Calanoida ( Order )
    Epacteriscoidea ( Superfamily )
Epacteriscidae Fosshagen,1973; emend.: Fosshagen, Boxshall & Iliffe, 2001 ( Epacteriscoidea )
Ref.: Fosshagen, 1973 (p.146); Razouls, 1982 (p.595); Bowman & Abele, 1982 (p.9); Barr, 1984 (p.160, Rev., emend.); Fosshagen & Iliffe, 1985 (p.346, Rev.); Park, 1986 (p.192, Rem.); Huys & Boxshall, 1991 (p.406, 419); Razouls, 1993 (p.306); Bradford-Grieve, 1999 b (p.25, Def.); Jaume & Humphreys, 2001 (p.157, Genera key, Rem.); Ohtsuka & Huys, 2001 (p.461); Fosshagen & al., 2001 (p.246, Rem., Genera Key, Phylogeny); Boxshall & Halsey, 2004 (p.14; 49; 111: Def.; p. 112: key of genera); Vives & Shmeleva, 2007 (p.992); Fosshagen & Iliffe, 2007 (p.73, Rem.: p.89, Genera Key: p.91); Blanco-Bercial & al., 2011 (p.103, Fig.3, molecular biology, phylogeny); Bradford-Grieve & al., 2014 (p.507, 528: Rem. Table 1, fig.11, cladistic analysis)
Bradford-Grieve J.M., (2002 onwards). Key to calanoid copepod families. Version 1 : 2 oct 2002. http://www.crustacea.net/crustace/calanoida/index.htm
Rem.: For Fosshagen (1973), the position of this family is difficult to situate. The author includes the family initially in the superfamily Centropagina (Gurney,1931) but underlines certain characteristics that make them come closer to the Arietellidae, the Ridgewayiidae, or even the Augaptilidae. It appears to be difficult to make the partitioning between anatomical convergences due to life strategies and phylogenetic relationships.
Hyperbenthic forms and submarine caves.
Type genus: Epacteriscus Fosshagen, 1973.
19 G.: Azygonectes, Balinella, Bofuriella, Bomburiella, Bunderia, Caiconectes, Cryptonectes, Edaxiella, Enantiosis, Enantronia, Enantronoides, Epacteriscus, Erebonectes, Erebonectoides, Gloinella, Iboyella, Miheptneria, Minnonectes, Oinella.
For Bradford-Grieve & al. (2014, p.528) the genera included in this family are: Balinella, Bofuriella, Bomburiella, Bunderia, Cryptonectes, Edaxiella, Enantiosis, Enantronia, Enantronoides, Epacteriscus, Erebonectes, Gloinella, Iboyella, Mikheptneria, Minnonectes, Oinella. Total 15 genera.

Diagnosis after Bradford-Grieve & al. (2014, p.528) :
Female caudal rami usually of equal lengths (except Balinella longer on right; Gloinella longer on ri!ght); male caudal rami symmetrical or asymmetrical, seta II spiniform, setiform, or apparently absent in Miheptneria, female seta VI tending to be asymmetrical, with 1 seta having one side bearing short setules, the other with long setules (except Balinella, Cryptonectes, Oinella, Bunderia, Edaxiella, Enantroides).
Female A1 ancestral segment IV with aesthetasc (except for Oinella).
Md with exopod and basis forming major axis of palp; endopod poorly developed , 2- or 1-segmented, or absent, being at most half length of exopod segment 1, which when present, with at most 2 setae, segment 2 with 4 or fewer setae (except Erebonectes, Miheptneria, which have 7 and 9 setae, respectively).
Mx1 coxal epipodite with seta 9 absent (except Miheptneria), and tendency for further setae to be lost up to seta 6.
Mx2 basis usually more than twice estimated length of coxa, basal endite with low profile, and endopod setae spine-like (except Miheptneria, which has elongate basal endite and normal setae).
Mxp endopod segments 2-6 shorter than length of coxa (not including praecoxa) and endopod setae spine-like (eccept Miheptneria, Bomburiella, Edaxiella, which appear to have normal setae on endopod).
P3 endopod segment 1 with outer distal corner bifid or trifid.
Male left P5 without inner process present on exopod segment 2 (only exception Balinella).

