              |--P. mysis (Claus 1863) N41a
    |              |    |--P. m. mysis N42b
    |              |    `--P. m. harringtoni Willey 1935 N42b
    |              |--P. paramysis Monard 1928 N42b
    |              |--P. royi (Monard 1928) N42b
    |              `--P. sarsi Sewell 1940 W67
    |--Diosaccidae S03
    |    |  i. s.: Pseudamphiascopsis ismaelensis (Monard 1936) W67
    |    |         Bulbamphiascus imus (Brady 1872) W67
    |    |         Haloschizopera junodi (Monard 1935) W67
    |    |         Paramphiascella robinsoni (Scott 1902) W67
    |    |         Melima W67
    |    |         Helmutkunzia W86
    |    |         Typhlamphiascus Lang 1948 R86
    |    |           `--T. latifurcata Por 1968 R86
    |    |         Pseudostenhelia Wells 1967 W67
    |    |           `--*P. prima Wells 1967 W67
    |    |         Paramphiascoides Wells 1967 W67
    |    |           `--*P. mixtus Wells 1967 W67
    |    |         Actopsyllus Wells 1967 W67
    |    |           `--*A. longipes Wells 1967 W67
    |    |         Schizoperoides Por 1968 R86
    |    |           `--S. expeditionis Por 1968 R86
    |    |         Goffinella Wilson 1932 W86
    |    |           `--G. stylifer Wilson 1932 W86
    |    |         Balucopsylla Rao 1972 W86
    |    |           `--B. similis Rao 1972 W86
    |    |         Protopsammotopa Geddes 1968 W86
    |    |           |--P. norvegica Geddes 1968 W86
    |    |           `--P. wilsoni Wells 1977 W86
    |    |         Robertgurneya W67
    |    |           |--R. rostrata (Gurney 1927) W67
    |    |           |--R. similis (Scott 1896) W67
    |    |           `--R. spinulosa (Sars 1911) W67
    |    |         Psammotopa Pennak 1942 W86
    |    |           |--P. chappuisi Noodt 1955 W86
    |    |           |--P. phyllosetosa (Noodt 1952) W86
    |    |           |--P. polyphylla Noodt 1955 W86
    |    |           `--P. vulgaris Pennak 1942 W86
    |    |--Diosaccinae N41b
    |    `--Stenheliinae N41b
    |         |--Stenhelia N41b
    |         `--Pseudomesochra Scott 1902 [incl. Stenheliopsis Sars 1906] N41b
    |--Canthocamptidae S03
    |    |--Nannomesochra arupinensis (Brian 1925) W67
    |    |--Bryocamptus S-MR03
    |    |--Attheyella S-MR03
    |    |--Elaphoidella KM13
    |    |--Moraria S-MR03
    |    |--Australocamptus hamondi Karanovic 2004 KM13
    |    |--Gulcamptus Miura 1969 R86
    |    |    `--G. uenoi Miura 1969 R86
    |    |--Antarctobiotus Chappuis 1930 R86
    |    |    `--A. bahamondei Ebert & Noodt 1975 R86
    |    |--Epactophanoides Borutzky 1966 R86
    |    |    `--E. udegeicus Borutzky 1966 R86
    |    |--Stygepactophanes Moeschler & Rouch 1984 R86
    |    |    `--*S. jurassicus Moeschler & Rouch 1984 R86
    |    |--Echinocamptus Chappuis 1928 R86
    |    |    |--E. echinatus (Mrázek 1893) [=Bryocamptus (Echinocamptus) echinatus] FS14
    |    |    `--E. georgevitchi (Chappuis 1924) B01
    |    |--Paracamptus Chappuis 1929 R86
    |    |    |--P. baikalensis W56
    |    |    |--P. gasparoi Stoch 1998 B01
    |    |    |--P. nakamurai Chappuis 1958 R86
    |    |    |--P. reductus Wilson 1956 W56
    |    |    `--P. schmeili W56
    |    |--Morariopsis Borutzky 1931 R86
    |    |    |--M. kieferi Petkovski 1959 R86
    |    |    `--M. scotenophila (Kiefer 1930) B01
    |    |--Canthocamptus Westwood 1836 R86
    |    |    |--C. insoletus Chappuis 1928 R86
    |    |    |--C. morimotoi Miura 1969 R86
    |    |    `--C. staphylinus D01
    |    |--Spelaeocamptus Chappuis 1933 R86
    |    |    |--S. incertus Petkovski 1956 R86
    |    |    |--S. neotropicus Noodt 1963 R86
    |    |    `--S. spelaeus (Chappuis 1925) R86
    |    |--Mesochra Boeck 1865 R86, S03
    |    |    |--M. hiunumaensis Kikuchi 1972 R86
    |    |    |--M. lilljeborgi [incl. Laophonte hecate Brehm 1910] N41b
    |    |    |--M. pygmaea (Claus 1863) W67
    |    |    `--M. rostrata (Gurney 1927) W67
    |    |--Antrocamptus Chappuis 1956 R86
    |    |    |--A. catherinae Chappuis & Rouch 1960 R86
