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Accepted by G. Bird: 25 Aug. 2015; published: 17 Sept. 2015
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN
1175-5334
(online edition)
Copyright © 2015 Magnolia Press
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http://dx.doi.org/10.11646/zootaxa.4018.4.3
http://zoobank.org/urn:lsid:zoobank.org:pub:A297C919-7B2B-482E-A3FD-97CD53C9B3EB
Two new species of family Neotanaidae (Peracarida: Tanaidacea) from the
Antarctic and Mid-Pacific Oceans
CATARINA L. ARAÚJO-SILVA
1,2,4
, ELSA FROUFE
1
& KIM LARSEN
3
1
CIIMAR (Interdisciplinary Centre of Marine and Environmental Research), AEE (Laboratory of Aquatic Ecology and Evolution),
Porto University, Rua dos Bragas, 289, 4050-123 Porto, Portugal.
2
Capes Foundation, Ministry of Education of Brazil, Brasília–DF, 70040–020 Brazil.
3
ENSPAC Department of Environmental, Social and Spatial Change, 11.2, Roskilde University Center. DK-4000 Roskilde, Denmark.
4
Corresponding author (E-mail: katelas@hotmail.com)
Abstract
Samples collected from the Antarctic (ANDEEP/2002) and Mid-Pacific (
BIONOD/2012
) Oceans allowed analyses of
several specimens of the family Neotanaidae. From these surveys two new species are described: Neotanais bicornutus
and Venusticrus thor. The new material led to a re-diagnosis of Venusticrus, and N. rotermundiae is now assigned to this
genus. The male of N. bicornutus shares a number of characters with the "robustus" species group, but differs by having
a pleotelson about 1.5 times as wide as long, cheliped carpus about 1.5 times as long as cephalothorax, cheliped propodus
with two long dorsal projections, and uropod endopod article 1 with 8–10 fine setae proximal to mid-length on outer mar-
gin. The N. bicornutus preparatory female differs from all species by a combination of characters including the number of
setae on dorsal margin of cheliped carpus (about 15 setae), the uropod attachment slightly posterior to mid-length, uropod
basal article about 2.7 times as long as endopod article 1. The female of V. thor differs from those of V. insolitus, V. glan-
durus and V. rotermundiae by the body proportions, the pleon having three lateral setae on epimera, pleon with a blunt
ventral keel, pereopodal setation, number of setae on maxilliped endite and basis as well as other characters. Total genomic
DNA was extracted from two specimens of V. t h o r and sequences of two genes, i.e., cytochrome oxidase 1 (COI) and ri-
bosomal (28S) were obtained.
Key words: Neotanais, Venusticrus, COI, 28S, ANDEEP, Antarctic, Mid-Pacific
Introduction
The superfamily Neotanaoidea Sieg, 1980 consists of one family, Neotanaidae Lang, 1956 and is considered one of
the largest tanaidacean families, comprised of 51 (with two described herein) species (Anderson 2013; Wi et al.
2014, 2015). This family is restricted to deep water and currently holds four genera (Carololangia Gardiner, 1975;
Herpotanais Wolff, 1956a; Neotanais Beddard, 1886; Venusticrus Gardiner, 1975). Neotanaids are geographically
widespread with 24 species originally described from the Pacific and two, four and 19 from the Indian, Antarctic
and Atlantic oceans, respectively. Despite the distributional separation, the morphology of the females is highly
conservative while the males are polymorphic. This might suggest gene flow or genetic lag between populations
(Gardiner 1975) which potentially could be attributed to the higher mobility of this taxon relative to
Tanaidomorpha (Larsen 2005).
Before the type genus Neotanais was elevated to the family rank by Lang (1956), the genus was first included
within the Tanaididae (Beddard, 1886) and later Paratanaidae (Lang, 1949). Afterwards, Sieg (1980) created the
suborders Apseudomorpha, Neotanaidomorpha and Tanaidomorpha; he considered Neotanaidomorpha had too
many shared characters with both Apseudomorpha and Tanaidomorpha to be placed within either and elevated the
taxon to suborder rank.
Recently, Kakui et al. (2011) presented a phylogenetic study of the Tanaidacea based on a molecular analysis.
Their results showed that the suborder Neotanaidomorpha nested within Tanaidomorpha, with strong support, and
it was reduced to superfamily rank. This result conflicts with previous phylogenies based on morphological
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characters (Larsen & Wilson 2002). Also the general life-style between neotanaids and tanaidomorphans differs
considerably e.g., neotanaids are free-living surface dwellers (Gardiner 1975; Thistle et al. 1985) while
tanaidomorphans are mostly, if not exclusively, tube dwellers (Johnson & Attramadal 1982a, b; Mendoza 1982;
Holdich & Jones 1983).
This study aims to analyse morphologically several neotanaid specimens and describe one new species of
Neotanais collected in the Antarctic (ANDEEP/2002) and one of Venusticrus, sampled from Manganese Nodules-
Pacific Ocean (BIONOD/2012). In addition, mtDNA COI and r28S genes were sequenced from two Venusticrus
specimens and submitted to GenBank.
Material and Methods
Taxonomy
The specimens of Neotanais bicornutus sp. nov. (see below) were collected from the survey ANDEEP/2002
(Antarctic) and Venusticrus thor sp. nov. (see below) from BIONOD/2012 (Mid-Pacific) (for information on that
material see Larsen & Araújo-Silva (2014a, b).
Terminology follows Larsen (2003). Dissections were made with chemically sharpened tungsten wire needles
and then placed on slides with glycerin and chlorazol black, covered by a cover slip and sealed with nail polish.
