Table Of Content© Zoological Institute, St.Petersburg, 2005
Three bathypelagic scolecitrichids new to the North Pacific,
with comments on diagnostic features of Scolecitrichidae
and Tharybidae and on the positions of Heteramalla, Rythabis
and Parkius (Copepoda: Calanoida)
N.V. Vyshkvartzeva
Vyshkvartzeva, N.V. 2005. Three bathypelagic scolecitrichids new to the North Pacific,
with comments on diagnostic features of Scolecitrichidae and Tharybidae and on the
positions of Heteramalla, Rythabis and Parkius (Copepoda: Calanoida). Zoosystematica
Rossica, 13(2), 2004: 157-180.
Additional descriptions and illustrations of the females of Heteramalla sarsi Roe, 1975,
Amallothrix aspinosa Roe, 1975 and Landrumius gigas (A. Scott, 1909), collected in the
central and northwestern Pacific for the first time, are presented. The diagnostic signifi-
cance of the setation of the maxillules, maxillae and maxillipeds in Scolecitrichidae, Thary-
bidae and Diaixidae is analyzed. The proportions of maxillule expressed as indices show-
ing length or width of its different parts in relation to the width of the first inner lobe are
proposed here as the main distinguishing characters separating Scolecitrichidae and Thary-
bidae, more informative than some diagnostic features of its setation. The relationships
between the Bradfordian families and between sections of the family Scolecitrichidae are
discussed, mainly on the basis of setation of the maxilla and maxilliped. The systematic
positions of A. aspinosa and the genera Heteramalla, Puchinia, Cenognatha, Plesioscole-
cithrix, Rythabis and Parkius are discussed. A detailed comparative analysis of charac-
ters of the genus Heteramalla is provided. Heteramalla and Puchinia belong to the group
of early offshoots of the family Scolecitrichidae, not to the central core, where they were
placed by Ohtsuka with coauthors (2003) and by Boxshall & Hulsey (2004). Rythabis has
the setation of maxillule closer to the ancestral type common to all Bradfordian families,
but the proportions of maxillule typical to Tharybidae. Since the indices of maxillule do
not overlap in Scolecitrichidae and Tharybidae, and the maxillule and other mouthparts
show different evolutionary trends of specialization, Rythabis is considered as a member
of the Tharybidae, where it was placed by Schulz (in Schulz & Beckmann, 1995). Parkius
has the proportions of the maxillule intermediate between those in Scolecitrichidae and
Tharybidae and unique apomorphic features recorded earlier by Ferrari & Markhaseva
(1996). These unique progressive features of Parkius can be a result of important biolog-
ical transformations, compared with the other Bradfordian families; therefore the status
of the separate family Parkiidae, fam. dist., is restored. The male considered as Racovit-
zanus sp. (Grice & Hulsemann, 1967) actually belongs to Parkius. Swimming legs 5 of
this male are peculiar, fitting those neither of Scolecitrichidae nor of Tharybidae. Their
structure shows a separate evolutionary trend and also justifies the family status of Parki-
idae.
N.V. Vyshkvartzeva, Zoological Institute, Russian Academy of Sciences, Universitetskaya
nab. 1, St.Petersburg 199034, Russia.