Key to genera after Boxshall & Halsey (2004, p.112) :
1 - P3 and 4 with 2 outer spines on 3rd exopodal segment ……. 2.
1’ – P3 and P4 with 3 outer spines on 3rd exopodal segment …… 3.
2 – P2 with strong outer spine on basis ……. Erebonectes.
2’ – P2 with unarmed basis ……. Erebonectoides.
3 – Gnathobase of Md projecting frontally, with tip modified into coarse rake-like blade ; labrum elongate, tapering to a single point …….. Epacteriscus.
3’ – Gnathobase of Md not projecting frontally, and not of this form ; labrum typically with rounded tip, not as above …….. 4.
4 – Female A1 with modified flattened setae with flagellate tips on proximal segments (I-V typically) …….. 5.
4’ – Female A1 with normal taperinf setae on proximal segments ……… 9.
5 – Syncoxal setal formula of Mxp 1,2, 4,4 ; rostrum variable in form ……. 6.
5’ – Syncoxal setal formula of Mxp 0,0,2,4 ; rostrum vestigial, represented by weakly rounded lobes on frontal margin …….. Bomburiella.
6 – Mx1 with 6 or 7 well developed setae on exopod ; proximal segments of male left P5 exopod strongly modified (where known) …….. 7.
6’ – Mx1 with 1 well developed and 2 vestigial setae on exopod ; proximal segments of male left P5 exopod unmodified ……… Bunderia.
7 – A2 with 1st endopodal segment about equal to or shorter than compound distal segment ; anal somite minute, completely concealed beneath posterior rim of preceding somite …… 8.
7’ – A2 with 1st endopodal segment distinctly longer than compound distal segment ; anal somite well developed and conspicuous …….. Enantronia.
8 – Md palp with single minute seta on basis and with endopod located distally on basis ; Mx1 with 3 or 4 setae on coxal and proximal basal endites ……. Iboyella.
8’ – Md palp with free endopodal segment located in middle of basis, and basis unarmed ; Mx1 with 2 and 1 setae on coxal and proximal basal endites ……. Enantiosis.
9 – Md palp uniramous ; lacking any trace of endopod and lacking any setae on basis ; rostrum scarcely defined, represented by paired filaments carried on anterior margin of dorsal cephalic shield …….. 10.
9’ – Md palp biramous, with 1 or 2-segmented endopod, or with endopod represented by 2 setae on small process ; rostrum typically well-developed, bifurcate or elongate with bifid tip, or (rarely) weakly developed ……….. 11.
10 – Md palp with 1 extremely long seta plus 5 normal setae on exopod ; female A1 with rounded process on segment IX ; A2 with short endopod, 1st segment markedly shorter than second …….. Oinella.
10’ – Md palp without unusually long seta on exopod ; female A1 without rounded process on segment IX ; A2 with long endopod, 1st segment markedly longer than second …… Enantronoides.
11 – A2 well developed, with endopod slightly longer than exopod ……… Balinella.
11’ – A2 small, with endopod markedly shorter than exopod ……… 12.
12 – Rostrum elongate, indented at tip with short rostral filaments ; female P5 lacking inner seta on 1st exopodal segment …….. Gloinella.
12’ – Rostrum not elongate ; female P5 with inner seta on 1st exopodal segment ……… 13.
13 – Rostrum bifid with pointed branches ; anal somite minute, completely concealed beneath posterior rim of preceding somite, giving urosome a 3-segmented appearance ……… Edaxiella.
13’ – Rostrum vestigial, represented by weakly rounded lobes on frontal margin ; anal somite well developed, urosome conspicuously 4-segmented …….. Bofuriella.
Family Epacteriscidae - Plate 1Issued from : J.M. Bradford-Grieve, G.A. Boxshall & L. Blanco-Bercial in Zool. J. Linn. Soc., 2014, 171. [p.525, Fig.11].
Strict consensus of one tree after one round of successive weighting. Clade numbers above the line. The out-group is Disseta palumbii Giesbrecht, 1889.