    |    |    |--A. chappuisi Rouch 1970 R86
    |    |    |--A. coiffaiti Chappius 1956 R86
    |    |    |--A. longifurcatus Rouch 1970 R86
    |    |    `--A. stygius Rouch 1970 R86
    |    |--Maraenobiotus Mrazek 1893 R86
    |    |    |--M. brucei R86
    |    |    |    |--M. b. brucei R86
    |    |    |    `--M. b. carpathicus Chappuis 1928 R86
    |    |    `--M. insignipes R86
    |    |         |--M. i. insignipes R86
    |    |         `--M. i. kyzylkumicus Borutzky 1972 R86
    |    `--Ceuthonectes Chappuis 1924 R86
    |         |--C. chappuisi Rouch 1980 R86
    |         |--C. gallicus Chappuis 1928 R86
    |         |--C. hungaricus Ponyi 1958 R86
    |         |--C. mirabilis Miura 1964 R86
    |         |--C. rouchi Petkovski 1984 R86
    |         |--C. serbicus Chappuis 1924 [incl. Morariodes colchidana Borutzky 1930, C. colchidanus] R86
    |         `--C. vievilleae Rouch 1980 R86
    |--Ameiridae S03
    |    |  i. s.: ‘Amphiascus’ mucronatus Brady 1910 N41b
    |    |         ‘Amphiascus’ proximus Scott 1914 N41b
    |    |--Psammoleptomesochra australis (Mielke 1994) [=Paramesochra australis] S03
    |    |--Pseudameira W67
    |    |--Proameira W67
    |    |--Nitocrella R86
    |    |--Leptomesochra W67
    |    |--Parapseudoleptomesochra S-MR03
    |    |--Nitokra PH13
    |    |--Stygonitocrella R86
    |    |--Sicamaeira W86
    |    |--Inermipes humphreysi Lee & Huys 2002 HA13
    |    |--Biameiropsis barrowensis Karanovic 2006 HA13
    |    |--Pseudoleptomesochra Lang 1965 W86
    |    |    `--P. typica Lang 1965 W86
    |    |--Psyllocamptus Scott 1899 R86
    |    |    |--P. minutus Sars 1911 W67
    |    |    `--P. monachus Chappuis 1938 R86
    |    |--Paraleptomesochra Wells 1967 W86
    |    |    |--*P. minima Wells 1967 W67
    |    |    `--P. wellsi Rao 1972 W86
    |    |--Philoleptomesochra Wells 1967 W67
    |    |    |--*P. reductiforma Wells 1967 W67
    |    |    `--P. secunda Wells 1967 W67
    |    |--Karllangia Noodt 1964 W86
    |    |    |--*K. arenicola Noodt 1964 W67, W86
    |    |    |--K. psammophila Wells 1967 W86
    |    |    `--K. tertia Kunz 1975 W86
    |    |--Sarsameira Wilson 1924 [incl. Pseudosarsameira] W67
    |    |    |--S. exilis (Scott & Scott 1894) W67
    |    |    |--S. minor Wells 1967 W67
    |    |    `--S. pendula (Shen & Bai 1956) W67
    |    |--Psammameira Noodt 1952 W67
    |    |    |--*P. hyalina Noodt 1952 W67
    |    |    |--P. grandis (Nicholls 1939) [=Ameira grandis] W67
    |    |    `--P. reducta Wells 1967 W67
    |    |--Nitocrellopsis Petkovski 1976 R86
    |    |    |--N. elegans (Chappuis & Rouch 1959) R86
    |    |    |--N. intermedia (Chappuis 1937) R86
    |    |    `--N. ioneli (Dumont & Decraemer 1974) R86
    |    |--Praeleptomesochra Lang 1965 W86
    |    |    |--P. africana (Kunz 1951) W86
    |    |    |--P. phreatica Pesce 1981 R86
    |    |    |--P. pygmaea (Vervoort 1964) W86
    |    |    `--P. similis Lang 1965 W86
    |    |--Ameira W86
    |    |    |--A. exigua W67
    |    |    |--A. leptoderma (Klie 1950) W67
    |    |    |--A. longipes Boeck 1864 W67
    |    |    |--A. parvula (Claus 1866) [=Amphiascus parvulus] N41b
    |    |    `--A. simulans W67
    |    |--Pseudoleptomesochrella Lang 1965 W86
    |    |    |--P. bisetosa Lindgren 1975 W86
    |    |    |--P. halophila (Noodt 1952) W86
    |    |    |--P. incerta (Chappuis & Delamare Deboutteville 1956) W86
    |    |    |--P. marina (Chappuis & Rouch 1960) W86
    |    |    `--P. pontica Apostolov 1969 W86
    |    |--Parevansula Guille & Soyer 1966 W86
    |    |    |--P. elegans (Marinov 1974) W86
    |    |    |--P. mediterranea Guille & Soyer 1966 W86
    |    |    |--P. reductiforma (Wells 1967) W86
    |    |    |--P. secunda (Wells 1967) W86
    |    |    |--P. vermiformis Moore 1976 W86
    |    |    `--P. wellsi (Marinov 1973) W86
    |    `--Interleptomesochra Lang 1965 W86
    |         |--I. attenuata (Scott 1896) W86
    |         |--I. boguensis Lindgren 1975 W86
