Phaennidae Family - Marine Planktonic Copepods

Calanoida ( Order )	details
Clausocalanoidea ( Superfamily )	details

Abbreviations or symbols

Phaennidae Sars, 1902 ( Clausocalanoidea )

Ref.:

Sars, 1902 (1903) (p.42); Gurney,1931 a (p.84); Rose, 1933 a (p.125); Brodsky, 1950 (1967) (p.83, 225); Bradford, 1969 b (p.502, Table 3); Mazza, 1967 (p.173); Bradford, 1973 (p.137, Def.); Andronov, 1974 a (p.1005); Bowman & Abele, 1982 (p.9); Razouls, 1982 (p.279); 1993 (p.310); Brodsky & al., 1983 (p.144,146); Park, 1983 a (p.317, Def.); Bradford & al., 1983 (p.58); Mauchline, 1988 (p.735, 740 : cuticular pores); Huys & Boxshall, 1991 (p.356); Chihara & Murano, 1997 (p.852); Bradford-Grieve & al., 1999 (p.880, 902, 904, 927, Key of Genera); Ohtsuka & Huys, 2001 (p.461); Ohtsuka & al., 2003 (p.53, 62: Rem.); Boxshall & Halsey, 2004 (p.15; 49; 158: Def.; p.160: Key of genera); Ferrari & Markhaseva, 2005 (p.45, Rem.); Markhaseva & Ferrari, 2005a (p.111, fig.31, Rem.); Vives & Shmeleva, 2007 (p.678, part. Genera Key); Blanco-Bercial & al., 2011 (p.103, Table 1, Fig.2, 3, 4, Biol. molecular phylogeny); Markhaseva & al., 2014 (p.63, 67, 73, 75, Table 1, 2, 3, 4, Rem., Def.) ; Laakmann & al., 2019 (p.330, Table 1, fig.1, 2, 3, Table A, phylogenetic relationships)
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.:

8 G + 1 G.: Brachycalanus, Cephalophanes, Cornucalanus, Onchocalanus, Phaenna, Phaennocalanus, Talacalanus, Xanthocalanus + Kirnesius.
Schulz (1998, p.42, 48), Vyshkvartzeva (2000, p.217) and Ohtsuka & al., 2003 (p.61, 62) regard the genus Xantharus as belonging to the Scolecitrichidae family. The belonging of this genus to this family is discussed by Schulz & Kwasniewski (2004, p.158).
Markhaseva & Semenova (2005) include provisionally the genus Kirnesius in this family.

The types of setae on the ramus of Mx2 include this family in the 'Bradfordian' grioup of families.

For Markhaseva, Laakmann & Renz (2014, p.73) the family includes 9 genera: Brachycalanus, Cephalophanes, Cornucalanus, Onchocalanus, Phaenna, Phaennocalanus, Talacalanus, Xanthocalanus, Kirnesius.
Type: Phaenna Claus, 1863.
The diagnosis of this family is based on the following combination of characters: the apomorphy within ''Bradfordians'' is the presence of a maximum of 2 worm-like sensory setae on Mx2 endopod, setae are in combination: 1w+7br, 1w+8br, 2w+6br; 2w+7br and 2w+6br+1sc.

Diagnosis after Bradford, Haakonssen & Jillett (1983, p.58), and Boxshall & Halsey (2004, p.158) :
- Cephalosome and pedigerous somite 1 separate, pedigerous somites 4 and 5 partly fused or completely separate.
- Rostrum a bifurcate plate or represented by paired rostral filaments, rarely absent.
- Urosome female short, 4-segmented, notprojected ventrally.
- Genital aperture female located medially on ventral surface ; copulatory pore (s) contained within median genital aperture.
Urosome male 5-segmented.
- Single genital aperture male located ventrolaterally at posterior rim of genital somite on left side.
- Caudal rami short, with up to 5 setae.
- A1 24-segmented in female ; ; segmental homologies (based on Xanthocalanus marlyae Campaner : segment 1 (I) free, segment 2 (II-IV) triple , segments 3 (V) to 7 (IX) separate, segmenys 8 (X) and 9 (XI) fused, segments 10 (XII) to 23 (XXVI) separate, apical (24th) segment double (XXVII-XXVIII). Aestgetascs presenbt on segments III, IV, VII, IX, XI, XIV, XVI, XXI, XXVII-XXVIII.
- A1 non-geniculate on both sides in male ; often with partial segmental fusions in proximal part of antebbule ; sometimes with segments 20 and 21 (XXIII-XXIV) fused on right side only.
- A2 biramous with separate coxa and basis ; coxa with 1 seta ; basis with 2 setae ; endopd 2)segmented, compound distal segment bilobed, setation formula 1, 8, 7 ; exopod indistinctly 6 or 7-segmented, segmental fusions indeterminate except for IX-X, setation formula 0, 1, 1, 1, 1, 1, 4, setation often further reduced. Exopod at most one and one-half times as long as endopod.
- Md biramous ; coxa with well developed gnathobase and distal palp consisting of basis, with 3 setae ; 2-segmented endopod and 5-segmented exopod ; endopodal segments 1 and 23 with 0/2 and 9 setae ; exopodal sdetation formula 1, 1, 1, 1, 2.
- Mx1 with slender well developed praecoxal arthrite bearing 9 to 14 elements ; coxa with endite bearing 2 to 5 setae and with up to 9 setae on epipodite ; basis without outer seta, with proximal endite bearing 3 or 4 setae and distal group of 3 (Phaenna) to 6 setae representing distal endite ; endopod indistinctly 2 or 3-segmented, setation formula often 3, 3, 4/5 ; total number of endopodal setae ranging from 6 in Phaenna to 13 in Brachycalanus ; exopod 1-segmented with 9 or 10 setae, 5 in Phaenna.
- Mx1 males usually with reduced setation on arthrite and endites (inner an douter lobes).
- Mx2 indistinctly segmented, short ; praecoxa and coxa partly fused, setation formula of endites 5 (rarely 4), 3, 3, 3, with 1 element of distal endite sometimes curved, spine-like ; basis with 4 setae or strongly curved claw plus 1 or 2 modified sensoriform filaments and 1 or 2 setae ; free endopd typically with 1 vermiform and 7 brush-like sensoriform filaments ; additional seta on endopoid in Brachycalanus.
- Mx2 often reduced in males.
- Mxp 7-segmented ; syncoxa with endite setation formula 1, 2, 2, 3 ; one element of 2 nd endite sometimes modified as sensoriform filament ; proximal element of 3rd endite always modified as brush-like sensoriform filament ; basis with 3 setae, plus 2 setae on incorporated 1st endopodal segment ; free endopod 5-segmented, segmental setation formula 4, 4, 2/3, 2/3+1, 4.
- Setation on Mx2 males, especially on syncoxa, often reduced.
- Swimming legs P1 to P4 biramous, typically with 3-segmented rami, except 1-segmented endopod of P1 and 2-segmented endopod of P2.
- Endopods of P2 to P4 ornamented with strong spinules on posterior surface.
- Exopods of P2 to P4 sometimes ornamented with spinules.
- Inner seta on basis of P1 situated on anterior surface of basis and passing across face of endopod ; absent in most Onchocalanus species.