Family Epacteriscidae - Plate 2Issued from : J.M. Bradford-Grieve, G.A. Boxshall & L. Blanco-Bercial in Zool. J. Linn. Soc., 2014, 171. [p.528, Table 7].
Spine and seta formulae of swimming legs in Epacteriscidae.
Roman numerals = robust setae; Arabic numerals = setae. Outer border setation listed in first in each segmental group, and separated by ;

Family Epacteriscidae - Plate 3Issued from : G.A. Boxshall & S.H. Halsey in An Introduction to Copepod Diversity. The Ray Society, 2004, No 166, Part. I. [p.111].
Armature formula of swimming legs P1 to P4 and P5 female (f).

Nota: Female P5 comprising 3-segmented protopod retaining hoop-like vestige of praecoxa laterally, and 3-segmented rami; setation sometimes reduced.
- Male P5 asymmetrical; both legs typically with 3-segmented rami; exopod 2-segmented due to fusion of 2nd and 3rd segments fused on left side in Epacteriscus. Endopods similar to those of female with armature 0-1; 0-1; 2,2,2; slightly transformed in some genera. Exopods usually highly transformed; most complex structures usually present on left exopod; modified with a slender spiniform distal segment, long and curved on right side in Erebonectes and Enanttiosis.

Family Epacteriscidae - Plate 4Issued from : G.A. Boxshall & S.H. Halsey in An Introduction to Copepod Diversity. The Ray Society, 2004, No 166, Part. I. [p.113, Fig.21].
Epacteriscidae. A, Erebonectes nesioticus habitus male; B, A1; C, A2; D, Epacteriscus rapax habitus female; E, P5. [Fosshagen & Iliffe, 1985: A; Huys & Boxshall, 1991: B-C, E; Fosshagen, 1973: D]

Family Epacteriscidae - Plate 5Issued from : J.M. Bradford-Grieve in NIWA Biodiv. Memoir 111, 1999 [p.26]. Armature formula of the swimming legs P1 to P5.
(1) Azygonectes Fosshagen & Iliffe, 2004
Ref.: Fosshagen & Iliffe, 2004 (p.119); 2007 (p.80: emendation, 91); Bradford-Grieve & al., 2014 (p.507, 522: Rem., Table 1, fig. 9, 10, 11, cladistic analysis)
Rem.: Total: 2 spp.
Bradford-Grieve & al. (2014, p.530) consider the genus, after phylogenetic analysis, of uncertain placement in the family (incertae sedis).
(2) Balinella Fosshagen, Boxshall & Iliffe, 2001
Ref.: Fosshagen & al., 2001 (p.292, 316); Bradford-Grieve, 2004 (p.287); Fosshagen & Iliffe, 2007 (p.91); Bradford-Grieve & al., 2014 (p.507, 529: Rem., Table 1, fig. 9, 10, 11)
Rem.: Total: 1 sp.
(3) Bofuriella Fosshagen, Boxshall & Iliffe, 2001
Ref.: Fosshagen & al., 2001 (p.286, 316); Bradford-Grieve, 2004 (p.287); Fosshagen & Iliffe, 2007 (p.85: emendation, p.92)
Rem.: 3 spp.
(4) Bomburiella Fosshagen, Boxshall & Iliffe, 2001
Ref.: Fosshagen & al., 2001 (p.282, 316); Bradford-Grieve, 2004 (p.287); Fosshagen & Iliffe, 2007 (p.92); Bradford-Grieve & al., 2014 (p.507, 522: Rem., Table 1, fig. 9, 10, 11, cladistic analysis)
Rem.: Total: 1 sp.
(5) Bunderia Jaume & Humphreys, 2001
Ref.: Jaume & Humphreys (p.158, 168); Bradford-Grieve, 2004 (p.287); Fosshagen & Iliffe, 2007 (p.92); Bradford-Grieve & al., 2014 (p.507, 522: Rem., Table 1, fig. 9, 10, 11, cladistic analysis)
Rem.: Total: 1 sp.
(6) Caiconectes Fosshagen & Iliffe, 2007
Ref.: Fosshagen & Iliffe, 2007 (p.74, 91); Bradford-Grieve & al., 2014 (p.507, 530, Table 1, fig. 9, 10, 11).
Rem.: Total: 1 sp.
Bradford-Grieve & al. (2014, p.530) consider the genus, after phylogenetic analysis, of uncertain placement in the family (incertae sedis).
(7) Cryptonectes Fosshagen & Iliffe, 2004
Ref.: Fosshagen & Iliffe, 2004 (p.122); 2007 (p.91); Bradford-Grieve & al., 2014 (p.507, 522: Rem., Table 1, fig. 9, 10, 11, cladistic analysis)
Rem.: Total: 1 sp.
(8) Edaxiella Fosshagen, Boxshall & Iliffe, 2001
Ref.: Fosshagen & al., 2001 (p.247, 316); Fosshagen & Iliffe, 2007 (p.92); Bradford-Grieve & al., 2014 (p.507, 522: Rem., Table 1, fig. 9, 10, 11, cladistic analysis)
Rem.: Total: 1sp.
(9) Enantiosis Barr,1984; emend. Fosshagen, Boxshall & Iliffe, 2001
Ref.: Barr, 1984 (p.161); Fosshagen & Iliffe, 1985 (p.346); Huys & Boxshall, 1991 (p.62, 63); Razouls, 1993 (p.306); Mauchline, 1998 (p.70: F; p.73: M; comme Enantiosus); Bradford-Grieve, 1999 b (p.26, Def.); Fosshagen & al., 2001 (p.258, 316); Bradford-Grieve, 2004 (p.287); Fosshagen & Iliffe, 2007 (p.92); Bradford-Grieve & al., 2014 (p.507, 522: Rem., Table 1, fig. 9, 10, 11, cladistic analysis)
Rem.: Total: 7 spp.
Remarks on dimensions and sex ratio:
The mean female size is 1.328 mm (n = 11, SD = 0.115), and the mean male size is 1.235 mm (n = 13, SD = 0.085). The size ratio male: female is 0.923.
(10) Enantronia Fosshagen, Boxshall & Iliffe, 2001
Ref.: Fosshagen & al., 2001 (p.251, 316); Vives & Shmeleva, 2007 (p.993); Fosshagen & Iliffe, 2007 (p.91); Bradford-Grieve & al., 2014 (p.507, 522: Rem., Table 1, fig. 9, 10, 11, cladistic analysis)
Rem.: Total: 1sp.
Type: Enantronia canariensis Fosshagen, Boxshall & Iliffe, 2001.