Habitus illustrations were made from holotypes and allotypes while appendages were dissected and drawn from
paratypes via a camera lucida attached to an Olympus compound microscope. Drawings were made with the aid of
the computer program CorelDraw. Type material of N. bicornutus is kept at the Natural History Museum,
Hamburg, Germany (DZMB ZMH) while V. t h o r at the Muséum National d’Histoire Naturelle Paris, France
(MNHN).
Molecular
Total genomic DNA was extracted from two specimens of V. t h o r and sequences of two genes, i.e., cytochrome c
oxidase subunit 1 (COI) and 28S ribosomal RNA (28S) were obtained. Primers used, PCR reactions, and
sequencing conditions for both genes are described in Larsen et al. (2014), except for the DNA template volumes
that were 1.0 μL. The cycle parameters are as follow: initial denaturation at 94°C (3 min), denaturation at 94°C (30
s), annealing temperature at 45°C (COI) and 46ºC (28S) both for 1 min and extension at 72°C (1 min 30s) repeated
for 44 cycles, with a final extension of 10 min at 72°C. Sequences were read on an ABI-310 and are available on
GenBank (accession numbers: 28S - KT592230, KT592231; COI - KT592232, KT592233).
Systematics
Order Tanaidacea Dana, 1849
Suborder Tanaidomorpha Sieg, 1980
Superfamily Neotanaoidea Sieg, 1980
Family Neotanaidae Lang, 1956
Genus Neotanais Beddard, 1886
Diagnosis. For females, see Weigmann & Guerrero-Kommritz (2009: 21)
Gender. Masculine
Type species. Neotanais americanus Beddard, 1886
Generic remarks. According to Larsen (2005) and Bamber (2007) the following combination of characters
have been useful to distinguish the species (females): relative length/width (l/w) of cephalothorax, pereonites,
pleotelson, cheliped propodus, antennae, and uropods; the shape of pleonal ventral keel; number of cheliped dorsal
carpal setae; the l/w of the cheliped fixed finger and dactylus; proportions of article 1 and relative lengths of the
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distal articles of the antennule; relative length of the uropod exopod with the first article of the uropod endopod and
the adult size.
Neotanais bicornutus sp. nov.
(Figs 1–3)
Material examined. Holotype: female with oostegites (ZMHK 45086), body total length: 9.7 mm, ANT XIX-3
ANDEEP I Sta. 42-2/EBS-SUPRA, 59º 24’ 10’’ S, 57º 21’ 15’’ W, 27 January 2002, depth 3683 m.
Allotype: copulatory male (damaged, partly dissected—ZMHK 45088), body total length: 20.6 mm, ANDEEP
II Sta. 140-8/EBS-Supra, 58° 9’ 35’’ S, 24° 32’ 14’’ W, 22 March 2002, depth 2947 m.
Paratype: one female without oostegites, body total length: 5.21 mm (dissected, body partially preserved -
ZMHK 45087), same locality of allotype.
Etymology. The name reflects the presence of two projections found on the dorsal male chelipedal propodus,
similar to a "cornus”, i.e. horn, in Latin.
Diagnosis. Female, with oostegites: body six times as long as wide. Cephalothorax marginally longer than first
two pereonites combined, no apparent rostrum or setae. Pereonites rectangular, without setae. Pleonites narrower
than pereon and pleotelson, pleon lateral epimera with three plumose setae, mid-ventral keel very weekly
pronounced. Pleotelson 0.6 times as long as wide, no setation. Antennule as long as cephalothorax, article 1 about
3.3 times as long as wide. Maxilliped basis broad, without proximal seta, endite with three spiniform and one
bipinnate inner distal setae. Cheliped carpus with two simple ventral setae and about 15 simple dorsal setae.
Uropod attachment slightly posterior to mid-length, basal article about 2.7 times as long as article 1 of endopod,
exopod biarticulate, marginally shorter than endopod article 1.
Male, copulatory male: larger than female, about 7.2 times as long as wide. Body with no apparent setae.
Cephalothorax about 1.5 times as long as the first two pereonites combined. Pereonites rectangular, without setae.
Pleon and pleotelson naked, uropod attached mid-length on pleotelson. Antennule 0.8 times as long as
cephalothorax, article 1 about 7.5 times as long as wide. Maxilliped basis robust, naked. Endite naked. Palp of four
slender articles with one simple seta on article 3 and four simple distal setae on article 4. Cheliped sclerite, basis,
ischium and merus naked; carpus about 1.5 times as long as cephalothorax, widening distally with one large ventral
protuberance bearing one strong spine; propodus with two long dorsal projections; fixed finger with an inner-
medial deep groove and a tooth distally. Uropod endopod article 1 with 8–10 fine setae proximal to mid-length on
outer margin.
Description. Female. Body based on female holotype and appendages from dissected paratype. Body (Fig. 1A,
B): heavily calcified, dorsoventrally flattened, about six times as long as wide. Cephalothorax: about 1.2 times as
long as wide and 1.1 times as long as the first two pereonites combined, no apparent rostrum or setae. Pereon:
about 4.1 times as long as wide, 60% of total body length, no dorsal relief. All pereonites rectangular, no plumose
or simple setae; pereonite 1 shortest, 0.4 times as long as wide; pereonites 2–6 subequal: 0.7, 0.76, 0.81, 0.87, 0.68
times as long as wide, respectively. Pleon: about 1.8 times as long as wide, 20% of total body length; all pleonites
subequal, narrower than pereon and slightly narrower than pleotelson, lateral epimera with about three plumose
setae, mid-ventral keel very weekly pronounced (Fig. 1D). Pleotelson 0.6 times as long as wide, no apparent
setation; uropod attachment slightly posterior to mid-length.