    |         |--I. elongata (Božić 1955) W86
    |         |--I. eulitoralis (Noodt 1952) W86
    |         |--I. noodti Galhano 1968 W86
    |         |--I. reducta Lang 1965 W86
    |         `--I. tenuicornis (Sars 1911) W86
    `--Laophontidae [Namakosiramiidae] S03
         |--Heterolaophonte quinquespinosa (Sewell 1924) W67
         |--Klieonychocamptus W67
         |--Pseudocletopsyllus Vervoort 1964 W67
         |--Mictyricola Nicholls 1957 W67
         |--Pholenota Vervoort 1964 W67
         |--Novolaophonte viatorum Cottarelli, Saporito & Puccetti 1983 W86
         |--Laophonte N41b
         |--Platychelipus Brady 1880 N41b
         |--Sarsocletodes Wilson 1924 [=Pseudocletodes Sars 1921 (preoc.), Pseudoplatychelipus Lang 1936] N41b
         |--Donsiella Stephenson 1935 N41b
         |--Psammolaophonte Wells 1967 W86
         |    `--*P. spinicauda Wells 1967 W67
         |--Apolethon Wells 1967 W67
         |    `--*A. fumator Wells 1967 W67
         |--Esola Edwards 1891 N41b
         |    `--E. spelea (Chappuis 1938) R86
         |--Cubanocleta Petkovski 1977 R86
         |    `--C. noodti Petkovski 1977 R86
         |--Mexicolaophonte Cottarelli 1977 W86
         |    `--M. arganoi Cottarelli 1977 W86
         |--Arenolaophonte Lang 1965 W86
         |    `--A. stygia Lang 1965 W86
         |--Stygolaophonte Lang 1965 W86
         |    `--S. arenophila Lang 1965 W86
         |--Pseudolaophonte Scott 1896 N41b
         |    `--P. spinosa (Thompson 1893) [incl. P. aculeata Scott 1896] N41b
         |--Harrietella Scott 1906 N41b
         |    `--H. simulans (Scott 1894) N41b
         |--Hemilaophonte Jakubisiak 1932 N41b
         |    `--H. janinae Jakubisiak 1932 N41b
         |--Lobitella Monard 1934 N41b
         |    `--L. apoda Monard 1934 N41b
         |--Onychocamptus PH13
         |    |--O. bengalensis (Sewell 1934) PH13
         |    `--O. mohammed (Blanchard & Richard 1891) S-MR03 (see below for synonymy)
         |--Tapholeon Wells 1967 W67
         |    |--*T. ornatus Wells 1967 W67
         |    `--T. uniarticulatus Wells 1967 W67
         |--Paralaophonte W67
         |    |--P. brevirostris (Claus 1863) W67
         |    |--P. congenera (Sars 1908) W67
         |    |--P. quinquespinosa P71
         |    `--P. tenera (Sars 1920) W67
         |--Asellopsis Brady & Robertson 1873 N41b
         |    |--A. duboscqui Monard 1926 N41b
         |    |--A. hispida Brady & Robertson 1873 N41b
         |    |--A. intermedia (Scott 1895) N41b
         |    `--A. littoralis Nicholls 1939 N41b
         |--Klieonychocamptoides Noodt 1958 W86
         |    |--K. arenicola (Chappuis & Delamare Deboutteville 1956) W86
         |    |--K. arganoi Cottarelli & Mura 1980 W86
         |    |--K. itoi Mielke 1981 W86
         |    `--K. remanei Noodt 1958 W86
         |--Echinolaophonte Nicholls 1941 N41b
         |    |--*E. horrida (Norman 1876) [=Laophonte horrida] N41b
         |    |--E. armiger (Gurney 1927) (see below for synonymy) N41b
         |    |--E. brevispinosa (Sars 1908) [=Laophonte brevispinosa] N41b
         |    `--E. mirabilis (Gurney 1927) [=Laophonte mirabilis] N41b
         |--Laophontina Norman & Scott 1905 N41b
         |    |--L. acantha Noodt 1955 W86
         |    |--L. distincta Wells 1967 W86
         |    |--L. dubia Norman & Scott 1905 N41b
         |    |--L. noodti Kunz 1983 W86
         |    |--L. reducta Coull & Zo 1980 W86
         |    |--L. triarticulata Coull & Zo 1980 W86
         |    `--L. variabilis Coull & Zo 1980 W86
         `--Aphrolaophonte Chappuis 1960 W86
              |--A. aequatorialis Cottarelli & Mura 1982 W86
              |--A. brevipes (Chappuis 1954) W86
              |--A. leonis Cottarelli & Mura 1982 W86
              |--A. michaelae Cottarelli & Mura 1982 W86
              |--A. monodi Chappuis 1960 W86
              |--A. pori Masry 1970 W86
              |--A. renaudi (Chappuis & Delamare Deboutteville 1956) W86
              `--A. schmidti Mielke 1981 W86

