Distribution of haploniscids (Isopoda, Asellota, Haploniscidae) in Icelandic waters, with description of Haploniscus astraphes n. sp. from the Iceland basin and the Southeast Atlantic Ocean

2017·3 MB·English

by  Wiebke Brökeland

Zootaxa 4231 (3): 301–326 ISSN 1175-5326 (print edition)
Article ZOOTAXA
http://www.mapress.com/j/zt/
Copyright © 2017 Magnolia Press ISSN 1175-5334 (online edition)
https://doi.org/10.11646/zootaxa.4231.3.1
http://zoobank.org/urn:lsid:zoobank.org:pub:1226613C-0001-4369-93C9-3D0B1A40BE99
Distribution of haploniscids (Isopoda, Asellota, Haploniscidae) in Icelandic
waters, with description of Haploniscus astraphes n. sp. from
the Iceland basin and the Southeast Atlantic Ocean
WIEBKE BRÖKELAND1 & JÖRUNDUR SVAVARSSON2,3
1Senckenberg am Meer Wilhelmshaven, Abt. DZMB, Südstrand 44, 26382 Wilhelmshaven, Germany. E-mail: [email protected]
2Marine Biology Centre, Institute of Biology, University of Iceland, Sturlugata 7, 101 Reykjavík, Iceland; 3The University of Iceland's
Research Centre in Suðurnes, Garðvegi 1, 245 Sandgerði, Iceland. E-mail: [email protected]
Abstract
Ten species of Haploniscidae Hansen, 1916 were sampled in Icelandic waters during expeditions in the framework of the
BIOICE project. Nine of these were known from the North Atlantic Ocean, i.e. Haploniscus aduncus Lincoln, 1985, H.
ampliatus Lincoln, 1985, Haploniscus angustus Lincoln, 1985, H. bicuspis (Sars, 1877), H. foresti Chardy, 1974, H.
hamatus Lincoln, 1985, H. spinifer Hansen, 1916, Antennuloniscus simplex Lincoln, 1985 and Chauliodoniscus armadil-
loides (Hansen, 1916). All but H. bicuspis and H. angustus were restricted to the Atlantic Ocean south of the Greenland-
Iceland-Faeroe Ridge (GIF Ridge), while H. bicuspis occurred at considerable depth ranges both north and south of the
GIF Ridge. A new species, Haploniscus astraphes n. sp., is described based on material from the Denmark Strait, North
Atlantic and the Guinea Basin, South Atlantic. H. astraphes n. sp. belongs to a group of Haploniscus species closely re-
lated to the genus Antennuloniscus and shares several characters with species from that genus, especially the spine row on
pleopod 1, the stout sensory seta on the carpus of pereopod 7 and characters of the antennae. H. astraphes n. sp. is char-
acterized by a rectangular body shape, the straight frontal margin of the head and the strongly convex posterior margin of
the pleotelson.
Key words: BIOICE, DIVA, Haploniscidae, Haploniscus, Antennuloniscus, Chauliodoniscus, Greenland-Iceland-Faeroe
Ridge
Introduction
The northernmost part of the North Atlantic Ocean is characterized by the extensive Greenland-Iceland-Faeroe
Ridge (GIF Ridge). This is the only large submarine ridge in the North Atlantic Ocean stretching in easterly-
westerly direction, separating the deeper parts of the Nordic Seas and the Arctic Ocean from the deep North
Atlantic Ocean. This ridge shapes the distribution of water masses in this area, having generally colder water
masses (often <0°C) to the north of the ridge, while warmer water masses (often >0°C) characterize the area south
of the ridge (see Stefánsson 1962; Hansen & Østerhus 2000). The colder water masses flow southward, either
along East Greenland or crossing the ridge usually in deep channels (deepest channel of 840 m; Hansen & Østerhus
2000), while the warmer water masses flow northwards usually in shallower waters.
The ridge and the associated water masses shape the distribution and diversity of benthic invertebrates in the
area (Svavarsson et al. 1993). Most of the species have their distribution either to the north or to the south of the
ridge (see data in Hansen 1916; Negoescu & Svavarsson 1997; Weisshappel 2001; Dijkstra et al. 2009; Brix &
Svavarsson 2010). The ridge may accordingly serve as barrier for dispersal of deep-water species and thus prevent
species from entering the Nordic Seas and the Arctic Ocean.