Antennule (Fig. 2A): as long as cephalothorax. Article 1 longest, about 3.3 times as long as wide, with one
setulose and four simple distal setae. Article 2 about three times shorter than article 1, with four simple distal setae.
Article 3 about 1.2 times as long as wide, 1.6 times shorter than previous article, with two simple distal setae.
Article 4 1.2 times as long as article 3 and three times as long as wide, naked. Articles 5–7 marginally shorter than
article 4, with one simple and one annulated aesthetasc on article 5; article 6 with one annulated aesthetasc; article
7 with three simple distal setae.
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FIGURE 1. Neotanais bicornutus sp. nov., female with oostegites (ZMHK 45086): A, holotype habitus dorsal view; B,
holotype habitus lateral view. Female paratype (ZMHK 45087): C, cheliped; D, pleon and pleotelson; E, pleopod. Copulatory
male allotype (ZMHK 45088): F, habitus dorsal view; G, pleon; H, cheliped; I, pleotelson and uropod. Scale bars: A, B, F, H =
2 mm; C = 0.5 mm; D, E = 0.2 mm; G, I = 1 mm.
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FIGURE 2. Neotanais bicornutus sp. nov., female paratype: A, antennule; B, antenna; C, labrum; D, left mandible; E, right
mandible; F, maxillule, outer endite; G, maxillule, inner endite; H, labium; I, maxilla; J, maxilliped; K, uropod; copulatory male
allotype: L, antennule and antenna; M, maxilliped. Scale bars: A–J = 0.2 mm; K = 0.5 mm; L, M = 2 mm.
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Antenna (Fig. 2B): marginally shorter than antennule (0.8 times). Article 1 broader than others articles, about
1.6 times as long as wide and, as long as articles 3 and 4 combined, naked. Article 2 slender than article 1, about 2.6
times as long as wide, with two simple distal setae. Article 3 as long as wide and, as long as articles 7–8 combined,
0.4 times as short as article 2, with one simple distal seta. Article 4 about two times as long as wide, naked. Article
5 marginally longer than article 2, about 3.3 times as long as wide, with two simple distal setae. Article 6 about 1.2
times as long as article 3, naked. Articles 7 and 8 both short, with three and five simple distal setae, respectively.
Mouthparts (Fig. 2C–J): labrum (Fig. 2C) subtriangular, with several fine setules at the tip, anterior lateral
margins depressed. Mandibles (Fig. 2D, E) molar with several distal pointed denticles. Left mandible (Fig. 2D)
incisor longer than lacinia mobilis, with several blunt denticles, setal row with two spiniform- serrated setae;
lacinia mobilis well developed with four crenulated denticles. Right mandible (Fig. 2E) incisor with several blunt
crenulated denticles, setal row with four spiniform- serrated setae. Labium (Fig. 2H) palp elongated with several
setules at tip and two small spiniform setae, lobes with distal and lateral rows of setules. Maxillule (Fig. 2F, G)
outer endite (Fig. 2F) with eight spiniform distal setae as well as several fine lateral setae; inner endite (Fig. 2G)
one long and two short bipinnate setae and one short spiniform seta. Maxilla (Fig. 2I) inner lobe of fixed endite
with several distal setae; outer lobe with two spiniform setae; inner lobe of movable endite with three spiniform
setae, outer lobe with two spiniform and three bifurcate spiniform setae.
Maxilliped (Fig. 2J) endite with three inner distal spiniform setae, one bipinnate spiniform and several fine
setae along the distal margin. Basis broad, palp pedestal (see Larsen 2005: 11) with one long simple distal seta.
Palp article 1 naked, as long as article 3; article 2 shortest, with one outer distal simple seta and six inner simple
setae; article 3 about 1.4 times as long as article 2, with eight simple inner setae; article 4 slender and longest than
the other articles, four times as long as wide, with three pinnate and four simple setae. Epignath not recovered.
Cheliped (Fig. 1C): sclerite rectangular (illustrated on Fig. 1B). Basis tapering distally, about 1.1 times as long
as wide, naked. Ischium very short, hardly visible. Merus with one simple seta. Carpus about 1.1 times as long as
basis, about twice as long as wide; with two simple ventral setae and, 15 simple setae along dorsal margin.
Propodus slightly shorter than basis, no setae. Fixed finger with one simple inner proximal seta, three distal and
two on outer margin next to unguis; dactylus shorter than fixed finger with two short spines in the inner margin.
Pereopod 1 (Fig. 3A): coxa small, naked. Basis slender, about 4.1 times as long as wide, slightly longer than
the three next articles combined, with four dorsoproximal and two plumose ventral setae. Ischium with two simple
setae. Merus about 1.5 times as long as carpus, with two bipinnate medial setae. Carpus shortest, about 1.7 times as
long as wide, with three simple medial setae, one spiniform- serrated distal and, three long- bipinnate distal setae.
Propodus as long as dactylus and unguis combined, about 2.9 times as long as wide, with one setulose medial, two
simple distal, one spiniform- serrated hook-like, and four spiniform- bipinnate setae. Dactylus and unguis thin,
naked.