The mouthparts of Phaennidae are adaptated for seizing and chewing prey (Arashkevich, 1969). The mandible is heavily sclerotized and the ventral teeth have sharp crowns. The lobes (endites) of Mx1 have dagger-shaped setae. The lobes (endites) of Mx2 have curved setae which may be especially large lobes 4 and 5 ; the endopod has a tuft of sensory filaments. According to Matthews (1964) benthic forms appear to be scavengers.

Key to genera after Boxshall & Halsey ( 2004, p.160) :
1 – Frontal margin of head occupied by conspicuous lenses of enlarged nauplis eye …….. Cephalophanes.
1’ – Nauplius eye without conspicuous lenses …… 2.
2 – Basis of P1 lacking inner seta ……… Onchocalanus (part.)
2’ – Basis of P1 with inner seta ……… 3.
3 – P5 absent in female …….. Phaenna.
3’ – Female with uniramous 2 or 3-segmented P5 ……. 4.
4 – Female with 1 large curved claw on 2nd and 3rd free endopodal segments of female Mxp ……… Cornucalanus.
4’ – 2nd and 3rd free endopodal segments of Mxp without modified claws in female …… 5.
5 – Mx1 with 2 setae on coxal endite ………. 6.
5’ – Mx1 with 3, 4 or 5 setae on coxal endite …….. 7.
6 – A1 short, not reaching end of prosome ……. Brachycalanus.
6’ – A1 reaching beyond end of prosome ……Onchocalanus magnus (Wolfenden).
7 – 2 brush-like sensoriform elements much thicker than remaining 5 elements and all 7 of similar length ; female P5 lacking surface spinulation and armed with single apical spine …..Phaennocalanus.
7’ – Brush-like sensoriform elements not of this pattern ; female P5 with surface spinulation and typically armed with 2 to 4 spines …..8.
8 – Modified razor-like setae present on praecoxal arthrite of Mx1 and on free endopodal segments of Mxp …….. Talacalanus.
8’ – Setae on praecoxal arthrite of Mx1 and on endopod of Mxp not razor-like ……… Xanthocalanus.

For Markhaseva, Laakmann & Renz (2014, p.73), the diagnosis of this family is base on the combination of the following characters; the apomorphy (character derived from and differing from an ancestral condition) for the family within ''Bradfordians'' is the presence of a maximum of 2 worm-like sensory setae on the Mx2 endiopod, setae are in combination: 1w+7br, 1w+8br, 2w+6br, 2w+7br and 2w+6br+1sc. [w = worm-like seta, br = brush-like seta, sc = sclerotised seta].
General characters :

1 - Proximal part of right A1 in males shows several types of fusions: 1st type observed in Cornucalanus, Onchocalanus, Cephalophanes and some species of Xanthocalanus with only ancestral segments X-XI fused (shared with Kyphocalanidae, Parkiidae, 3 diaixid genera and 2 genera of unclear family placement; 2nd type in some species of Xanthocalanus with segments X-XI fused, segments XI and XII incompletely or completely fused, and segments XII-XV fused (segments XIV-XV can be only incompletely fused); 3rd type found in Phaenna, with segments X-XI fused and segments XII-XVI fused and segments XII--XVI fused (see fig. 5).

2 - A2 coxa with 1 seta; basis with 1 seta (Phaennocalanus, Phaenna, with 1-2 setae in Onchocalanus and some species of Xanthocalanus and with 2 setae in the remaining genera; endopod segment 1 with 2 setae (Cornucalanus, Phaennocalanus, Talacalanus), 1-2 setae (Brachycalanus, Onchocalanus, Xanthocalanus), and 1 seta (Phaenna, Cephalophanes); the setal formula of exopod is: 0, 0-0-1, 1, 1, 1, 1, 1, 3 (Brachycalanus and some species of Xanthocalanus; 0, 0-0-1-1, 1, 1, 1, 1, , 3 (Onchocalanus; 0, 0-0-0, 1, 1, 1, 1, 1, 3 (Talacalanus, Xanthocalanus) and 0, 0-0-0-1, 1, 1, 1, 1, 3 in the remaining genera (see Table 1).

3 - Md endopoal segment 2 with 9 setae; exopod 5-segmented with 6 setae (in Xanthocalanus 4-5 setae); gnathobase without hump (crest), usually elongate (see fig. 6b, Table 2).

4 - Mx1 setal formula, typically: 10-14, 2-5, 3-4, 4-5, 6-11, 10, 9 setae; Phaenna deviates in praecoxal arthrite wiyj 9, coxal and distal basal endites with 1 and 3 setae respectively, and exopod with 5 setae, some species of Brachycalanus also can have praecoxal arthrite with only 9 setae, and in Xanthocalanus exopod can bear 9-10 setae; distal basal endite separate from endopod (see Table 3).
5 - Mx2 endites setal formula: 5, 3, 3, 3, 4; some species of Brachycalanus can have 4-5 setae on the proximal endite of the praecoxa (see Table 4).

6 - Mxp praecoxal endites setal formula: 1, 2, 2 setae (fig. 6f); praecoxal setae: 1sc + 2sc + 2sc, 1sc + 2sc + (1sc+1br); 1w + 2sc + (1sc+1br), and 1w + (1sc+1w) + (1sc+1br); basis with 3 proximal and 2 distal setae (see Table 5).

7 - Male P5, uniramous (rarely biramous) ; asymmetrical in different degree; of simple structure, (segments of cylindrical shape); right coxa and basis significantly shorter than left; exopods 3-segmented, solmetimes in right leg: 1-2 segmented (type of leg shared with 3 diaixid genera (see fig. 7 a-d).