The aim of the international BIOICE project was partly to explore the distribution and species composition in
Icelandic waters north and south of the GIF Ridge. The family Haploniscidae is particularly interesting for
evaluating patterns of distribution in this area. The family is one of the most pronounced deep-sea families among
the asellote isopod crustaceans (see Svavarsson et al. 1993) currently holding 142 species (Boyko et al. 2008
Accepted by K. Meland: 8 Dec. 2016; published: 13 Feb. 2017 301
Licensed under a Creative Commons Attribution License http://creativecommons.org/licenses/by/3.0
onwards; Ahyong et al. 2011) of small isopods mostly restricted to the deep-sea. Among the many species sampled
during the BIOICE project were ten species of the Haploniscidae. One of the species was new to science while the
other nine were already known, i.e. H. aduncus Lincoln, 1985, H. ampliatus Lincoln, 1985, H. angustus Lincoln,
1985, H. bicuspis (Sars, 1877), H. foresti Chardy, 1974, H. hamatus Lincoln, 1985, H. spinifer Hansen, 1916,
Antennuloniscus simplex Lincoln, 1985 and Chauliodoniscus armadilloides (Hansen, 1916). Here we describe the
new haploniscid species and examine the distribution of the other species in the area.
Material and methods
The study area. The area is the northernmost part of the North Atlantic Ocean and the southern part of the Nordic
Seas (the Iceland, Greenland and Norwegian Seas). This area is characterized by the extensive submarine ridge
extending from Greenland to the Faeroe Islands, the Greenland-Iceland-Faeroe Ridge (GIF Ridge; often referred to
as the Greenland-Iceland-Scotland Ridge; the GIS Ridge). The deepest saddle depth of the ridge is between the
Faeroe Islands and Scotland at around 840 meters, while the deepest saddle depth in the Denmark Strait between
Greenland and Iceland is around 620 meters. Between Iceland and the Faeroes the deepest saddle depth is around
480 meters (Hansen & Østerhus 2000).
The area is characterized by several primary water masses, often overlying each other (see Stefánsson 1962;
Hansen & Østerhus 2000; Malmberg & Valdimarsson 2003), and all differing in either temperature (total range =
−0.9 to 12°C) and salinity (total range = 34.30–35.45). These are modified North Atlantic Waters (MNAW; 7.0–
8.5°C, salinity 35.10–35.30), Labrador Sea Water (LSW; 3–4°C, salinity 34.90–34.95), Iceland Sea Overflow
Water (ISOW; 2–3°C, salinity 34.85–35.00), Modified East Icelandic Water (MEIW; 1–3°C, salinity 34.70–34.90),
Norwegian Sea Deep Water (NSDW; <−0.5°C, salinity <–34.90), Norwegian Sea Arctic Intermediate Water
(NSAIW; −0.5–0.5°C, salinity 34.85–34.90), Arctic/Polar Water (A/PW; 0–2°C, salinity 34.30–34.90) and Coastal
Water (CW; 5–12°C, salinity <34.50).
Additional material of Haploniscus astraphes n. sp. was sampled during the DIVA 2 expedition (DIVA:
Latitudinal Gradients of Deep-Sea BioDIVersity in the Atlantic Ocean) in the Guinea and Angola Basins in the
southeast Atlantic Ocean in 2005. The major bottom water mass in these basins is the North Atlantic Deep Water
(NADW) which partly mixes with Circumpolar Deep Water (CDW) passing through the Romanche and Chain
Fracture Zones in the Mid-Atlantic Ridge.
Sampling, identifying and measuring. The BIOICE samples were taken with a modified Rothlisberg-Pearcy
epibenthic sled (RP sled; Rothlisberg & Pearcy 1977, Brattegard & Fosså 1991), a Sneli sled (Sneli 1998), a
Triangle dredge and an Agassiz trawl. The RP sled samples were elutriated through a 0.5 mm sieve, and then the
remaining sediment and animals were sieved through a series of sieves (4, 2, 1 and 0.5 mm) and preserved in 5%
buffered formalin. The Sneli sled samples were sieved through 2, 1 and 0.5 mm sieves, while the Agassiz samples
and the Triangle dredge samples were hand-picked on deck. Most specimens were collected in the RP sled. The
DIVA samples were taken with a Brenke sledge (Brenke 2005) and preserved in ethanol having been sorted upon
arrival at the deck.