Pereopod 2 (Fig. 3B): articles with similar length and armament as pereopod 1 except: basis with two ventral
and one setulose dorsal setae. Carpus about 0.8 times as long as merus, with four simple medial setae. Propodus
with seven spiniform- bipinnate ventral setae. Dactylus with one fine simple proximal seta; unguis broken.
Pereopod 3 (Fig. 3C): articles with similar length to pereopod 2 except: coxa with one simple seta. Basis about
three times as long as wide, with two dorsal and one ventral setulose setae. Carpus about 1.2 times as long as
merus, with three simple medial and four bipinnate ventral setae. Propodus with three simple distal setae.
Pereopod 4 (Fig. 3D): coxa absent. Basis about 4.2 times as long as wide, with one dorsoproximal and three
setulose ventral setae. Ischium with three simple setae. Merus shortest, about 0.8 times as long as carpus, with three
bipinnate distal setae. Carpus marginally shorter than propodus, with six bipinnate and one small simple distal
setae. Propodus as long as dactylus and unguis combined (without the terminal seta), with five spiniform bipinnate,
three simple and one setulose setae. Dactylus with denticles arranged in a ring around the insertion of unguis;
unguis slender with two ventral rows of small spines.
Pereopod 5 (Fig. 3E): articles with similar length to pereopod 4 except: basis with two setulose ventral setae.
Pereopod 6 (Fig. 3F): articles with similar length to pereopod 5 except: basis with one setulose dorsoproximal
seta. Ischium with two simple setae. Merus with two bipinnate distal setae. Carpus with five bipinnate distal and
three simple setae. Propodus with seven bipinnate, two simple and about five spiniform serrated setae next to
dactylus insertion.
Pleopods (Fig. 1E): basal article about 1.6 times as long as wide, with three plumose setae. Exopod with 13
plumose setae. Endopod with three inner and eleven outer plumose setae.
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FIGURE 3. Neotanais bicornutus sp. nov., female paratype: A, pereopod 1; B, pereopod 2; C, pereopod 3; D, pereopod 4; E,
pereopod 5; F, pereopod 6. Scale bars: A, C–F = 0.5 mm; B = 0.2 mm.
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Uropod (Fig. 2K): basal article naked, about 2.7 times as long as article 1. Endopod (damaged) with five
articles. Articles 1–4 bearing one to four simple setae, except article 4, naked. Article 5 longer, bearing five simple
distal setae. Exopod biarticulate, marginally shorter than article 1 of endopod, with one simple seta each article.
Description. Copulatory male allotype. Body (Fig. 1F): heavily calcified, no apparent setae. Dorsoventrally
flattened, about 7.2 times as long as wide. Cephalothorax: about 1.4 times as long as wide and 1.5 times as long as
the first two pereonites combined, with visible rostrum, with two strong anterolateral spines. Pereon: about 3.7
times as long as wide, about 50% of total body length. All pereonites rectangular, no plumose or simple setae;
pereonite 1 shortest, 0.3 times as long as wide; pereonites 2 and 3 subequal, 0.5 and 0.7, respectively; pereonites 4
and 5 similar; pereonite 6 about 0.6 times as long as wide. Pleon: no mid-ventral spur (Fig. 1G); about 1.6 times as
long as wide, 30% of total body length; all pleonites subequal, as wide as cephalothorax and pereonite 1, lateral
epimera with no seta. Pleotelson (Fig. 1I) about 0.6 times as long as wide, no apparent seta; uropod attachment
mid-length on pleotelson.
Antennule (Fig. 2L): damaged. Article 1 longest, 0.8 times as long as cephalothorax, about 7.5 times as long as
wide, with five proximal and six simple subdistal setae. Article 2 about 2.1 times as long as wide, twice as long as
articles 3 and 4 combined, with five simple medial setae. Article 3 naked. Article 4 short bearing several
aesthetascs. Other articles missed.
Mouthparts and maxilliped (Fig. 2M): all reduced, except: maxilliped basis robust, longer than wide, naked.
Endite naked. Palp article 1 naked, short; article 2 about 2.2 times as long as article 1, naked; article 3 about 1.3
times as long as article 2, with one simple proximal seta; article 4 longest, about 1.2 times as long as article 3, with
four simple distal setae.
Cheliped (Fig. 1H): sclerite (not illustrated), naked. Basis short, naked. Ischium shaped as a narrow incomplete
band extending from under the merus. Merus short, naked. Carpus 1.5 times as long as cephalothorax, widening
distally, with one simple seta and one visible protuberance bearing one long strong spine on ventral side. Propodus
with two strong dorsal projections and a third less visible (next to dactylus insertion), with slight crenation and one
simple seta next to dactylus insertion. Fixed finger inner margin with one simple seta, one deep groove medial and
one tooth distally. Dactylus marginally longer than fixed finger, with three proximal denticles and a row of
denticles distally.
Uropod (Fig. 1I): basal article attached at mid-lateral margin, about 1.2 times as long as article 1 of endopod.
Endopod damaged, article 1 with 8–10 fine setae proximal to mid-length on outer margin. Exopod biarticulate, half
size of endopod article 1, with one simple seta on each article.
Pleopods (not illustrated): basal article apparently naked. Exopod with two apparent articles (at least a fusion
line), first article bearing three and second with about 25–30 plumose setae. Endopod with about 30 inner and three
plumose outer setae.
Remarks. The specimens of Neotanais bicornutus sp. nov. were collected in the Antarctic Ocean and when
compared to Gardiner’s (1975) Neotanais scheme, it mostly resembles with the "robustus" group, which hitherto
held N. robustus Wolff, 1956b, N. antarcticus Kussakin, 1967 and N. tuberculatus Kudinova-Pasternak, 1970.