All genera share the Mx2 endopod with a maximum number of 2 worm-like sensory setae. Minor variations in the Mx2 endopod setation that were describede, i.e. 1w + 8br , 2w + 7br and 2w + 6br + 1sc (Campaner, 1978; Ferrari & Markhaseva, 2000; Markhaseva & Schnack-Schiel, 2003), retain Bradford's robust diagnosis. The placement of Kirnesius in under question, until a female is discovered.

Issued from : E.L. Markhaseva, S. Laakmann & J. Renz in Mar. Biodiv., 2014, 44 [p.70, Fig. 5].
Schematic view of right A1 in males of ''Bradfordian'' family Phaennidae.
Black arrows: fused ancestral segments; Roman numerals: ancestral segments; dotted line between antennule segments: incomplete fusion.

Issued from : E.L. Markhaseva, S. Laakmann & J. Renz in Mar. Biodiv., 2014, 44 [p.73, Table 1].
A2 armament (number of seta) in different ''Bradfordian'' genera (females). Phaennidae.
c: coxa; b: basis; End1: endopod segment 1; End2: endopod segment 2; Exp: exopod.

Issued from : E.L. Markhaseva, S. Laakmann & J. Renz in Mar. Biodiv., 2014, 44 [p.74, Table 2].
Md armament (number of seta) in different ''Bradfordian'' genera (females). Phaennidae.
b: basis; End1: endopod segment 1; End2: endopod segment 2; Exp: exopod; gn: gnathobase.

Issued from : E.L. Markhaseva, S. Laakmann & J. Renz in Mar. Biodiv., 2014, 44 [p.76, Table 3].
Mx1 armament (number of seta) in different ''Bradfordian'' genera (females). Phaennidae.
pa: praecoxal arthrite (setal formula of praecoxal arthrite: terminal+posterior+anterior setae); ce: coxal endite; bp: proximal basal endite; bd: distal basal endite; End: endopod; Exp: exopod; Epi: epipodite.

Issued from : E.L. Markhaseva, S. Laakmann & J. Renz in Mar. Biodiv., 2014, 44 [p.78, Table 4].
Mx2 armament (number of seta) in different ''Bradfordian'' genara (females). Phaennidae.
at: attenuation; pe: praecoxal endite; ce: coxal endite; bp: proximal basal endite; bd: distal basal endite; el: enditic-like lobe of proximal endopodal segment; End: endopod; w: worm-like seta; br: brush-like seta; sc: sclerotised seta.

Issued from : E.L. Markhaseva, S. Laakmann & J. Renz in Mar. Biodiv., 2014, 44 [p.80, Table 5].
Mxp setation (number of seta) in different ''Bradfordian'' genera (females). Phaennidae.
at: attenuation; pes: praecoxal endites of syncoxa (from proximal to distal); ces: coxal endite of syncoxa; bp: basis proximal; bd: basis distal; End: endopod.
For more detailed morphology of the seta on the praecoxal endites of the maxilliped syncoxa see Markhaseva & Ferrari (2005) and Markhaseva & al. (2008).

Issued from : G.A. Boxshall & S.H. Halsey in An Introduction to Copepod Diversity. The Ray Society, 2004, No 166, Part. I. [p.158].
Armature formula of swimming legs P1 to P4.

Nota:
- P5 female small, uniramous, typically 3-segmented, comprising unarmed coxa, basis and 1-segmented exopod. Exopod bearing up to 4 elements, often highly ornamented with surface spinules, rarely unornamented. Legs sometimes reduced, absent in Phaenna.
- Male P5 asymmetrical, left leg longer than right: typically uniramous but rudimentary endopods sometimes present. Left leg primitively 5-segmented, with slender coxa, basis and 3-segmented exopod. Right leg 3, 4 or 5-segmented, consisting of coxa, basis and 1 to 3-segmented exopod.
Eggs released into water.

Issued from : S. Laakmann, E.L. Markhaseva & J. Renz in Mol. Phylog. Evol., 2019, 130. [p.331, Table 1].
compilation of information on relationships among ''Bradfodian'' genera from Markhaseva & Ferrari 2005) and Markhaseva & al. (2014).
Abbreviations: A2, antenna; Md, mandible; Mx1, maxillule; Mx2, maxilla; Mxp, maxilliped ; P5, leg 5. w, worm-like sensory seta; b, brush-like sensory seta; s, sclerotized seta.

(1) Brachycalanus Farran, 1905

Species, varieties and details

Ref.:

Farran, 1905 (p.41); van Breemen, 1908 a (p.66); A. Scott, 1909 (p.81); Sewell, 1929 (p.176); Wilson, 1932 a (p.75); Rose, 1933 a (p.139); Tanaka, 1960 a (p.102); Bradford, 1973 (p.138); Campaner, 1978 a (p.976, Redef.); Razouls, 1982 (p.279); Bradford & al., 1983 (p.60, Def.); Othman & Greenwood, 1988 a (p.356, Rem.); Razouls, 1993 (p.310); Mauchline, 1998 (p.84: F); Ohtsuka & al., 2003 (p.62: Rem.); Bradford-Grieve, 2004 (p.287); Boxshall & Halsey, 2004 (p.161); Markhaseva & al., 2014 (p.73, Table 1, 2, 3, 4, Rem.); Renz & Markhaseva, 2015 (p.96, Table 4, fig.3, sex ratio, biogeography); Laakmann & al., 2019 (p.330, Table 1)

Rem.:

type: Xanthocalanus atlanticus (Wolfenden,1904). Total: 7 spp. (1 doubtful) + 1 undet.

Redefinition after Campaner (1978 a, p.976), and [Bradford & al., 1983, p.60] (Female) :
- Body robust and oval dorsally.
- Cephalosome and thoracic segment 1 separated, thoracic segments 4 and 5 complete or incompletely separated or totally fused.
- Rostrum short with 2 filaments.
- Abdomen 4-segmented, densely covered with scale-like spines (anal segment extremely reduced).
- A1 short, 24-segmented [with large aesthetascs].
- Exopod of A2 little greater than endopd.
Inner lobe 1 [2] and 3 of Mx1 with respectively 2 and 4 setae.
- Endopod of Mx2 with 1 vermiform [worm-like] seta and ( ?5) 7-8 brush-like setae [7 sensory filaments].
- The dorsal and dorsal surfaces of basis 1-2, and especially of exopod and endopod, from P1-P5 densely covered with scale-like spines.
- P5 with 3 segments, last one with 2 lateral and 2 terminal aculeate spines.