The specimens were identified, measured and dissected using a Leica MZ APO12.5 or a Leica MZ9.5
dissecting microscope, drawings were prepared using Leica compound microscopes (DMLS2, DM2500) equipped
with camera lucida.
Body length was measured in lateral view along the midline from the anterior margin of the head to the
posterior margin of the pleotelson. Body segment and limb article lengths were also measured along the midline.
Other measurements refer to the greatest length/width of an article or segment. The differentiation in peduncular
and flagellar articles of the antennae follows Lincoln (1985a) and Wägele (1983). Allocation to ontogenetic stages
follows Brökeland (2010). Examined DIVA specimens are deposited at the Zoological Museum Hamburg (ZMH),
while the BIOICE specimens of Haploniscus astraphes and specimens of the other haploniscid species are
deposited at the Icelandic Museum of Natural History (IMNH).
Taxonomy
Family Haploniscidae Hansen, 1916
302 · Zootaxa 4231 (3) © 2017 Magnolia Press BRÖKELAND & SVAVARSSON
Genus Haploniscus Richardson, 1908
Haploniscus astraphes n. sp.
(Figs 1–9)
Holotype: DIVA2-HA307, male stage V, 1.01 mm, RV Meteor, M63/2, DIVA 2 expedition, EBS station 89-6,
0°42.95’N, 5°31.29’W, 5142 m, Guinea Basin, southeast Atlantic, ZMH K-42450.
Paratypes: DIVA2-HA308, 1 manca stage III, same locality as holotype, ZMH K-42451; DIVA2-HA515, 1
male stage V, 1.04 mm, DIVA2-HA414, 1 female stage IV or V, 1.06 mm, DIVA2-HA415, 1 manca stage III, 0.92
mm, EBS station 90-7, 0°40.49’N, 5°29.71’W, 5142 m, Guinea Basin, ZMH K-42502; DIVA2-HA153, 1 female
stage IV or V, 1.24 mm, EBS station 45-3, 9°53.76’S, 0°53.60’E, 5657 m, Angola Basin, ZMH K-42503.
BIOICE station 2909, RV Bjarni Sæmundsson, North Atlantic: 2 females stage IV or V, 1.25 and 1.09 mm, RP sled,
65°15.61’N, 28°50.15’W, 1300 m, 25 August 1996, 3.96°C, Denmark Strait, south of Greenland-Iceland Ridge,
IMNH 10370.
Etymology. The Greek astraphes, “straight”, refers to the straight outlines of head and pereon in this species.
Diagnosis. Body rectangular. Head rectangular in males, trapezoidal in females. Vertex straight, without
rostrum. Clypeus slightly prolonged, dorsally keeled. Pleotelson terminal margin strongly convex. Antenna 2
peduncular article 1 small, article 3 with long tooth, curved backwards, articles 5 and 6 short, not fused, article 6
with strong apical projection, flagellum small with 8–9 articles. Mandibular palp article 3 with 6 pectinate setae.
Maxilliped with 2 retinaculae. Outer apical comb on carpus of pereopods 4–6 composed of large spines, much
larger than inner comb; dorsal sensory stout seta on carpus of pereopod 7; distalmost ventral scale on propodus
drawn out towards dactylus. Pleopod 2 of males with short stylet of the endopod, operculum of females nearly
circular. Pleopod 3 exopod with 3 simple setae. Uropods stout, ramus length about 1.8 width.
Description of male (stage V). Body (Fig. 1) length 3.3 width, rectangular, lateral margins nearly parallel.
Head length 0.7 width, rectangular; vertex straight, without rostrum; anterolateral margins slant in lateral view;
clypeus slightly prolonged, keeled. Pereonites 4 and 5 longest, pereonite 7 shortest; anterolateral angles of
pereonites 2–5 acute. Pleotelson as wide as pereonite 7 basally, length 1.1 width, lateral margins convex, slightly
concave distally; terminal margin strongly convex; pleotelson processes short, slightly exceeding terminal margin;
dorsal surface with two indistinct keels, ventral surface with faint cuticular ledge around branchial chamber.