Diagnosis after Vives & Shmeleva (2007, p.993) :
- As in the family.
- Caudal rami larger than urosomal segments 3 and 4, each; left seta V very long.
- P1-P4 with segmentation as in the family.
- P1 with 1 inner seta on the basipod 1 (= coxa)..
Genus Enantronia - Plate 1Issued from : F. Vives & A.A. Shmeleva in Fauna Iberica, 26, 2007. [p. 994, Fig. 573, A-D]. Enantronia canariensis Female.
A, habitus (dorsal view); B, urosome (lateral view); C, Mxp; D, P1.
(11) Enantronoides Fosshagen, Boxshall & Iliffe, 2001
Ref.: Fosshagen & al., 2001 (p.255, 316); Bradford-Grieve, 2004 (p.287); Fosshagen & Iliffe, 2007 (p.91); Bradford-Grieve & al., 2014 (p.507, 522: Rem., Table 1, fig. 9, 10, 11, cladistic analysis)
Rem.: Total: 1 sp.
(12) Epacteriscus Fosshagen,1973; emend. Fosshagen, Boxshall & Iliffe, 2001
Ref.: Fosshagen, 1973 (p.146); Huys & Boxshall, 1991 (p.53); Razouls, 1993 (p.306); Mauchline, 1998 (p.68); Bradford-Grieve, 1999 b (p.26, Dιf.); Fosshagen & al., 2001 (p.276, 316, Rem.); Bradford-Grieve, 2004 (p.287); Fosshagen & Iliffe, 2007 (p.91); Bradford-Grieve & al., 2014 (p.507, 522: Rem., Table 1, fig. 9, 10, 11, cladistic analysis)
Rem.: type: Epacteriscus rapax. Total: 3 spp.
Genus Epacteriscus - Plate 1Issued from : G.A. Boxshall & S.H. Halsey in The Ray Society, 2004. [p.113, Fig. 21, D, E]. Epacteriscus rapax Female.
D, habitus (lateral view); E, P5.
(13) Erebonectes Fosshagen,1985; emend. Fosshagen, Boxshall & Iliffe, 2001
Ref.: Fosshagen & Iliffe, 1985 (p.346); Huys & Boxshall, 1991(p.50, 51); Razouls, 1993 (p.306); Mauchline, 1998 (p.67); Bradford-Grieve, 1999 b (p.26, Dιf.); Fosshagen & al., 2001 (p.291, 316, Rem.); Bradford-Grieve, 2004 (p.287); Fosshagen & Iliffe, 2007 (p.91); Bradford-Grieve & al., 2014 (p.507, 522: Rem., Table 1, fig. 9, 10, 11, cladistic analysis)
Rem.: Total: 1 sp. Type: Erebonectes nesioticus Fosshagen, 1985.