Material and methods The abbreviations used in this paper are as fol-
lows: Ce – cephalosome, Pr – prosome, SmP1-
The examined specimens were taken from SmP5 – somites bearing 1st-5th swimming legs,
plankton samples obtained during the 19th and Ur – urosome, Ur1-Ur5 – 1st-5th urosomal
39th cruises of R/V “Vitjaz” and the 40th cruise somites, A1 – antennule, A2 – antenna, Md –
of R/V “Akademik Korolev” with use of big mandible, Mx1 – maxillule (Li1 – praecoxal ar-
Judey net, mesh size 0.33 mm, and preserved in thrite or gnathobase, Li2 – coxal endite, Li3 and
4% formaldehyde. The examined specimens are Li4 – basal endites, Le1 – coxal epipodite), Mx2
kept at the Zoological Institute, St.Petersburg. – maxilla (Li1 and Li2 – praecoxal endites, Li3
The techniques of measurements, preparation of and Li4 – coxal endites, Li5 – basal endite, Wo –
slides and drawings are described by Vyshk- worm-like seta, Br – brush-like seta), Mxp –
vartzeva (2000). maxilliped (Li1-Li4 – 1st-4th endites of syncoxa),
158 N.V. Vyshkvartzeva: Scolecitrichidae and Tharybidae (cid:127) ZOOSYST. ROSSICA Vol. 13
P1-P4 – 1st-4th pairs of swimming legs, P5 – wider than long, with highly convex sides; in late-
5th pair of legs, Re1-Re7 – 1st-7th segments of ral view, it slightly deeper than long, with a large
exopod, Ri1-Ri3 – 1st-3rd segments of endopod. genital prominence; spermatheca very long, fin-
The measurements of different parts of Mx1 ger-like, directed anterodorsally and reaching
were made using the author’s original illustra- anterodorsal angle of genital double-somite (Fig.
tions, both published or unpublished, and em- 4). Genital field in ventral view (Fig. 5) with
ploying the works of previous investigators (see ovoid operculum widened distally; long vesicles
references in the text). The length of Mx1 Li1 directed anteriorly, lateral skeletal plates not visi-
was measured along the distal margin, from dis- ble. Ur2-Ur4 wider than long. Caudal rami about
tal marginal seta to the point of attachment of as long as wide. Dorsal seta II shorter than cau-
Li2. The width of Li1 and Le1 was measured from dal ramus; ventral seta VII slightly longer than
the proximal marginal seta to the distal marginal caudal ramus; proximal part of apical seta VI
seta. The length of Li2+Li3+Li4+Ri was meas- thicker than that of apical setae III-V.
ured from the point of attachment of Li2 to the A1 (Figs 6, 7) with 24 free segments, reaching
distal margin of Ri. Additionally, the length of SmP5. Md and A2 in Figs 8, 9 and 10, respec-
the longest distal marginal seta of Li1 was meas- tively. Mx1 (Fig. 11): Li1 with 7 marginal and 1
ured. The indices representing the ratios of the posterior setae, Li2-Li4 with 0, 3 and 4 setae,
following distances to the width of Mx1 Li1: (A) respectively, endopod two-segmented with 3 and
the length of Li1, (B) the length of the longest 4 setae, exopod with 10 setae, Le1 with 7 long
distal marginal seta of Li1, (C) the width of Le1, and 2 shorter setae. Mx2 (Fig. 12) with 5 setae
(D) the length of Li2+Li3+Li4+Ri, are calculat- on Li1, 3 setae on each of Li2-Li4 and 4 setae on
ed for the taxonomic groups under discussion. Li5 (all setae tapering and plumose). Endopod
of Mx2 with 3 long worm-like sensory setae dis-
TAXONOMIC ACCOUNT tally, 3 long brush-like setae with small apical
brushes and two shorter but strongly thickened
Heteramalla sarsi Roe, 1975 setae each with large globular apical brush; cen-
(Figs 1-19) tral part of each brush with numerous thin set-
ules, surrounded on periphery by longer flattened
Heteramalla sarsi Roe, 1975: 342 (341: remark); Brad- setules with fused proximal parts. Syncoxa of
ford et al., 1983: 87 (88: remark); Razouls, 1995: 346. Mxp (Fig. 13) with 4 inner lobes bearing, respec-
Heteramalla dubia: Sars, 1907: 17 (part); A. Scott, 1909: tively: 1 seta; 1 aesthetasc-like thin-walled seta
86-87 (part), pl. 33, figs 1-9; Sars, 1924: pl. 39, figs 1-
(similar to worm-like but with pointed tip); 1
16; 1925: 142-144 (part); Rose, 1933: 135 (part), fig.
short plumose + 1 long aesthetasc-like setae, simi-
130 (after Sars, 1924); ?Wilson, 1942: 69, 161, 189
(part); ?Wilson, 1950: 238, 368 (sex not indicated); lar to that of Li2; 3 plumose distal setae. Basis of
Grice & Hulsemann, 1965: 221, 223, 235; Vervoort, Mxp with 2 medial and 1 distal setae and a row
1965: 28-30 (part); ?Grice & Hulsemann, 1967: 16 (sex of spinules reaching proximal medial seta. Ri1-
not indicated). Ri6 with 2, 4, 4, 3, 3, and 2 setae, respectively;
Ri1 fused with Ri2; outer setae of Ri5 and Ri6
Material. 1 F, 14º00´N 133º00´W, R/V “Akademik absent.