Antenna 1 (Fig. 4A) length 0.3 body length, peduncular article 1 broadest, length 1.6 width, with 2 penicillate
setae distally; article 2 length 0.9 article 1 length, 2.2 width, with 1 simple and 4 penicillate setae distally; article 3
length 0.4 article 1 length, width 0.4 article 1 width, with 1 simple seta distally. Flagellum with 4 articles; article 1
shortest, with 2 penicillate setae distally; article 2 length 2.0 article 1 length, with 1 simple seta, article 3 length
1.45 article 1 length, with 1 aesthetasc and 1 simple seta, article 4 length 2.3 article 1 length, with 2 aesthetascs, 1
penicillate seta and 3 simple setae.
Antenna 2 (Fig. 4B) length 0.6 body length, peduncular article 1 minute (lost during dissection), article 2
longer, with 1 simple seta; article 3 length 1.7 article 2 length, 1.9 width, with 3 simple setae, dorsal tooth slightly
longer than article width, curved backwards, with at least one simple seta; article 4 length 0.8 article 2 length, as
wide as long, with 1 simple seta; article 5 short, length 1.1 article 2 length, 1.4 width, with about 4 simple setae in
distal half; article 6 short, length without tooth 1.1 article 2 length, 1.65 width, with at least 3 penicillate setae and
about 11 simple setae, distal tooth length about 0.7 article length. Flagellum small, total length 2.6 peduncular
article 6 length, with 9 articles, decreasing in length and width from articles 1–7, articles 8 and 9 slender; with up to
5 simple setae per article.
Mandible (Figs 5A, B) incisor with 5 cusps; lacinia mobilis of left mandible with 5 teeth; spine row with 2
serrated and 2 setulated spines, spine row of right mandible with 3 serrated and 3 setulated spines. Molar with tooth
row and opposing cuticular ledge indistinctly separated into teeth, 4–5 strong setae proximally of tooth row,
accessory teeth row proximally of cuticular ledge and 2–3 setae on molar trunk. Palp slightly shorter than
mandibular body, article 1 with 1 simple distal seta, article 2 with 3 pectinate setae distally, article 3 with 6
pectinate setae along inner margin.
Maxilla 1 (Fig. 6A) lateral lobe length 3.5 width, distal margin with 12 stout setae, outer margin with rows of
slender setae, inner margin with 2 setae distally, surface with few scattered rows of small setae; mesial lobe length
about 0.7 outer lobe length, width 0.6 outer lobe width, distal margin with 2 stout 4 long and several smaller setae,
surface and outer margin with several slender setae.
HAPLONISCIDS IN ICELANDIC WATERS Zootaxa 4231 (3) © 2017 Magnolia Press · 303
FIGURE 1. Haploniscus astraphes n. sp. male holotype, ZMH K-42450. A, habitus, dorsal view; B, habitus, lateral view; C,
posterior body, ventral view. Scale bar = 1mm.
304 · Zootaxa 4231 (3) © 2017 Magnolia Press BRÖKELAND & SVAVARSSON
FIGURE 2. Haploniscus astraphes n. sp. female paratype, ZMH K-42503. A, habitus, dorsal view; B, habitus, lateral view; C,
posterior body, ventral view. Scale bar = 1mm.
HAPLONISCIDS IN ICELANDIC WATERS Zootaxa 4231 (3) © 2017 Magnolia Press · 305
FIGURE 3. Haploniscus astraphes n. sp. female paratype, ZMH K-42503. Head, A, ventral view; B, lateroventral view. Scale
bar = 0.1mm.
Maxilla 2 (Fig. 6B) lateral and middle lobes subequal in length and width, length about 5.5 width; mesial lobe
width 1.8 middle lobe width, length 1.3 middle lobe length, 3.9 width. Proximal inner margin of mesial lobe with
about 3 long setae; surface and distal inner margin with rows of stout and slender setae; distal margin with 2 short
serrated stout setae and 5 strong setae. Lateral and middle lobe each with 1 small stout seta and 2 long and 1 shorter
serrated stout setae distally. Inner margin of middle endite with several strong setae, outer margin of lateral endite
with numerous slender setae.