Type-species :

Campaner (1978 a, p.976) disagrees with Grice (1972), who placed B. minutus in this genus based mostly on the absence of rostral filaments.
Xanthocalanus ordinarius Grice (1972) is doubtless a new species of Brachycalanus.
Since B. gigas A. Scott, 1909, was definitely removed to the genus Lophothrix, the genus hitherto includes 3 species : B. atlanticus (Wolfenden, 1904), B. ordinarius (Grice, 1972), B. bjornbergae Campaner, 1978.
B. minutus Grice, 1972, is a species incertae sedis.

Remarks on dimensions and sex ratio:

The mean female size is 2.371 mm (n = 14; SD = 0.7179). No male has been observed yet.

(2) Cephalophanes Sars, 1907

Species, varieties and details

Ref.:

Sars, 1907 a (p.15); Wolfenden, 1911 (p.283); Sars, 1925 (p.154); Steuer, 1926 b (p.214); Rose, 1933 a (p.141); Vervoort, 1957 (p.93, Rem.); Tanaka, 1960 a (p.124); Bradford, 1973 (p.138); Razouls, 1982 (p.280); Park, 1983 a (p.319, Def.); Bradford & al., 1983 (p.60, Def.); Mauchline, 1988 (p.735: cuticular pores); Razouls, 1993 (p.310); Chihara & Murano, 1997 (p.852); Boxshall & Halsey, 2004 (p.160); Markhaseva & Ferrari, 2005a (p.111, fig.31, Rem.); Kuriyama & Nishida, 2006 (p.293, Rem.: p.313); Vives & Shmeleva, 2007 (p.679, spp. Key); Markhaseva & al., 2014 (p.73, Table 1, 2, 3, 4, Rem.); Laakmann & al., 2019 (p.330, Table 1)

Rem.:

Type: Cephalophanes refulgens Sars, 1907. Total: 3 spp.

Diagnosis after Park (1983 a, p.319) , and Bradford, Haakonssen & Jillett (1983, p.60) :
Female:
- Head and pedigerous segment and pedigerous segments 4 and 5 fully separate.
- Rostrum of 2 slender filaments.
- Forehead with a pair of large eyes, each occupying almost entirely half of forehead. A1 25-segmented.
- Exopod of A2 much longer than endopod.
- Mx2 terminated by 6 or 7 slender brush-like appendages and perhaps 1 worm-like appendage. Two of 4 setae on 5th basipodal lobe (= endite) of Mx2 large and spiniform, but not conspicuously larger than large setae of other basipodal lobes.
- Endopods of P2 to P4 armed posteriorly with relatively long spinules.
- P5 uniramous, 3-segmented, armed with spinules and spines (sometimes very fragile and likely broken off).
Male:
- P5 4-segmented on each side; right leg much shorter than left.

After Sars (1925, p.155), the bright device in C. refulgens) consists of two symmetric half , of prismatic shape, filling almost completely the frontal part of the head, and arranged so as to be contiguous to down, whereas their superior sharp edges are moved away enough one of the other one. Inside of every organ there is a dense bundle of radiant fibers wrapped with a pale pigment, doubtless the generators of the light, which is reflected by the external plane surface of organs provided of a chandelier very bright and pearly . This chandelier however faints after dipping of specimens in the alcohol, better kept in the formalin.

Remarks on dimensions and sex ratio:

The mean female size is 4.00 mm (n = 6; SD = 1.0738), and the mean male size is 3.688 mm (n = 4; SD = 0.4837). The size ratio (Male: Female) is 0.90.

(3) Cornucalanus Wolfenden, 1905

Species, varieties and details

Ref.:

Wolfenden, 1905 a (p.20); van Breemen, 1908 a (p.222); A. Scott, 1909 (p.83); Wolfenden, 1911 (p.281); Sars, 1925 (p.151); Sewell, 1929 (p.176); Wilson, 1932 a (p.74); Rose, 1933 a (p.138); Brodsky, 1950 (1967) (p.236); Vervoort, 1957 (p.90, Rem.); Tanaka, 1960 a (p.121); Razouls, 1982 (p.281); Bradford & al., 1983 (p.61, Def.); Park, 1983 a (p.351, Def.); Mauchline, 1988 (735: pores cuticulaires); Razouls, 1993 (p.310); Mauchline, 1998 (p.81: M; p.82: F); Bradford-Grieve & al., 1999 (p.929, spp. key); Boxshall & Halsey, 2004 (p.160); Markhaseva & Ferrari, 2005a (p.111, fig.31, Rem.); Vives & Shmeleva, 2007 (p.682, spp. Key); Markhaseva & al., 2014 (p.73, Table 1, 2, 3, 4, Rem.); Laakmann & al., 2019 (p.330, Table 1)

Rem.:

6 spp. + 1 unident.

Diagnosis after Bradford & al. (1983, p.61) :
- Head and pedigerous segment 1 usually separate, 4th and 5th thoracic segments fused or separate.
- Head with or without spine.
- A1 34-25-segmented in female, fewer segments in male.
- Mx2 with strong claw-like spine on lobe 5 as in Onchocalanus
- Mxp endopod segments 2 and 3 each with a strong spine.
- P5 uniramous, symmetrical in female; asymmetrical in male.

Diagnosis after Park (1983 a, p.351) :
Female :
- Head and 1st pedigerous segment separate, 4th and 5th thoracic segments partially fused.
- Rostral plate strong, divided distally into 2 large rami, each ramus often folloiwed by a delicate filaments.
- A2 exopod slightly longer than endopod.
- One of 3 setae on 4th basipodal lobe of Mx2 enlarged into a sttrong spine; 5th basipodal lobe with a greatly enlarged spiniform seta, strongly curved resembling a sickle, and a small seta in the form of a simple crankshaft.
- 2nd and 3rd endopodal segments of Mxp each with a large, claw-shaped seta.
- Basis of P1 with a well-developed inner seta.
- In P2 to P4, whole posterior surfaces of both rami armed with spinules.
- P5 uniramous, 3-segmented. Last 1 or 2 segments armed with patches of spinules.

Male:
- Habitus similar to female.
- Urosome 5-segmented.
- A1 12th and twenty-first segments fused on right side but separate on left.
- A2 and P1 to P4 similar those of female;
- Md, Mx1 and Mx2 similar in setation to those of female, but more or less reduced in their development.
- None of endopodal setae of Mxp enlarged into claws.
- Left P5 long, 5-segmented. Right leg very short, 3- to 5-segmented.