Gardiner (1975) recognized this group based mainly on copulatory male characters, although identification of
neotanaid males to species is difficult owing to their strong sexual polymorphism (Larsen & Błażewicz-
Paszkowycz 2003; Larsen 2005). Males are also frequently larger than females and show sexual dimorphism in the
proportions of the cephalothorax and the shape of the cheliped (Bamber 2007).
The male of N. bicornutus shares many characters with the "robustus" group by having a pleotelson about 1.5
times as wide as long (1.4 times in N. robustus, 1.7 in N. antarcticus), the pleonites are broad and rounded laterally
in dorsal view, lateral epimera without apparent seta, and they also bear a strong ventral spine arising from a
protuberance on the cheliped carpus (Fig. 1H). However, N. bicornutus differs from the "robustus" group by having
a dorsally naked cheliped carpus (while there are four setae in N. tuberculatus, five in N. antarcticus and apparently
two or three tiny setae in N. robustus). The uropod endopod article 1 has 8–10 fine setae proximal to mid-length on
outer margin, while these setae are lacking in the "robustus" group, but are found in N. hadalis Wolff, 1956a and N.
mesostenoceps Gardiner, 1975. It further differs in having a cheliped propodus with two strong and long dorsal
projections; within the "robustus" group only in N. robustus are there propodus projections, one small dorsal and
one longer ventral. The propodal dorsal crest is very slight in N. bicornutus, while in other species this character is
rather pronounced.
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The females vary mainly in the number of setae on the dorsal margin of the cheliped carpus; N. bicornutus has
about 15 setae, while there are nine in N. robustus, three in N. antarcticus and seven in N. tuberculatus. Also, the
pleonal epimera have three plumose setae, while the other "robustus" group species apparently lack this feature.
Neotanais bicornutus also resembles the Antarctic species N. krappschickelae Larsen & Błażewicz-Paszkowycz,
2003, in having a ring of spines on the unguis of pereopods 4–6 and the many dorsal setae on the cheliped carpus
(about 24 setae in N. krappschickelae and 15 setae in N. bicornutus). However, they can be easily distinguished by the
number of the plumose setae on almost all pereopod articles of both genders, the number of setae on the first
antennule article (about eleven plumose [described as bipinnate] in N. krappschickelae while N. bicornutus has two
simple and two plumose setae), the epimera setae (absent in N. krappschickelae while N. bicornutus has three), the
body size as well as other characters.
Genus Venusticrus Gardiner, 1975
Generic remarks. Neotanais rotermundiae Weigmann & Guerrero-Kommritz, 2009 shares several characters with V.
glandurus such as the uropod attached ventrally to the pleotelson (possibly an apomorphic character for Venusticrus)
and the distal bilobate and finely serrated terminal shield at insertion of the unguis of pereopods 4–6 (so far found only
in species of Venusticrus), therefore, the former is here transferred to Venusticrus (as V. rotermundiae comb. nov.). In
his revision of Neotanais, Larsen (1999) also transferred N. insolitus Gardiner, 1975 to Venusticrus, warranted by the
uropod attachment, the setation on the cheliped carpus, the slim cheliped basis and ventral pereopod attachments.
Nevertheless, V. insolitus, V. rotermundiae and the new species of Venusticrus described here, lack some of the
diagnostic characters listed by Gardiner (1975). Thus, a revised generic diagnosis is presented in the present study.
Gender. Masculine
Type species. Venusticrus glandurus Gardiner, 1975
Diagnosis. Female (modified after Gardiner 1975: 158). Pleonites with or without setae on lateral epimera.
Pleotelson acorn-shaped, marginally wider than long. Cheliped carpus with 12–18 simple dorsal setae; propodus
bearing a large, with brown-bordered dorsal crest (V. glandurus), or a distal margin curved and pointed (sometimes
with crenulated tip). Pereopod attachments not visible in dorsal view. Pereopods 4–6 with articles bearing long
plumose setae (V. glandurus) or with setulose, simple and bipinnate setae (all other species); dactylus with distal
bilobate and finely serrated terminal shield (as described by Weigmann & Guerrero-Kommritz (2009)); unguis
slender, with two fine ventral rows of spines. Uropod basal article attached posterior to mid-length on the ventral
margin of the pleotelson (not visible in dorsal view), exopod biarticulate.
Male (copulatory). Body heavily calcified, no apparent setae. Cephalothorax anterior margin relatively broad
widening posteriorly. All pereonites rectangular, no seta. Pleon lateral margin rounded. Antennule article 4 bearing
several long aesthetascs. Mouthparts all reduced. Cheliped sclerite, basis, ischium and merus naked, with two simple
ventral setae; fixed finger reduced, in the shape of a hammer. Uropod basal article attached posterior to mid-length on
the ventral margin of the pleotelson, endopod article 1 with several proximal to mid-length fine setae on outer margin.
Species included. Venusticrus glandurus Gardiner, 1975; V. insolitus (Gardiner, 1975); V. rotermundiae
(Weigmann & Guerrero-Kommritz, 2009) comb. nov.; V. thor sp. nov.
Venusticrus thor sp. nov.
(Figs 4–6)
Material examined. Holotype: female with oostegites (
MNHN-IU-2014-10184
), body total length: 6.6 mm, BIONOD
EBS-Supra Sta. 12–51, 12º 18’ 44’’N, 118º 8’ 2’’W, 11 April 2012, depth 4274 m.