Definition after Bradford-Grieve (1999, p.26) :
- As in the family.
- Urosome 4-segmented in female; 4-segmented in male.
- Caudal rami bear 6 plumose setae.
- Rostrum directed ventrally, bifurcate, with filaments.
- A1 27-segmented on the left in both male and female; right male 23-segmented, with a distinct geniculation between segments 18 and 19.
- A2 endopod longer than exopod which is 8-segmented.
- Md basipod with 4 setae; endopod 2-segmented, reduced; bladewith 4 strong rounded bicuspid teeth.
- Mx1 well-developed with a shape commonly found among calanoids, but with an elongate, slender endopod.
- Mx2 lobes 1-6 with 6, 3, 3, 3, 4, setae respectively.
- Mxp endopod with 5 free segments.
- All swimming legs on female bear 2 outer-edge spines on exopod segment 3; basipod 2 of P1-P5 with an outer-edge seta or spine.
- P5 male in the form of a complex grasping organ; rami 3-segmented, endopods only slightly modified, exopods hlghly modified.
Genus Erebonectes - Plate 1Issued from : G. A. Boxshall & S.H. Halsey in The Ray Society, 2004. [p.179, Fig. 21, A-C]. Erebonectes nesioticus Male.
A, habitus (dorsal view); B, A1; C, A2.
(14) Erebonectoides Fosshagen, Boxshall & Iliffe, 2001
Ref.: Fosshagen & al., 2001 (p.291, 316); Bradford-Grieve, 2004 (p.287); Fosshagen & Iliffe, 2007 (p.91); Bradford-Grieve, 2014 (p.507, 530, fig.9, 10, 11)
Rem.: 1 sp.
Bradford-Grieve & al. (2014, p.530) consider the genus, after phylogenetic analysis, of uncertain placement in the family (incertae sedis).
(15) Gloinella Fosshagen, 2001
Ref.: Fosshagen & al., 2001 (p.301, 316); Bradford-Grieve, 2004 (p.287); Fosshagen & Iliffe, 2007 (p.92); Bradford-Grieve & al., 2014 (p.507, 522: Rem., Table 1, fig. 9, 10, 11, cladistic analysis)
Rem.: Total: 1 sp.
(16) Iboyella Boxshall & Jaume, 2003
Ref.: Boxshall & Jaume, 2003 (p.86); Fosshagen & Iliffe, 2007 (p.92); Bradford-Grieve & al., 2014 (p.507, 522: Rem., Table 1, fig. 9, 10, 11, cladistic analysis)
Rem.: Total: 1 sp.
(17) Miheptneria Andronov, 2007
Ref.: Andronov, 2007 (p.672); Bradford-Grieve & al., 2014 (p.507, 522: Rem., Table 1, fig. 9, 10, 11, cladistic analysis)
Rem.: Type: Miheptneria abyssalis. Total: 1 species.

After Bradford-Grieve & al., 2014 (p.508, 530) the genus and species belong to the Epacteriscidae.
(18) Minnonectes Fosshagen & Iliffe, 2004
Ref.: Fosshagen & Iliffe, 2004 (p.128: male diagnosis); 2007 (p.88: female diagnosis, p.92); Bradford-Grieve & al., 2014 (p.507, 529: Rem., Table 1, Fig. 9, 10, 11)
Rem.: Total: 1sp.
(19) Oinella Fosshagen, Boxshall & Iliffe, 2001
Ref.: Fosshagen & al., 2001 (p.297, 316); Bradford-Grieve, 2004 (p.287); Fosshagen & Iliffe, 2004 (p.132); 2007 (p.91); Bradford-Grieve & al., 2014 (p.507, 522: Rem., Table 1, fig. 9, 10, 11, cladistic analysis)
Rem.: Total: 1 sp.

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