Podogennonta incertae sedis:
  Antiboreodiosaccus Lang 1944 S03
  Dactylopina Brady 1910 S03
  Flavia Brady 1899 S03
  Mawsonella Brady 1918 S03
  Pyrocletodes Coull 1973 S03
  Tisemus Monard 1928 S03
  Metahuntemannia Smirnov 1946 S03
    |--M. spinipes Dahms & Pottek 1992 S03
    `--M. triarticulata Schriever 1984 S03
  Schizacron vervoorti (Fiers 1987) S03
  Psammonitocrella Rouch 1992 GL04

Echinolaophonte armiger (Gurney 1927) [=Laophonte armiger; incl. L. hystrix Brian 1928, L. steueri van Douwe 1929] N41b

Lourinia Wilson 1924 N41a [=Jurinia Claus 1866 (preoc.) N41a; incl. Ceylonia Thompson & Scott 1903 N41a, Ceyloniella Wilson 1924 N41a; Ceyloniellidae, Louriniidae S03]

Lourinia armata (Claus 1866) W67 [=Jurina armata N41a, Ceylonia armata N41a, Ceyloniella armata N41a; incl. Ceylonia aculeata Thompson & Scott 1903 N41a, Ceyloniella aculeata N41a, Ceylonia aculeata var. adriatica Brian 1923 N41a, Ceyloniella aculeata var. adriatica N41a]

Metis ignea [incl. Ilyopsyllus coriaceus Brady & Robertston 1873, I. natans Williams 1907, I. sarsi Sharpe 1911] B26

Onychocamptus mohammed (Blanchard & Richard 1891) S-MR03 [=Laophonte mohammed S-MR03; incl. L. humilis Brian 1929 N41b]

*Type species of generic name indicated

REFERENCES

[BK77] Barel, C. D. N., & P. G. N. Kramers. 1977. A survey of the echinoderm associates of the north-east Atlantic area. Zoologische Verhandelingen 156: 1–159.