Pereopods (Figs 7, 8): Basis with 1–2 long simple setae dorsally and 1 simple seta ventrally (probably broken
off on pereopod 7); basis of pereopods 2–7 with 1 penicillate seta. Ischium of pereopods 1–4 with 1 seta, ischium
of pereopods 5–7 with 2 setae ventrally. Merus with 2 setae distoventrally and distodorsally each, merus of
pereopod 6 with 1 seta on ventral margin. Ventral margins of carpus, propodus and dactylus fringed by comb-like
scale rows, distalmost scale on propodus drawn out towards dactylus; surface of carpus and propodus with some
scale rows. Carpus of pereopod 1 with 3 setae ventrally, carpus of pereopods 2–7 with 2 setae ventrally; distal
carpus with 1 simple seta dorsally, distal carpus of pereopods 2–7 with 1 penicillate seta (probably broken off on
pereopod 3) dorsally, carpus of pereopod 7 with one very stout sensory seta; apical combs of peropods 2–7 of
different size, outer comb distinctly larger, more spinose than inner comb, outer combs of pereopods 4–6 composed
of very large spines. Propodus with 2–4 setae ventrally, pereopods 1–3 with 1 long simple seta distodorsally, seta
shorter on remaining pereopods (broken off on pereopod 4), pereopods 2–7 with penicillate seta distodorsally
(broken off on pereopods 4 and 6); with small apical comb. Dactylus with 2–5 setae near insertion of unguis and
triangular accessory claw.
Pleopod 1 (Fig. 9A) length 0.8 ventral pleotelson length, 1.85 width, neck (narrowest part) width 0.5
maximum width. Lateral margins concave, distal margins rounded, smooth, with some setae.
Pleopod 2 (Fig. 9B) basipod semicircular, length 1.8 width, distal two thirds of lateral margin with long setae,
endopod short, not reaching tip of basipod.
Uropod (Fig. 1C) stout, ramus length 1.8 width, reaching terminal margin of pleotelson.
Description of female (stage IV or V). Body (Figs 2, 3) length 3.2 width. Head trapezoidal, tapering towards
vertex; pereonite 7 and pleotelson slightly narrower than pereonite 6. Pleotelson length 1.3 width, lateral margins
convex basally, concave distally; terminal margin strongly convex; pleotelson processes short, reaching terminal
margin; dorsal surface with two indistinct keels.
Antenna 1 (Fig. 4D) peduncular article 1 with 3–4 penicillate setae distally. Flagellum with 3 articles; article 1
shortest, with 2 penicillate setae distally; articles 2 and 3 of subequal length, each 2.0 article 1 length, article 2 with
1 simple seta, article 3 with 2 aesthetascs, 1 penicillate seta and 1 simple seta.
306 · Zootaxa 4231 (3) © 2017 Magnolia Press BRÖKELAND & SVAVARSSON
Antenna 2 (Fig. 4C) peduncular article 1 minute, article 2 with 2 simple setae; article 3 length 1.55 article 2
length, 2.25 width, with 1 simple setae, dorsal tooth slightly longer than article width, with at least one simple seta;
article 4 length 1.3 width; article 5 length 0.8 article 2 length, 1.2 width, with about 2 simple setae distally; article
6, without tooth as long as article 2, length 2.1 width, with at least 2 penicillate setae and 7 simple setae. Flagellum
total length 2.3 peduncular article 6 length, with 8 articles, articles 2–8 of subequal length, decreasing in width,
more slender than in male; with up to 4 simple setae per article.
Maxilliped (Fig. 6C) endite apical margin with at least 2 small fan setae, several small stout setae and simple
setae; apical medial margin with 1 serrated and 1 simple stout seta dorsally and a row of strong setae on ventral
surface; few simple setae on dorsal surface; 2 retinaculae. Epipod slightly longer than endite.
Pleopod 2 (Fig. 9C) nearly circular, shoulders indistinct, slightly wider than long, ventral suface smooth with
shallow depression basally, lateral margins with 1 seta each, distal margin with 6 setae.
Pleopod 3 (Fig. 9D) endopod length 1.65 width. Exopod length 0.4 endopod length, 1.2 width, with 3 simple
setae.
Pleopod 4 (Fig. 9E) endopod length 1.4 width. Exopod length 0.45 endopod length, 3.1 width, plumose seta
length about 2.5 exopod length.
Pleopod 5 (Fig. 9F) length 2.1 width.
Remarks. Only a few specimens of Haploniscus astraphes n. sp. were found in both localities, the Guinea and
Angola Basins and the North Atlantic (Fig. 10A, dot), and the two specimens from the Denmark Strait were both
juvenile females. With such a long distance and a depth difference of about 3800 m between the localities it might
seem doubtful that the specimens from the North Atlantic Ocean actually belong to the same species as those from
the South Atlantic Ocean. Examination of the available specimens revealed no morphological differences, as to
justify the erection of a separate species. Therefore the specimens from the North Atlantic are allocated to H.
astraphes n. sp. until more material allows the analysis of further characters.