Allotype: copulatory male (
MNHN-IU-2014-10185
), body total length: 10.6 mm, same locality as holotype.
Paratypes: one female with oostegites (
MNHN-IU-2014-10186
), (dissected), body total length 7.1 mm, same
locality as holotype. One female, manca II (
MNHN-IU-2014-10187
) body total length: 2.2 mm, same locality as
holotype. Two females with oostegites (used for DNA extraction, 28S - KT592230, KT592231; COI - KT592232,
KT592233), same locality as holotype. One female, manca II; one female, without oostegites (
MNHN-IU-2014-10188
),
BIONOD EBS-Supra Sta. 12–80, 14º 3’ 23’’N, 130º 2’ 23’’W, 21 April 2012, depth 4986 m.
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FIGURE 4. Venusticrus thor sp. nov., female with oostegites (MNHN-IU-2014-10184): A, holotype habitus dorsal view; B,
holotype habitus lateral view; female paratype (MNHN-IU-2014-10186): C, cheliped; D, uropod; manca female paratype
(MNHN-IU-2014-10187): E, habitus dorsal view; F, cheliped. Scale bars: A, B, E = 1 mm; C, D, F = 0.5 mm.
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TWO NEW ANTARCTIC AND PACIFIC NEOTANAIDS
Etymology. The name reflects the strong shape of male cheliped dactylus and fixed finger, as if holding a
hammer, like the god Thor from Nordic mythology. Noun in apposition.
Diagnosis. Female with oostegites. Body with pigmentation along the pereonites and pereopods, no apparent
setae. Cephalothorax as long as the first two pereonites combined, with no apparent rostrum or setae. Pleonites
lateral epimera with three plumose setae, ventral keel weekly pronounced. Labrum subtriangular, naked;
maxilliped endite with five setulose spiniform distal setae, pedestal palp with one long distal seta, and basis with
two simple proximal long setae. Cheliped carpus with about twelve simple dorsal setae; propodus with a distal
margin curved and pointed crenulated at tip. Uropod attached mid-length on ventral margin, basal article about 1.8
times as long as article 1 of endopod; exopod biarticulate, half as long as endopod article 1.
Male (copulatory): see diagnosis of Venusticrus.
Description. Body based on female holotype and appendages from dissected paratype.
Body (Fig. 4A, B): heavily calcified, dorsoventrally flattened, about 7.9 times as long as wide, with
pigmentation along the pereonites and pereopods; no apparent setae. Cephalothorax: about 1.3 times as long as
wide, as long as the first two pereonites combined; no apparent rostrum or setae. Pereon: about 4.6 times as long as
wide, about 57% of total body length. All pereonites rectangular; pereonite 1 shortest, about 0.5 times as long as
wide; pereonite 2–5 similar. Pereonite 6 slightly short than previous pereonite, about 0.8 times as long as wide.
Pleon (Fig. 5K): about 2.4 times as long as wide, about 28% of total body length; all pleonites subequal, with three
plumose setae on lateral epimera, ventral keel present but blunt and soft. Pleotelson (Fig. 5L): rounded, marginally
wider than long (1.1 times), uropod attached to mid-length on ventral margin.
Antennule (Fig. 5A): marginally longer than cephalothorax. Article 1 longest, about 2.9 times as long as wide,
about three times as long as article 2, with two plumose subdistal and two simple distal setae. Article 2 slightly
longer than articles 3, about 1.7 times as long as wide, with two plumose and three simple distal setae. Article 3
about 1.4 times as long as wide, as long as article 4, naked. Article 4 as long as articles 5–7 combined, with three
simple distal setae. Article 5 and 6 subequal with one annulated aesthetasc each. Article 7 with three simple distal
setae.
Antenna (Fig. 5B): about 1.5 times as long as antennule. Article 1 broader than other articles, as long as wide,
as long as articles 3 and 4 combined, naked. Article 2 longest, about 3.5 times as long as wide and 3.8 times as long
as article 3, with one fine spiniform distal and one plumose setae. Article 3 as long as wide, with one plumose distal
seta. Article 4 about 1.8 times as long as article 3, with one plumose distal seta. Article 5 about 2.5 times as long as
wide, marginally shorter than articles 6 and 7 combined, with two strong and four short simple setae, one setulose
distal seta. Article 6 about 2.8 times as long article 7, with four simple distal setae. Articles 7–9 subequal, bearing
three, two and four simple distal setae, respectively.
Mouthparts (Fig. 5C–J): labrum (Fig. 5C) subtriangular, naked. Mandibles (Fig. 5D, E) heavily calcified,
molar with several pointed distal denticles. Left mandible (Fig. 5D) incisor with about five blunt denticles, setal
row with two serrated setae; lacinia mobilis marginally longer than incisor with one or two blunt crenulated
denticles. Right mandible (Fig. 5E) incisor with several blunt distal denticles, setal row with two serrated setae.
Labium (Fig. 5H): palp elongated with several setules at tip, with two outer denticles and two inner small spiniform
setae, lobes with distal and lateral setules. Maxillule (Fig. 5F, G) outer endite (Fig. 5F) with seven spiniform and
three bipinnate distal setae; inner endite (Fig. 5G) with one plumose and three short spiniform distal setae. Maxilla
(Fig. 5I) with several setules on ventral and lateral margins; inner lobe of fixed endite with several simple distal
setae; outer lobe with two spiniform and three bifurcate spiniform setae; inner lobe of movable endite with three
spiniform setae; outer lobe with two bipinnate and three spiniform setae.