Korolev”, cruise 40, depth 2610-3920 m, XI.1985; 1 F,
P1 as in Fig. 14; its basis without inner seta;
14º00´N 132º30´W, R/V “Akademik Korolev”, cruise 40,
depth 1395-2610 m, 28.X.1985; 1 F, 45°20´N 153º54´E, exopod two-segmented, endopod one-segment-
R/V “Vitjaz”, cruise 39, depth 0-4000 m, 19.VIII.1966. ed. Re1 without outer spine and inner seta; Re2
Description of female. Body length 3.41-3.78 and Re3 completely fused, compound segment
mm (n = 3). Prosome elliptical in lateral (Fig. 1) with 2 outer spines, 4 inner and 1 apical setae
and dorsal views. Forehead low, not produced (i.e., outer spine and inner seta of former Re2
compared with base of rostrum, broadly round- retained, as well as armament of Re3 typical to
ed in lateral (Fig. 2) and dorsal views. Rostrum scolecitrichids). Endopod one-segmented, with
(Fig. 3) directed ventrally, large, single, distally 3 inner and 2 apical setae and a group of spinules
divided into two strong conical processes. Ce and on posterior surface; outer lobe of endopod lack-
SmP1 separated by a thin suture, as well as SmP4 ing. P2 and P4 as in Figs 15 and 18, respectively,
and SmP5. Posterolateral corners of SmP5 not with segmentation and setation typical to scole-
produced, laterally rounded, dorsally triangular, citrichids. P3 (Figs 16, 17) with three-segment-
reaching the proximal part of genital double- ed exopod and two-segmented endopod (two dis-
somite (Fig. 4). tal segments fused; a thin suture visible on ante-
Urosome (Fig. 4) about 0.3 times as long as rior surface); outer setae of Re1-Re3 rudimenta-
prosome. Urosomites and caudal rami with fol- ry, small; apical spine of Re3 1.3 times as long
lowing relative lengths: 29:20:19:16:16=100. as segment, finely serrated. P5 (Fig. 19) minute,
Posterior margins of Ur1-Ur3 with short spinules. slightly asymmetrical, coxa fused with intercoxal
Genital double-somite in dorsal view distinctly sclerite, basis longer than wide; basis of right leg
ZOOSYST. ROSSICA Vol. 13 (cid:127) N.V. Vyshkvartzeva: Scolecitrichidae and Tharybidae 159
Figs 1-9. Heteramalla sarsi Roe, 1975, female. 1, body, right lateral view; 2, forehead, right lateral view; 3, rostrum; 4,
SmP5 and Ur, right lateral view; 5, genital double-somite, ventral view; 6, A1; 7, distal segments of A1; 8, Md; 9,
gnathobase of Md.
slightly longer than left; terminal segment of right (T. Scott, 1894), collected in the northeastern At-
leg small, with an apical spine; spine of right leg lantic close to the type locality, where female
slightly shorter than left one. specimens had been collected, according to Sars
Male unknown. (1907, 1924-1925). Roe compared Atlantic fe-
Remarks. Sars (1907) erected the genus Heter- males with Sars’ description and Atlantic males
amalla for a female which he believed, with some with T. Scott’s type specimens collected in the
doubts, to be conspecific with T. Scott’s (1894) Gulf of Guinea and found that females and males
Amallophora dubia described from male, as both previously considered as conspecific, apart from
have two much enlarged brush-like sensory se- the normal sexual differences, clearly differ in
tae on the endopod of Mx2. At the moment, such several (five) features, and thus belong not only
setae were known only in the males of the genus to different species but also to different genera.