H. astraphes n. sp. strongly resembles species of the genus Antennuloniscus Menzies, 1962. The latter genus
has been characterized by the following apomorphies: clypeus strongly prolonged, article 1 of antenna 2 small,
articles 5 and 6 of antenna 2 fused (suture may be visible), and pleopod 1 with spine row near distal end of
transverse groove (Brökeland 2006).
H. astraphes n. sp. is similar in the shape of the head and body to Antennuloniscus simplex Lincoln, 1985,
which also occurs in Icelandic waters. This latter species is different from other Antennuloniscus species in having
the clypeus less prolonged and the antenna 2 articles 5 and 6, while definitely fused, are separated by a more
distinct suture than in most other species of the genus. The flagellum of antenna 2 is, however, smaller in A.
simplex than in H. astraphes and has only six articles, while the flagellum of H. astraphes has eight or nine articles.
At least three other species of Haploniscus, i.e. H. rostratus (Menzies, 1962), H. ingolfi Wolff, 1962, and H.
pygmaeus Birstein, 1969, seem to be closely related to Antennuloniscus (Brökeland 2010). Together with H.
astraphes n. sp. they share some characters with the Antennuloniscus species, such as article 3 of antenna 2 being
longer than wide, article 6 of antenna 2 having a large terminal projection and therefore the flagellum being
inserted subapically. An interesting feature is the location of the dorsal sensory stout seta on the carpus of pereopod
7 instead of pereopod 6, where it is located in the majority of haploniscid species. As far as could be determined
from the species descriptions, Antennuloniscus species also have the sensory stout seta on pereopod 7, as seen in H.
ingolfi (but not H. rostratus). Adult males of H. rostratus have the typical spine row on pleopod 1 (see Brökeland
2010).
However, the similarities between H. astraphes n. sp. and Antennuloniscus also include several other
characters, which are found in many or even all species of Antennuloniscus. One of these is the small article 1 of
antenna 2, formerly an autapomorphy of Antennuloniscus. The clypeus of H. astraphes n. sp. is slightly prolonged
(Fig. 3), but not as much as in most Antennuloniscus species. The armature of the pereopods is similar, with the
distalmost ventral comb on the propodus drawn out towards the dactylus and one of the apical combs on the carpus
composed of large spines. Within Antennuloniscus, preserved specimens often have antenna 1 directed posteriorly
or laterally and therefore ventrally of antenna 2. A similar position of antenna 1 could be observed in some
specimens of the new species. The proportions of articles 5 and 6 of antenna 2 are similar to those of most
Antennuloniscus species. Like H. astraphes n. sp. many Antennuloniscus species have only two retinaculae on the
maxilliped, in contrast to the usual three in most other haploniscids. Since no adult males of H. astraphes n. sp.
were found, similarities of pleopod 1 remain unclear.
HAPLONISCIDS IN ICELANDIC WATERS Zootaxa 4231 (3) © 2017 Magnolia Press · 307
FIGURE 4. Haploniscus astraphes n. sp. male paratype, ZMH K-42502 (A, B), female paratype, ZMH K-42502 (C, D). A,
antenna 1; B, antenna 2; C, antenna 2; D, antenna 1. Scale bar = 0.1mm.
308 · Zootaxa 4231 (3) © 2017 Magnolia Press BRÖKELAND & SVAVARSSON
FIGURE 5. Haploniscus astraphes n. sp. male paratype, ZMH K-42502 (A, B), female paratype, ZMH K-42502 (C, D). A,
right mandible; B, left mandible; C, right mandible; D, left mandible. Scale bar = 0.1 mm.
HAPLONISCIDS IN ICELANDIC WATERS Zootaxa 4231 (3) © 2017 Magnolia Press · 309
FIGURE 6. Haploniscus astraphes n. sp. male paratype, ZMH K-42502 (A, B and E, F), BIOICE female paratype, IMNH
10370 (C). A, maxilla 1; B, maxilla 2; C, maxilliped; D, maxilliped, detail of distomedial margin of endite; E, maxilliped,
retinaculae. Scale bars: A, B = 0.5 mm; C = 0.1 mm; D, E = 0.01 mm.
310 · Zootaxa 4231 (3) © 2017 Magnolia Press BRÖKELAND & SVAVARSSON