For Park (1983 a, p.352) the females of the genus are very characteristic: a strong, sickle-shaped spine and a small, crankshaft-shaped seta on the 5th Mx2 lobe and 2 of the endopodal setae of Mxp greatly enlarged into claws.
The males are very similar to those of Onchocalanus but can be distinguished from them by the shape of the strong, spiniform setae on the 4th and 5th Mx2 lobes and the presence of an inner seta on the basis of P1.

Remarks on dimensions and sex ratio:

The mean female size is 6.907 mm (n = 10; SD = 1.7039), and the mean male size is 6.723 mm (n = 6; SD = 1.1541). The size ratio (Male: Female) is 0.86 (n = 3; SD = 0.0752). The sex ratio (female: male) is 2.

(4) Kirnesius Markhaseva & Semenova, 2005

Species, varieties and details

Ref.:

Markhaseva & Semenova, 2005 (p.40); Markhaseva & Schulz, 2009 (p.22, Rem.); Laakmann & al., 2019 (p.330, Table 1)

Rem.:

Type: Kirnesius groenlandicus Markhaseva & Semenova, 2005. Total; 1 sp.
For Markhaseva & Schulz (2009, p.22) the family placement in Phaennidae is only provisionally due to the presence of rudimentary oral parts and the uniramous P5. Segmentation and setation of swimming legs and the structure of A1 of the species-type are typical of the superfamily CVlausocalanoidea. The presence of A1 brush-like aesthetascs is unique in the order Calanoida, and is an apomorphiy that separates the new genus from all other described clausocalanoideans.

Diagnosis from Markhaseva & Semenova (2005, p.40) :
Male:
- Cephalosome anf 1st pedigerous somite separated, 4th and 5th incompletely separated.
- Rostrum large, thick, oval-rectangular and lacking filaments.
- A1 non-geniculate on both sides. Brush-like aesthetascs present in proximal part of A1 on articulated segments 2 and 5 (ancestral segments II-IV and VII respectively). A1 slightly asymmetrical, 23-segmented on right, 22 segmented on left side, due to fusion of segments 18 and 19 (ancestral segments XXII-XXIII).
- Mouthparts moderately reduced.
- Mx2 with 8 sensory setae on exopod, praecoxal endite with 5 setae.
- Syncoxa of Mxp with 4 setae on praecoxal lobes: 1 scletotized seta on proximal lobe, 1 sensory worm-like plus 1 sclerotized on medial lobe and 1 sclerotized on distal lobe.
- P1 endopod 1-segmented.
- P2 endopod 2-segmented.
- P3 - P4 endopods 3-segmented.
- P1 - P2 exopods 3-segmented, P3 - P4 endopods broken.
- P1 exopodal segment 3 with 3 medial setae.
- P2 exopodal segment 3 with 4 medial setae.
- P2 - P4 endopods and coxopods poorly spinulated.
- P5 uniramous, 5-segmented on both sides, of simple stucture.
Female unknown.

Remarks on dimensions and sex ratio:

Body size in only one male: 1.6 mm.

(5) Onchocalanus Sars, 1905

Species, varieties and details

Ref.:

Sars, 1905 b (p.19); van Breemen, 1908 a (p.65); A. Scott, 1909 (p.82); Wolfenden, 1911 (p.275); Sars, 1925 (p.144); Sewell, 1929 (p.176); Rose, 1933 a (p.136, spp. Key); Vervoort, 1950 a (p.3, spp. Key F); Brodsky, 1950 (1967) (p.234); Tanaka, 1960 a (p.115); Bradford, 1973 (p.136, 138); Razouls, 1982 (p.283); Gardner & Szabo, 1982 (p.269); Bradford & al., 1983 (p.61, Def., Rem,.p.66); Park, 1983 a (p.327, Def., spp. Key); Mauchline, 1988 (p.735: cuticular pores); Razouls, 1993 (p.310); Chihara & Murano, 1997 (p.853); Mauchline, 1998 (p.81: M; p.82: F); Bradford-Grieve & al., 1999 (p.929, spp. Key); Boxshall & Halsey, 2004 (p.160, 161); Markhaseva & Ferrari, 2005a (p.111, fig.31, Rem.); Vives & Shmeleva, 2007 (p.689, spp. Key); Markhaseva & al., 2014 (p.73, Table 1, 2, 3, 4, Rem.); Laakmann & al., 2019 (p.330, Table 1)

Rem.:

type: Onchocalanus trigoniceps Sars,1905. Total: 10 spp.

Diagnosis after Park (1983 a, p.327) :
Female:
- Cephalosome and 1st pedigerous segment usually separate, 4th and 5th thoracic segments usually separate.
- Rostrum large plate, distally divided into 2 strong spiniform rami.
- A2 with the two rami about equal in length.
- 5th basipodal lobe (= endite) of Mx2 with a large spiniform seta, smoothly curved like a claw.
- None of endopodal setae of Mxp enlarged into a strong claw.
- Basis of P1 without an inner seta.
- Whole posterior surface of both rami of P2 to P4 densely armed with spinules.
- P5 uniramous, 3-segmented, with whole surface densely covered by relatively long spinules.

Male:
- Similar in habitus to female.
- Urosome 5-segmented.
- 12th and 21th segments separate in left A1 but fused in right.
- Cephalosomal appendages from A1 to Mxp similar to those of female but more or less reduced in size and development of setae.
- P1 to P4 almost identical with yhose of female.
- P5 strongly asymmetrical, with long, 5-segmented left leg and small, 3- or 4- segmented right leg.