Xanthocalanus (Phaennidae). Roe (1975) exam- Therefore, the species described from female by
ined females and males of Heteramalla dubia Sars (1907) was given a new name, Heteramalla
160 N.V. Vyshkvartzeva: Scolecitrichidae and Tharybidae (cid:127) ZOOSYST. ROSSICA Vol. 13
Figs 10-13. Heteramalla sarsi Roe, 1975, female. 10, A2; 11, Mx1; 12, Mx2; 13, Mxp.
sarsi Roe, 1975. At the same time, A. dubia Scolecitrichidae (Bradford et al., 1983; Roe,
T. Scott, 1894 has been placed to a new genus, 1975; Vyshkvartzeva, 2003).
Scopalatum Roe, 1975, because the genus Amal- Female specimens of Heteramalla sarsi, de-
lophora was not valid, as based on a male speci- scribed above from the North Pacific, generally
men which actually belongs to the genus Xantho- agree well with Sars’ (1924-1925) description of
calanus Giesbrecht, 1892. All remaining Amal- his Atlantic specimens, except that they lack a
lophora spp. were recognized as belonging to seta on Li2 of Mx1 (this feature conforms with
other new or rediagnosed genera of the family Roe’s (1975) Atlantic specimens, but possibly it
ZOOSYST. ROSSICA Vol. 13 (cid:127) N.V. Vyshkvartzeva: Scolecitrichidae and Tharybidae 161
Figs 14-19. Heteramalla sarsi Roe, 1975, female. 14, P1; 15, P2 (Re2-3 detached); 16, P3; 17, apical spine of P3; 18,
P4; 19, P5.
is a variable feature in this species) and bear one trast to this, no lobe was pictured by Sars (1924:
plumose and one aesthetasc-like seta on Li3 of Fig. 14). At last, Grice & Hulsemann (1965) and
Mxp syncoxa (this feature is described here for Roe (1975) reported that females of H. sarsi were
the first time). Sars (1924: Pl. 39, Fig. 11) did lacking P5. In our specimens, P5 is small and
not describe Mxp, but in his illustration of Mxp rudimentary, as described by Sars (1924-1925),
only one seta with a tapering tip, which is treat- A. Scott (1909) and Rose (1933), but slightly
ed here as a transformed, aesthetasc-like seta, is asymmetrical. In some other species of Scole-
shown on Li3. The other authors dealing with citrichidae, P5 is sometimes not observed (i.e.,
this species (A. Scott, 1909; Rose, 1933; Roe, Scolecithricella tenuiserrata Giesbrecht, 1892
1975; etc.: see Table 1) gave no information on and Scolecithrix bradyi Giesbrecht, 1888). It is
the morphology of Mxp. not clear if the above-mentioned absence of P5
There are some other discrepancies in the de- is a variability of the species or P5 has been bro-
scriptions of H. sarsi (Table 1). Sars (1907, 1924- ken during preparation of these minute legs.
1925), Rose (1933) and Roe (1975) noted that However, at present the conclusion should be
the endopod of P3 is two-segmented. A. Scott made that H. sarsi is a variable species.
(1909) described the endopod of P3 as three-seg- Up to now, the females of Heteramalla sarsi
mented but gave no illustration. In our specimens were known only from the Atlantic Ocean (Sars,
from the North Pacific there is a thin suture visi- 1907, 1924-1925; Grice & Hulsemann, 1965;
ble on the ventral side of endopod just distal to Vervoort, 1965), the Gulf of Lions in the Medi-
the proximal inner seta, but the outer margin of terranean Sea (Rose, 1933) and in the Obi Strait
segment opposite to the inner seta in our speci- of Indo-Malayan waters (A. Scott, 1909) (Table
mens is not pointed but straight. The proximal 1). Heteramalla dubia was recorded from the In-
endopodal segments of swimming legs in Scole- dian Ocean (Grice & Hulsemann, 1967) and the
citrichidae usually have the outer distal corner Pacific Ocean: in the Hawaiian Islands region and
produced, as on Ri1 of P2 and P3 and on Ri1-2 in the central and southeastern Pacific (Wilson,
of P4 in H. sarsi (Figs 14-18). The 2nd and 3rd 1942, 1950). However, it is not clear which spe-
endopodal segments of P3 in this species are com- cies, Heteramalla sarsi or Scopalatum dubium,
pletely fused and this compound segment has 3 were encountered here since the sex of collected
inner setae, 2 apical setae and 4 rows of spines; specimens was not indicated. So, the above-de-
the second row in our specimens consists of 4 scribed females are the first reliable and the north-
spines and is situated on a raised lobe. In con- ernmost record of H. sarsi in the Pacific.