Maxilliped (Fig. 5J) endite with five inner distal setulose spiniform setae and, several fine simple setae. Basis
about 1.8 times as long as wide, with two proximal long simple setae, palp pedestal with one long simple distal
seta. Palp article 1, marginally longer than article 2 (about 1.1 times), about 1.6 times as long as wide, naked;
article 2 shortest, about 1.3 times as long as wide, with one simple outer distal seta and about five simple inner
setae; article 3 longest, about 1.5 times as long as article 2, and about 2.2 times as long as wide, with five simple
inner setae; article 4 slender, about 2.7 times as long as wide, with five pinnate and five simple setae. Epignath not
recovered.
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FIGURE 5. Venusticrus thor sp. nov., female paratype: A, antennule; B, antenna; C, labrum; D, left mandible; E, right
mandible; F, maxillule, outer endite; G, maxillule, inner endite; H, labium; I, maxilla; J, maxilliped; K, pleon and pleotelson; L,
detail of pleotelson and uropod ventral attachment; M, pleopod. Scale bars: A, L = 0.5 mm; B–J, M = 0.2 mm; K = 1 mm.
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TWO NEW ANTARCTIC AND PACIFIC NEOTANAIDS
Cheliped (Fig. 4C): sclerite rectangular (illustrated on Fig. 4B). Basis about 1.7 times as long as wide, naked.
Ischium short, hardly visible. Merus subtriangular, with one simple seta. Carpus marginally longer than basis, about
2.2 times as long as wide, with two simple ventral setae and, about 12 simple dorsal setae. Propodus slightly shorter
than basis, with a distal margin curved and pointed crenulated at tip, with one simple seta at the dactylus insertion.
Fixed finger with one simple inner proximal seta, three simple inner distal and two on outer margin next to unguis,
inner margin with three to four blunt denticles; dactylus marginally shorter than fixed finger, with one simple inner
proximal seta.
Pereopod 1 (Fig. 6G): coxa small, naked (not illustrated). Basis about three times as long as wide, about 1.7
times as long as merus, with three short setulose proximal setae. Ischium with two simple setae. Merus about 1.3
times as long as carpus, with two simple distal and one bipinnate setae. Carpus marginally shorter than propodus,
about 1.9 times as long as wide, with two spiniform, three bipinnate, one spiniform- serrated distal and, six simple
setae. Propodus as long as dactylus and unguis combined, about 2.7 times as long as wide, with one setulose medial,
three simple, one pinnate, five bipinnate and one hook-like spiniform serrated setae. Dactylus with one simple
proximal short seta; unguis naked.
Pereopod 2 (Fig. 6H): articles with similar length to pereopod 1 except: basis marginally broader, about 2.2
times as long as wide, with two setulose proximal setae. Carpus about 1.5 times as long as merus. Propodus with six
bipinnate and six simple setae. Dactylus marginally longer than propodus.
Pereopod 3 (Fig. 6I): articles with similar length to pereopod 2 except: coxa with one setulose seta. Basis with
three setulose setae. Carpus as long as propodus with seven simple setae. Propodus with five bipinnate setae.
Dactylus about 1.4 times as long as propodus.
Pereopod 4 (Fig. 6J): coxa absent. Basis as long as carpus and propodus combined, about 3.9 times as long as
wide, with two setulose proximal setae. Ischium with two simple setae. Merus shortest, about 0.6 times as long as
carpus, with one bipinnate and two simple distal setae. Carpus marginally longer than propodus and about 2.4 times
as long as wide, with six bipinnate and five simple setae. Propodus half size of dactylus and unguis combined, with
one setulose, six bipinnate and three simple setae. Dactylus with distal bilobate and finely serrated terminal shield at
insertion of unguis; unguis slender, with two fine ventral rows of setae.
Pereopod 5 (Fig. 6K): articles with similar length to pereopod 4 except: basis broader, about 2.5 times as long
as wide. Carpus with twelve bipinnate and one simple fine distal setae. Propodus with seven bipinnate and three
simple setae.
Pereopod 6 (Fig. 6L): articles with similar length to pereopod 5 except: basis with one setulose proximal seta.
Carpus with eight bipinnate and one simple fine distal setae.
Pleopods (Fig. 5M): basal article short and stout, with three plumose setae. Exopod with about 17 plumose
setae. Endopod with three inner and 15 outer plumose setae.
Uropod (Fig. 4D): basal article naked, about 1.8 times as long as article 1 of endopod. Endopod with 14 articles.
Articles 1–3 naked, subequal. Article 4 with one pair of plumose and other pair of simple setae. Article 5 with one
simple seta. Article 6 with one plumose seta and three simple setae. Articles 7, 9 and 11–13 subequal, naked. Article
8 as article 4. Article 10 with one plumose and three simple setae. Article 14 one plumose, four simple setae and one
aesthetasc. Exopod biarticulate, half the length of endopod article 1, with one simple seta on each article.
Manca II (Fig. 4E, F). Body (Fig. 4E) female paratype, length 2.2 mm, not calcified, about six times as long as
wide; no apparent pigmentation, naked. Cephalothorax: as adult female. Cheliped (Fig. 4F): sclerite rectangular.
Basis about 1.2 times as long as wide, naked. No apparent ischium. Merus as adult female. Carpus with six simple
dorsal and two ventral setae. Propodus with a distal margin very curved and pointed at tip, almost as reaching
dactylus. Fixed finger and dactylus with similar setation as adult female paratype. Pereopods 1–6 (not illustrated)
apparently as adult female but with less spiniform bipinnate setae on propodus. Uropods damaged.