Diagnosis from Bradford & al. (1983, p.61), after Vervoort, 1950 :
- Female with head and pediger segment 1, as well as pediger segments 4 and 5, imperfectly separated.
- Head crested in some species.
- Rostrum in form of sclerotised, bifurcated plate.
- A1 24-segmented; in female segments 8 and 9 fused; in male segments 8-13 fused.
- A2 endopod and exopod nearly equal in length.
- Md palp basipod 2 (= basis) of female with 3 setae , endopod segment 1 with 2 setae, endopod segment 2 with 9 setae; in male basipod 2 (= basis) large.
- Female Mx1 inner lobe 1 (= praecoxal arthrite) with 7 strong setae and 3 or 4 bristles; inner lobe 2 (= coxal endite) with 3 setae; inner lobe 3 (= basal endite) with 4 setae; outer lobe 1 (= basal exite1) with 9 setae; exopod with 10 setae; outer lobe 2 (= basal exite2 missing; basipod 2 (= basis) with 5 setae; endopod segments 1-3 with 3, 2 or 3 and 4 setae respectively.
- Female Mx2 lobe 5 with 1 seta claw-like and minutely denticulated near apex; endopod with 7 brush-like and 1 worm-like sensory filaments.
- Mxp basipod 2 (= basis) long with 1 proximal sensory appendage.
- Male Mx1 and 2 less well sclerotised and with weaker setae than female.
- Posterior surface of legs covered with spines.
- Female P5 3-segmented, 1st segment more or less fused with ventral surface of metasome; distal segment with 2 apical spines which may articulate with segment and an inner and outer spine; 1 of marginal spines, which usually are articulate, may be absent; posterior surface covered with strong hairs.
- Male P5 uniramous on each side; right leg short, 3-segmented; left leg elongate, about as long as urosome and consists of 2 basal and 3 terminal segments.

Remarks on dimensions and sex ratio:

The mean female size is 6.572 mm (n = 18; SD = 1.649), and the mean male size is 6.040 mm (n = 13; SD = 1.462). The size ratio (Male: Female) is 0.919.

(6) Phaenna Claus, 1863

Species, varieties and details

Ref.:

Claus, 1863 (p.188); Giesbrecht, 1892 (p.57, 293); Giesbrecht & Schmeil, 1898 (p.50); van Breemen, 1908 a (p.55); A. Scott, 1909 (p.79); Rose, 1933 a (p.126); Mori, 1937 (1964) (p.54); Bradford, 1973 (p.138); Razouls, 1982 (p.286); Park, 1983 a (p.318, Def.); Bradford & al., 1983 (p.67, Def.); Mauchline, 1988 (p.734: cuticular pores); Razouls, 1993 (p.310); Mauchline, 1998 (p.78); Boxshall & Halsey, 2004 (p.160); Mulyadi, 2004, Def.): Markhaseva & Ferrari, 2005a (p.111, fig.31, Rem.); Vives & Shmeleva, 2007 (p.687); Markhaseva & Renz, 2011 (p.68, Rem.); Markhaseva & al., 2014 (p.73, Table 1, 2, 3, 4, Rem.); Laakmann & al., 2019 (p.330, Table 1)

Rem.:

Type: Phaenna spinifera Claus, 1863. Total: 4 spp. (of which 2 doubtful).

Diagnosis after Park (1983 a, p.318) :
Female:
- Cephalosome and 1st pedigerous segment fully separate, 4th and 5th thoracic segments almost completely separate.
- Rostrum of 2 strong spiniform rami.
- Exopod of A2 considerably longer than endopod.
- 4th and 5th basipodal lobes (= endites) of Mx2 each with a large spine; that of 5th basipodal lobe much stronger and more curved.
2nd endopodal segment of P2 and 2nd and 3rd endopodal segments of P3 and P4 armed with relatively strong spinules.
- P5 absent.

Definition after Bradford, Haakonssen & Jillett (1983, p.67) and Vives & Shmeleva (2004, p.687) :
- Body globular.
- Cephalosome and pediger segment 1 separate, pediger segments 4 and 5 incompletely separate.
- Rostrum bifurcate, spiniforme.
- Urosomefemale 4-segmented.
- Urosome male 5-segmented.
- A1 female 24-segmented.
- Anal segment very short, more or less concealed.
- Caudal rami very short. , aboutb wider than long.
- A1 male 20-segmented, with short feeble setae. [18 -19 segments on left or right, respectively, after Vives & Shmeleva, 2007, p.657]
- A2 exopod longer than endopod.
- Masticatory process of Md and inner lobe 1 (praecoxal arthrite) of Mx1 very long and exopod short with only 5 setae.
- Mouthparts of male reduced.
- Mx2 with 1 worm-like and 7 brush-like elements.
- P5 female absent.
- P5 male uniramous on each side, left leg 5-segmented, right leg 4-segmented.

Remarks on dimensions and sex ratio:

In this genus, only two species are absolutely certain, the mean female size are 2.30 mm and 4.15 mm. Males in 1 or 2 species (P. spinifera and ? in P. zetlandica. If we consider the four species like correct we have: the mean female size is 2.903 mm (n = 6; SD = 1.0945) and the mean male size is 2.267 mm (n = 3; SD = 0.4041). The size ratio (male : female) is 0.781.

(7) Phaennocalanus Markhaseva, 2002

Species, varieties and details

Ref.:

Markhaseva, 2002 (p.312, Def.); Boxshall & Halsey, 2004 (p.161); Markhaseva & Ferrari, 2005a (p.11, fig.31, Rem.); Markhaseva & al., 2014 (p.73, Table 1, 2, 3, 4, Rem.); Laakmann & al., 2019 (p.330, Table 1)

Rem.:

Type: Phaennocalanus unispinosus Markhaseva, 2002. Total: Total: 1 sp.

Diagnosis from Markhaseva (2002, p.312) ;
- Rostrum with 2 thin, long filaments.
- Lobe 2 of Mx1 with 5 setae; exopod with with 10 setae.
- Mx2 lobe 1 with 5 setae; inner lobes 4-5 with one of setae spine-like; terminal part complrises distal basal lobe plus exopod with 1 worm-like and 7 brush-like densory setaze, the latter are: 2 thick brushes plus 5 thin brushes.
- Mxp syncoxa with 5 praecoxal setae in total, all are sclerotized.
- P5 3-segmented lacking surface spinulation, with 1 terminal spine.

For Markhaseva (2002, p.316) the genus differs from other phaennid genera by the combination of derived characters: 1- A1 8th, 13th and 18th segments with knife-like aesthetascs; 2- Mx2 brush-like setae are of nearly the same length, but not identical in structure (there are 2 thick brushes and 5 brushes are thin with small heads; 3- P1 exopodite 1 and 2 with lateral spines narrow abruptly at its proximal 1/3 length; 4- P5 3-segmented with 1 terminal spine, segments lacking surface spinulation.
Plesiomorphic character states separating the new genus from the other phaennids by the combination: 1- Mx1 inner lobe 2 with 5 setae; 2- Mxp syncoxa with 2 sclerotized setae on the distal praecoxal lobe (this is the 3rd group of setae on syncoxa from proximal to distal).
The knife-like aesthetascs of Phaennocalanus are similar to those of Brachycalanus. Aesthetascs of such shape were proposed as a synapomorphy for species of Brachycalanus (Ferrari & Markhaseva, 2000). This is no longer the case and the onky shared derived character state for species of Brachicalanus is epicuticularextension of female genital segment and following two urosomal somites.