162 N.V. Vyshkvartzeva: Scolecitrichidae and Tharybidae (cid:127) ZOOSYST. ROSSICA Vol. 13
In addition, it should be mentioned that Al- double-somite (Fig. 20), in dorsal view narrowly
zamora (1940: 6) misidentified another species rounded.
as “Heteramalla dubia”. Actually, his material Urosome (Figs 20, 31) about 0.2 times as long
fits neither Heteramalla sarsi nor Scopalatum as prosome. Urosomites and caudal rami with
dubium. following relative lengths: 33:23:18:12:14 = 100.
Posterior margins of Ur1-Ur3 with short spinules.
Amallothrix aspinosa (Roe, 1975) Genital double-somite in dorsal view (Fig. 31)
(Figs 20-36) distinctly longer than wide, parallel-sided; in late-
ral view (Fig. 23), it slightly deeper than long,
Scolecithricella aspinosa Roe, 1975: 321, Fig. 13. with genital prominence anteriorly and with con-
Amallothrix aspinosa: Vyshkvartzeva, 2000: 236 (re- cave posteroventral margin; spermatheca very
mark). long, directed anterodorsally and reaching antero-
dorsal angle of genital double-somite. Genital
Material. 2 F, 14º00´N 132º30´W, R/V “Akademik field ventrally (Fig. 24) with rectangular opercu-
Korolev”, cruise 40, depth 1395-2610 m, 28.X.1985.
lum, large rounded skeletal plates and long vesic-
Description of female. Body length 3.88 mm
les; vesicles directed anteriorly, with 2-3 bend-
(n = 2). Prosome oval and robust in lateral (Fig.
ings in distal half. Genital field with a cuticular
20) and dorsal views. Forehead low, not produced
structure looking as a thin line in shape of large
anteriorly as compared with base of rostrum,
shallow arc, curving on both sides and continu-
broadly rounded in lateral (Fig. 21) and dorsal
ed in short posteromedial shoulders; probably,
views. Rostrum (Fig. 22) consisting of two long,
this structure outlines an invagination of cuticle.
strong, sausage-shaped processes, distally con-
Ur2 as wide as long; Ur3-Ur4 wider than long.
tinuing into two short conical processes. Ce and
Caudal rami about as long as wide.
SmP1 fused; SmP4 and SmP5 separated by a thin
Both A1 broken. A2 as in Fig. 32; Md as in
suture. Posterolateral corners of SmP5 in lateral
Figs 25, 26; Mx1 as in Fig. 27; Mx2 as in Fig.
view not produced, broadly rounded, with a shal-
28; Mxp as in Fig. 29. P1 as in Fig. 33: Re1 and
low excision, reaching proximal part of genital
ZOOSYST. ROSSICA Vol. 13 (cid:127) N.V. Vyshkvartzeva: Scolecitrichidae and Tharybidae 163
Figs 20-30. Amallothrix aspinosa Roe, 1975, female. 20, habitus, right lateral view; 21, forehead; 22, rostrum; 23,
SmP5 and Ur, right lateral view; 24, genital double-somite, ventral view; 25, palp of Md; 26, gnathobase of Md; 27,
Mx1; 28, Mx2; 29, Mxp; 30, P5.
164 N.V. Vyshkvartzeva: Scolecitrichidae and Tharybidae (cid:127) ZOOSYST. ROSSICA Vol. 13
Figs 31-36. Amallothrix aspinosa Roe, 1975, female. 31, Ur; 32, A2; 33, P1; 34, P2; 35, P3; 36, P4 (34-36: two distal
segments of Re and Ri of P2-P4 detached).