[B26] Bigelow, H. B. 1926. Plankton of the offshore waters of the Gulf of Maine. Bulletin of the Bureau of Fisheries 40 (2): 1–509.

[B01] Brancelj, A. 2001. Male of Moraria radovnae Brancelj, 1988 (Copepoda: Crustacea), and notes on endemic and rare copepod species from Slovenia and neighbouring countries. Hydrobiologia 453/454: 513–524.

[B50] Brooks, J. L. 1950. Speciation in ancient lakes. Quarterly Review of Biology 25: 30–60.

[CCG01] Canivet, V., P. Chambon & J. Gibert. 2001. Toxicity and bioaccumulation of arsenic and chromium in epigean and hypogean freshwater macroinvertebrates. Archives of Environmental Contamination and Toxicology 40: 345–354.

[CMA05] Corgosinho, P. H. C., & P. Martínez Arbizu. 2005. Two new interstitial species of Remaneicaris Jakobi 1972 from the Ribeirão do Ouro River, Minas Gerais, Brazil, with a redefinition of the genus (Crustacea, Copepoda, Harpacticoida, Parastenocarididae). Senckenbergiana Biologica 85 (2): 147–162.

[D01] Daday, E. 1901. Édesvizi mikroszkópi állatok [Mikroskopische Süsswasserthiere]. In: Horváth, G. (ed.) Zichy Jenő Gróf Harmadik Ázsiai Utazása [Dritte Asiatische Forschungsreise des Grafen Eugen Zichy] vol. 2. Zichy Jenő Gróf Harmadik Ázsiai Utazásának Állattani Eredményei [Zoologische Ergebnisse der Dritten Asiatischen Forschungsreise des Grafen Eugen Zichy] pp. 375–470. Victor Hornyánszky: Budapest, and Karl W. Hierseman: Leipzig.

[FS14] Fiasca, B., F. Stoch, M.-J. Olivier, C. Maazouzi, M. Petitta, A. Di Cioccio & D. M. P. Galassi. 2014. The dark side of springs: what drives small-scale spatial patterns of subsurface meiofaunal assemblages? J. Limnol. 73 (1): 71–80.

[GL04] Galassi, D. M. P., & P. De Laurentiis. 2004. Towards a revision of the genus Parastenocaris Kessler, 1913: establishment of Simplicaris gen. nov. from groundwaters in central Italy and review of the P. brevipes-group (Copepoda, Harpacticoida, Parastenocarididae). Zoological Journal of the Linnean Society 140: 417–436.

[G88] Gray, J. 1988. Evolution of the freshwater ecosystem: the fossil record. Palaeogeography, Palaeoclimatology, Palaeoecology 62: 1–214.

[HA13] Humphreys, G., J. Alexander, M. S. Harvey & W. F. Humphreys. 2013. The subterranean fauna of Barrow Island, north-western Australia: 10 years on. Records of the Western Australian Museum Supplement 83: 145–158.

[KM13] Karanovic, T. & J. McRae. 2013. The genus Schizopera (Copepoda, Harpacticoida) in the Pilbara region of Western Australia, with description of a new species and its molecular and morphological affinities. Records of the Western Australian Museum 28 (2): 119–140.

[N41a] Nicholls, A. G. 1941a. Littoral Copepoda from South Australia. (I) Harpacticoida. Records of the South Australian Museum 6 (4): 381–427.

[N41b] Nicholls, A. G. 1941b. A revision of the families Diosaccidae Sars, 1906 and Laophontidae T. Scott, 1905 (Copepoda, Harpacticoida). Records of the South Australian Museum 7 (1): 65–110.