(8) Talacalanus Wolfenden, 1911

Species, varieties and details

Ref.:

Wolfenden, 1911 (p.279); Campaner, 1978 a (p.976, Rev.); Razouls, 1982 (p.297); Park, 1983 a (p.327, Rem.); Bradford & al.,1983 (p.70, Rem.); Mauchline, 1988 (p.735, Rem.); Razouls, 1993 (p.310); Mauchline, 1998 (p.81: M; p.82: F); Bradford-Grieve & al., 1999 (p.928: Rem.); Boxshall & Halsey, 2004 (p.161); Markhaseva & Ferrari, 2005a (p.111, fig.31, Rem.); Markhaseva & al., 2014 (p.73, Table 1, 2, 3, 4, Rem.); Laakmann & al., 2019 (p.330, Table 1)

Rem.:

type: Xanthocalanus greeni Farran,1905.
Genus maintained (Campaner, 1978 a; Bradford & al., 1983) or not according to the authors: Sars, 1925, p.134; Park, 1983 a et Tanaka & Omori, 1992 (p.268), very close to Xanthocalanus, but is distinguished from it mainly by the broad razor-like setae on praecoxal arthrite (inner lobe 1) of Mx1 and the endopod of Mxp

Redefinition after Campaner (1978, p.976) , and [Bradford & al; (1983, p.70)]:
- Body lengthened (about 6.0 mm or greater).
- Cephalosome and thoracic segment 1 separated, thoracic segments 4 and 5 fused.
- Anal segment strongly reduced.
- A1 with 24 segments (with few and short setae), reaching about the body length.
- Endopod and exopod of Md palp with nearly the same length, Basis 1+2 broad, with 3 setae.
- Inner lobe of Mx1 very developed, with razor-like type setae in female and fleshy-type (flagellum-shaped from the middle to the tip) setae in male, inner lobe 3 with 3 setae.
- Endopod of female Mx2 usually with [1] 2 vermiform setae +[7] 6 brush-like setae (1 short and broad) ; endopod of male with brush-like setae forming a bud-like structure, and with ( ?) some vermiform setae.
- Endopod of Mxp with razor-like type setae.
- Endopod of P2-P4 with 1 or 2 groups of scale-like spines.
- Female P5 with 3 segments, 3rd segment relatively short with 1 terminal plus 1 or 2 subterminal spines ; male P5 very asymmetrical in length, both with rudimentary endopod.

The genus Talacalanus differs from the genus Xanthocalanus mostly by :
The setation on Mx1 and endopod setation of Mxp, the relatively great body length.
Talacalanus specimen, an aspect which does not correspond to that figured by Wolfenden (1911, pl.31, fig.11).

Type-species : Xanthocalanus greeni Farran, 1905.
Other species :
Talacalanus calaminus Wolfenden, 1911. X. species 2 Grice, 1972 ; X. maximus Brodsky, 1950 (redescribed by Tanaka & Omori, 1967) certainly belong to this genus.

Provisional total: 2 spp. + 1 unident.

Remarks on dimensions and sex ratio:

The mean female size is 8.825 mm (n = 2, SD = 1.9805), and in male 7.360 mm (n = 4; SD = 0.5224). The size ratio (male: female) is 0.92.
The sex ratio is 1.

Xantharus Andronov, 1981

Ref.:

Andronov, 1981 (p.1719); Bradford & al., 1983 (p.60, 104, Rem.); Schulz, 1998 (p.42: Rem.: Scolecithrichidae); Vyshkvartzeva, 1999 (2000) (p.217: Rem: Scolecitrichidae); Ohtsuka & al., 2003 (p.61, 62: Rem.)

Rem.:

Cf. Scolecitrichidae

(9) Xanthocalanus Giesbrecht, 1892

Species, varieties and details

Syn.:

Amallophora (part.) T. Scott, 1894 b (p.54); Tanaka, 1960 a (p.102, Rem.); ? Talacalanus Wolfenden, 1911 (p.279); Campaner, 1978 a (p.976); Bradford & al., 1983 (p.70); Mauchline, 1988 (p.735); Park, 1983 a (p.327) considère le genre Talacalanus comme un synonyme.

Ref.:

Giesbrecht, 1892 (p.57, 286); Giesbrecht & Schmeil, 1898 (p.49, spp. Key); Sars, 1900 (p.48); 1902 (1903) (p.45); Esterly, 1906 a (p.68); van Breemen, 1908 a (p.57); A. Scott, 1909 (p.80); T. Scott, 1909 (p.125, Rem.); Wolfenden, 1911 (p.272); Sars, 1919 (1921) (p.2); 1925 (p.125); Sewell, 1929 (p.176); Wilson, 1932 a (p.68, Key spp.); Rose, 1933 a (p.127, spp. Key); Brodsky, 1950 (1967) (p.226, spp. Key); Vervoort, 1957 (p.95, Rem.); Tanaka, 1960 a (p.86); Bradford, 1973 (p.133, 139, Redef.); Campaner, 1978 a (p.968, Rev.); Razouls, 1982 (p.287); Park, 1983 a (p.325, Def. F.); Bradford & al., 1983 (p.70 & suiv., Def.); Mauchline, 1988 (p.735); Ferrari, 1992 (p.392, tab.3); Razouls, 1993 (p.310); Chihara & Murano, 1997 (p.854); Mauchline, 1998 (p.81: M; p.82, 84, 85: F); Bradford-Grieve & al., 1999 (p.929, spp. Key); Vyshkvartzeva, 2003 (p.45: Rem.); Bradford-Grieve, 2004 (p.640, Rem.); Boxshall & Halsey, 2004 (p.161); Markhaseva & Ferrari, 2005a (p.111, fig.31, Rem.); Vives & Shmeleva, 2007 (p.695, spp. Key); Markhaseva & al., 2014 (p.73, Table 1, 2, 3, 4, Rem.) ; Renz & Markhaseva, 2015 (p.96, Table 4, Fig.3, biogeography); Laakmann & al., 2019 (p.330, Table 1)