Re2 without outer spine, posterior surface of Re2 exopod and also on the basis and endopod; Mx2
and Re3 with minute spinules. P2, P3 and P4 as has three wide, flattened worm-like setae on the
in Figs 34, 35 and 36, respectively; distal seg- endopod and one worm-like seta on Li5. The lat-
ments of both rami broken, cuticle in distal part ter worm-like seta is not mentioned by Roe (1975:
of coxa of P2-P4 thin. P5 (Fig. 30) apparently Fig.13h), while other three ones are different from
symmetrical, three-segmented; 2nd and 3rd seg- those figured by Roe, not wrinkled and not as
ments partially fused, 3rd segment with a small long as shown by him. However, this author not-
terminal spine and a long, thin inner seta (appar- ed that one of the paratypes had simple, flattened
ently, in this specimen broken on one of P5). worm-like setae on the endopod of Mx2. Thus,
Male unknown. there are no significant differences between the
Remarks. Females from the northwestern Pa- Atlantic and Pacific specimens.
cific agree with the description of Scolecithri-
cella aspinosa Roe, 1975 from the northeastern Landrumius gigas (A. Scott, 1909)
Atlantic, apart from the following features: the (Figs 37-42)
body is slightly larger; A2 has a short seta on Re2
(ancestral segment Re IV) and no medial seta on Brachycalanus gigas A. Scott, 1909: 81, Pl. 35, Figs 10-18.
the terminal segment; the basis of Md bears three Lophothrix gigas: Vervoort, 1965: 58; Grice & Hulse-
inner setae; the syncoxa of Mxp lacks a seta on mann, 1968: 332, Figs 63-74.
Lophothrix thorsoni Björnberg, 1975: 178, Figs 15-30.
the first inner lobe (this seta is present in Roe’s
Landrumius gigas: Park, 1983: 195, Figs 17-18.
specimen); Mx1 has a group of spinules on the
ZOOSYST. ROSSICA Vol. 13 (cid:127) N.V. Vyshkvartzeva: Scolecitrichidae and Tharybidae 165
Figs 37-42. Landrumius gigas (A. Scott, 1909), female. 37, body, dorsal view; 38, body, left lateral view; 39, distal
endopodal complex of Mx2; 40, P1; 41, P3; 42, P5.
166 N.V. Vyshkvartzeva: Scolecitrichidae and Tharybidae (cid:127) ZOOSYST. ROSSICA Vol. 13
Material. 1 F, 30º48´N 155º02´E, R/V “Vitjaz”, cruise (1983) noted individual variations in both the
19, St. 3206, depth 0-5000 m, 19.X.1954. distal end of SmP5 and the rostrum among his
Description of female. Body length 7.8 mm six specimens. The differences observed in the
(n = 1). Prosome elongate, strongly built. Fore-
present specimen also appear to be attributable
head with a short, low and somewhat flattened
to individual variations within this variable spe-
crest clearly visible dorsally (Fig. 37), ventrally cies.
and laterally (Fig. 38). Head produced anterior
In addition to the southeastern (Grice & Hulse-
to base of rostrum, in lateral view broadly round-
mann, 1968; Björnberg, 1975; Park, 1983) and
ed, in dorsal view triangular. Rostrum (Fig. 38) northern Pacific records, the species was collect-
strongly developed, somewhat plate or sausage-
ed also in the waters of the Malay Archipelago
like, with rounded tip. Line of fusion between
(A. Scott, 1909) and in Antarctic waters (Park,
Ce and SmP1 very faint, like that between SmP4 1983).
and SmP5; both visible mainly laterally. Postero-
lateral corner of SmP5 produced posteriorly as a
DISCUSSION
rounded lobe, covering anterior half of genital
double-somite, and having a small distal knob
The phylogenetic relationships of the five
on dorsal side.