[N42a] Nicholls, A. G. 1942a. A review of the genus Zaus Goodsir, and a description of two species of Laophonte Philippi (Copepoda, Harpacticoida). Annals and Magazine of Natural History, series 11, 9: 119–127.

[N42b] Nicholls, A. G. 1942b. Marine Copepoda from Western Australia. I. Littoral harpacticoids from Rottnest Island. Journal of the Royal Society of Western Australia 27: 135–141.

[N44] Nicholls, A. G. 1944. Littoral Copepoda from the Red Sea. Annals and Magazine of Natural History, series 11, 11: 487–503.

[PP64] Peres, J. M., & J. Picard. 1964. Nouveau manuel de bionomie benthique de la mer Mediterranee. Recueil des Travaux de la Station Marine d'Endoume, Bulletin 31 (27): 5–137.

[PH13] Poore, G. C. B., & W. F. Humphreys. 2013. Bunderanthura bundera gen. et sp. nov. from Western Australia, first anchialine Leptanthuridae (Isopoda) from the Southern Hemisphere. Records of the Western Australian Museum 28 (1): 21–29.

[P71] Por, F. D. 1971. One hundred years of Suez Canal—a century of Lessepsian migration: retrospect and viewpoints. Systematic Zoology 20 (2): 138–159.

[R86] Rouch, R. 1986. Copepoda: Les harpacticoïdes souterrains des eaux douces continentales. In: Botosaneanu, L. (ed.) Stygofauna Mundi: A Faunistic, Distributional, and Ecological Synthesis of the World Fauna inhabiting Subterranean Waters (including the Marine Interstitial) pp. 321–355. E. J. Brill/Dr W. Backhuys: Leiden.

[SC17] Schwentner, M., D. J. Combosch, J. P. Nelson & G. Giribet. 2017. A phylogenomic solution to the origin of insects by resolving crustacean-hexapod relationships. Current Biology 27: 1818–1824.

[S03] Seifried, S. 2003. Phylogeny of Harpacticoida (Copepoda): Revision of “Maxillipedasphalea” and Exanechentera. Cuvillier Verlag: Göttingen.

[SS03] Seifried, S., & H. K. Schminke. 2003. Phylogenetic relationships at the base of Oligoarthra (Copepoda, Harpacticoida) with a new species as the cornerstone. Organisms Diversity & Evolution 3: 13–37.

[S-MR03] Suárez-Morales, E., & J. W. Reid. 2003. An updated checklist of the continental copepod fauna of the Yucatan Peninsula, Mexico, with notes on its regional associations. Crustaceana 76 (8): 977–991.

[WS10] Wang, M., S. Sun, C. Li & X. Shen. 2010. Distinctive mitochondrial genome of the calanoid copepod Calanus sinicus, a useful molecular marker for phylogenetic and population studies. In: China-Russia Bilateral Symposium: Proceedings of the China-Russia Bilateral Symposium of "Comparison on Marine Biodiversity in the Northwest Pacific Ocean", 10–11 October 2010, Qingdao (China) pp. 187–192. Institute of Oceanology, Chinese Academy of Sciences; A. V. Zhirmunsky Institute of Marine Biology, Far East Branch of the Russian Academy of Sciences.

[W81] Watson, L. 1981. Sea Guide to Whales of the World. Hutchinson: London.

[W67] Wells, J. B. J. 1967. The littoral Copepoda (Crustacea) of Inhaca Island, Mozambique. Transactions of the Royal Society of Edinburgh 67: 189–358.

[W86] Wells, J. B. J. 1986. Copepoda: Marine-interstitial Harpacticoida. In: Botosaneanu, L. (ed.) Stygofauna Mundi: A Faunistic, Distributional, and Ecological Synthesis of the World Fauna inhabiting Subterranean Waters (including the Marine Interstitial) pp. 356–381. E. J. Brill/Dr W. Backhuys: Leiden.

[W56] Wilson, M. S. 1956. North American harpacticoid copepods: 3, Paracamptus reductus, n. sp., from Alaska. Journal of the Washington Academy of Sciences 46 (11): 348–351.

Last updated: 6 March 2022.