Rem.:

types : Xanthocalanus agilis & minor Giesbrecht,1892.
Campaner (1978 a, p.968) classifies the species into 2 groups of which the types for each group are: X. agilis et X. minor .
‘’Xanthocalanus agilis Group’’
Characters :
Cephalosome and 1st thoracic segment usually separated, 4th and 5th thoracic segments separated or fused.
Female A1 usually with 24 segments.
Md palp with endopod and exopod nearly aqual in length.
Mx1 : the large setae of the inner lobe of peculiar type (aculeate-straigth or slightly curved setae, bearing sharp spinules and/or setules) ; inner lobe 3 with 3 or 4 setae.
Endopod of Mx2 with 2 vermiform-type setae and 5 brush-like type setae, sometimes with an extra short brush-like seta.
Endopodite de P2-P4 usually with 1 or 2 groups of large spines.
P5 female with 3 segments, the 2 nd segment not much wider than the 1 st or 3 rd segment, this last either relatively narrow, long and with 1 or 2 terminal plus 2 lateral spines, or short, broad and with 2 or 3 spines : male P5 very or strongly asymmetical in length.
List of species :
Type-species : Xanthocalanus agilis Giesbrecht, 1892
? X. subagilis Wolfenden, 1904.
Species definitely in this group : X. fallax Sars, 1921 (= X. borealis Sars, 1903, part.) ; X. pinguis Farran, 1905 ; X. profundus Sars, 1907. Species possibly in this group : X. mixtus Sars, 1920 ; X. tenuiserratus Wolfenden, 1911.
Doubtful species in this group (most of them related with the genus Cephalophanes Sars, 1907, excepting by the absence of the frontal lenses) : X. antarcticus Wolfenden, 1908 ; X. squamatus Farran, 1905 ; X. agilis Wolfenden, 1911 ; X. kurilensis Brodsky, 1950 ; X. pavlosky Brodsky, 1955. .

‘’Xanthocalanus minor Group’’
Characters differing from the ‘’X. agilis Group’’:
Thoracic segments 4 and 5 fused.
Inner lobe 3 of Mx1 usually with 4 setae.
Endopod of Mx2 with 1 longer and 1 shorter vermifom-type setae, 5 brush-like type setae, usually with an extra short but wider brush-like seta ;
Female P5 with 2 or 3 segments, the last narrowed from the middle to the tip and with 1 or 2 terminal and 2 lateral spines ; male P5 slightly asymmetrical in length, with rudimentary endopod or none.
List of species :
Type-species : Xanthocalanus minor Giesbrecht, 1892.
Species definitely in this group : X. borealis Sars, 1900 ( = X. hirtipes van Höffen, 1897) non X. borealis Sars, 1903 (see Sars, 1921, pp.2-4 ; Sars, 1924, pl.35) ; X. medius Tanaka, 1937 ; X. penicillatus Tanaka, 1960 ; X. legatus Tanaka, 1960, X. marlyae Campaner, 1978.
Species probably in this group : X. fragilis Aurivillius, 1898 ; X. amabilis Tanaka, 1960 ; X. muticus Sars, 1905.

Total: 50 spp. (of which 1 doubtful) + 4 unident.

Diagnosis from Bradford, Haakonssen & Jillett (1983, p.70), after Bradford, 1973 :
- Rostrum usually with 2 filaments.
- Head and pedigerous segment 1 more or less separate, 4th and 5th fused or separate.
- A1 of 23-25 segments in female, fewer in male.
Mx1 inner lobe1 with 4 posterior surface setae; none of setae on inner lobe 1 modified.
- Mx2 with lobes 4 and 5 each with one of setae stout and spine-like; endopod with brush-like sensory filaments all different in size and shape, 1 usually much shorter and thicker than rest.
- Endopod of Mxp without modified setae.
- Male mouthparts with varying degrees of degeneration.
- Mx2 endopod similar to that of female or with largest brush-like sensory filament so enlarged as to almost obscure remaining filaments.
- Female P5 symmetrical, uniramous, 2- or 3-segmented and covered with spinules.
- Male P5 styliform, usually present on both sides; usually uniramous; rudimentary endopods sometimes present.

Remarks: Bradford (1973) restricted to the Phaennidae those copepods having 1 worm-like and 7 brush-like sensory filaments on the terminal part of Mx2.
Among those species placed in Xanthocalanus existing descriptions of Mx2 differ. It still remains unclear whether of not all Xanthocalanus have 1 worm-like and 7 brush-like sensory filaments.
Not all of the species to set in the genus Xanthocalanus are described well enough to either verify or reject their assignment in this genus. The following do, differ significantly from the generic definition: X. alvinae, X. difficilis, X. distinctus, X, elongatus.
Campaner (1978) suggests that X. echinatus, X. incertus, X. dilatus, X. polaris, X. pulcher, X. tenuiremis, X. giesbrechti should also be removed from Xanthocalanus. Campaner (1978) suggests that some species possibly could be assigned to Cephalophanes if the presence of frontal lenses is not considered to be diagnostic..
For Bradfird & al. (1983, p.71) several species formerly included in Xanthocalanus have been tentatively assigned to the Tharybidae: ''X''. hispidus (= ? ''X". paululus); ''X"". macrocephalon, ''X''. paraincertus.
Four other species have now been removed to other genera by Campaner (1978): X. greeni, X. maximus to Talacalanus; X. watersae to Neoscolecithrix; and X. ordinarius to Brachycalanus.

Remarks on dimensions and sex ratio:

The mean female size is 3.328 mm (n = 33; SD = 1.6572) and the mean male size is 3.177 mm (n = 39; SD = 1.2602). The size ratio (Male : Female) is 0.925 (n = 12; SD = 0.1351). The sex ratio (Female : Male) is 1.5.
Probably, we have three groups of body size: Group 1 ( sizes > 4 mm), the mean female size is 5.451 mm (n = 12; SD = 1.3874), and in male 5.043 mm (n = 10, SD = 0.3072). In the Group 2 (sizes < 3 mm), the mean female size is 2.084 mm (n = 27; SD = 0.5927), and in male 2.201 mm (n = 21; SD = 0.4189); and in the Group 3 (body sizes between 3 and 4 mm) the mean female size is 3.384 mm (n = 15; SD = 0.1857) and the mean male size is 3.421 mm (n = 7; SD = 0.3483.
The size ratio (Male : Female) is to all species with male and female: 0.925 (n = 12; SD = 0.1351).
The sex ratio (Female: Male) is 1.5.