Bradfordian families, i.e., Phaennidae, Scole-
Urosome about 0.19 times the length of pro-
citrichidae, Tharybidae, Diaixidae and Parkiidae
some, consisting of 4 somites. Genital double-
(the latter family is not accepted by Boxshall &
somite in lateral view as long as remaining
Halsey, 2004) need a special comprehensive ana-
somites including caudal ramus, distinctly wider
lysis. Such a study is impossible until the bound-
than long, with a large and high genital promi-
aries, composition of genera included and diag-
nence, in dorsal view longer than wide, with par-
nostic characters of these families are clarified.
allel lateral margins. Spermatheca elongate, fin-
Below, the positions of Amallothrix aspinosa
ger-like. Ur2-Ur4 each distinctly wider than long.
(Roe, 1975), Heteramalla Sars, 1907, Puchinia
Caudal rami about as long as wide, divergent.
Vyshkvartzeva, 1989, Cenognatha Bradford-
A1 broken. Mouthparts as in L. antarcticus
Grieve, 2001, Plesioscolecithrix Markhaseva &
Park, 1983, except that in our specimen 6th short
Dahms, 2004, Rythabis Schulz, 1995 and Parkius
brush-like sensory seta on endopod of Mx2 (Fig.
Ferrari & Markhaseva, 1996 will be discussed,
39) not visible, possibly due to the fact that on
and comments on the relationships and on the
permanent slide it is masked by proximal parts
diagnostic significance of some characters in the
of other five brush-like sensory setae. P1, P3 and
families Scolecitrichidae, Tharybidae and, brief-
P5 as in Figs 40, 41 and 42, respectively.
ly, in Diaixidae, will be given.
Male unknown.
The five Bradfordian families are characterized
Remarks. The female of Landrumius gigas
by the possession of sensory setae on the endo-
collected in the North Pacific is similar to that
pod and some lobes of Mx2 and on the syncoxa
described and illustrated by Park (1983) from the
of Mxp, which is a unique character state among
southeastern Pacific. It differs from the latter in
the monophyletic superfamily Clausocalanoidea.
the shape of the forehead having a somewhat flat-
These families together comprise a monophylet-
tened crest in lateral view, the posterolateral cor-
ic group (Fleminger, 1957; Ferrari & Markhase-
ners of SmP5 bearing a small distal knob, the
va, 1996). The ancient ancestor of the Bradford-
rostrum devoid of distal points (which corres-
ian families is supposed to have all setae of Mx2
ponds to the rostrum, described by Björnberg
and Mxp as normal, sclerotized setae. Nine setae
(1975)), and the absence of distal vesicle of the
on the endopod of Mx2 and three setae on Li3 of
spermatheca. Specimens from the southeastern
the syncoxa of Mxp are symplesiomorphies of
Pacific reported by Grice & Hulsemann (1968)
Scolecitrichidae (Vyshkvartzeva, 1989a, 1989b,
and Björnberg (1975) also slightly differ from
2001) and the other Bradfordian families (Ferra-
those illustrated by Park (1983) in the shape of
ri & Markhaseva, 2000; Ohtsuka et al., 2003;
the forehead (the head is very slightly produced
Boxshall & Halsey, 2004). After separation of
anteriorly in Fig. 66 in Grice & Hulsemann
the Phaennidae lineage, the ancestor common for
(1968); the crest is not indicated by Björnberg
the remaining Bradfordian families seems to re-
(1975)), in the rostrum (devoid of distal points
tain these plesiomorphic states of Mx2 and Mxp.
in Björnberg’s description) and in the longer and
The diagnostic significance of the composition
pointed posterolateral corners of SmP5, almost
of sensory setae on the endopod of Mx2 in four
reaching the distal margin of genital double-
Bradfordian families (Scolecitrichidae, Tharybi-
somite. P5 of specimens illustrated by Grice &
dae, Parkiidae and Diaixidae) is controversially
Hulsemann (1968) differs from that of our and
discussed presently. Some other authors also con-
Park’s specimens in having the inner apical spine
sider that (1) the “ninth element may be modi-
much longer than the other three spines. Park
fied into a small sensoriform seta or can remain