Description. Copulatory male allotype. Body (Fig. 6A, B): heavily calcified, no apparent setae. Dorsoventrally
flattened, about 8.1 times as long as wide. Cephalothorax: anterior margin relatively broad, widening posteriorly,
about 2.2 times as long as wide and 1.2 times as long as the first three pereonites combined, naked, no apparent
rostrum. Pereon: about 4.6 times as long as wide, about 49% of total body length. All pereonites pentagonal, no
plumose or simple setae; pereonite 1 shortest, about 0.5 times as long as wide; pereonites 2 and 6 subequal, about
0.8 and 0.7 times as long as wide, respectively; pereonites 3–5 subequal, marginally longer than wide; pereonite 6
with two tiny genital cone (only one illustrated, Fig. 6B). Pleon: about 2.3 times as long as wide, 24% of total body
length. All pleonites subequal, lateral margin rounded, with one fine simple seta. Pleotelson (Fig. 6C): marginally
wider than long, naked.
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FIGURE 6. Venusticrus thor sp. nov., copulatory male allotype (MNHN-IU-2014-10185): A, habitus dorsal view; B, habitus
lateral view; C, pleotelson and uropods; D, antennule; E, antenna; F, pleopod; female paratype: G, pereopod 1; H, pereopod 2; I,
pereopod 3; J, pereopod 4; K, pereopod 5; L, pereopod 6. Arrow in Fig. 6B indicates one of two genital cone. Scale bars: A, B
= 2 mm; C–F = 0.2 mm; G–L = 0.5 mm.
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Antennule (Fig. 6D): article 1 longest, about 0.6 times as long as cephalothorax, about 6.6 times as long as
wide, with one medial and four simple distal setae. Article 2 about 2.8 times as long as article 3, with one simple
distal seta. Article 3 as long as articles 5 and 6 combined, naked. Article 4 rounded at basis bearing several long
aesthetascs, about 2.6 times as long as wide. Articles 5 and 6 bearing one long aesthetasc each. Article 7 with three
simple setae at tip.
Antenna (Fig. 6E): about 0.8 times as long as antennule. Article 1 broader than following articles, about 1.8
times as long as wide, naked. Article 2 slender, about 5.7 times as long as wide, about 1.1 times as long as articles
3–5 combined, with one simple subdistal seta. Article 3 as long as article 7, with one simple distal seta. Article 4
marginally shorter than article 6, naked. Article 5 about 3.3 times as long as wide, with two plumose and a row (7–
10) of subdistal setae. Article 6 about 0.8 times as long as articles 7–9 combined, with two simple distal setae.
Articles 7 marginally longer than article 8, both naked. Article 9 shortest, with six simple distal setae.
Mouthparts and maxilliped: all reduced.
Cheliped (Fig. 6B): sclerite rectangular, apparently naked. Basis stout. Ischium shaped as a narrow incomplete
band extending from under the merus. Merus short, naked. Carpus long as in "robustus" group, but thinner and
without protuberance or a long strong spine on ventral margin, about 1.9 times as long as cephalothorax, with two
ventral simple setae. Propodus about 0.6 times as long carpus, about 4.8 times as long as wide, with two simple
distal setae. Fixed finger reduced, with several blunt denticles on inner and outer margins bearing three and two
simple setae, respectively. Dactylus long, about 2.6 times as long as fixed finger and slightly shorter than propodus
(0.9 times), with a row of distal blunt denticles on inner and outer margins, naked.
Pleopods (Fig. 6F): basal article about 1.2 times as long as wide, naked. Exopod with two apparent articles (at
least a fusion line), article 1 with three small and one long plumose setae, article 2 with about 17 plumose setae.
Endopod with about 16 inner and four plumose outer setae.
Uropod (Fig. 6C): basal article attached mid-length on ventral margin. Endopod article 1 with several fine
setae proximal to mid-length on outer margin, other articles missed. Exopod biarticulate, with two simple distal
setae.
Remarks. Venusticrus thor sp. nov. is the fourth described species of Venusticrus and the first record for the
Pacific Ocean. The new species is similar its congeners in having characters such as: several dorsal setae on the
cheliped carpus (more than ten), the type of armament on the dactylus at the unguis insertion of pereopods 4–6 (see
Fig. 6J–L), the cheliped propodus with a dorsodistal margin that is long, curved and pointed at its tip (also seen in
V. insolitus and V. rotermundiae), and by the uropods attached mid-length on the ventral margin of the pleotelson—
a diagnostic character for Venusticrus.
The females in the Neotanaidae family are known to have a very conservative morphology so often require a
combination of characters to distinguish them. With the exception of V. glandurus, which differs from its congeners
mainly due to the antennule article 1 and the number of plumose setae on the pereopods and uropods, the females
of V. rotermundiae and V. insolitus share many characters with V. thor. However they can be separated from V. t h o r
as this new species has the following unique character combination: body length about 7.9 times as long as wide
(nine times in V. insolitus and V. rotermundiae); pleonal lateral epimera with three setae (one or two in V. insolitus
and absent in V. rotermundiae), pleon with a very weak straight mid-ventral keel (pronounced and posteriorly
directed in V. glandurus and V. insolitus); antennule article 1 about 2.9 times as long as wide (3.1 times in V.
insolitus and 2.7 times in V. rotermundiae), with two simple and two plumose setae (with several setulose setae in
V. rotermundiae and five simple setae in V. insolitus); maxilliped endite with five inner distal setulose spiniform
setae (two in V. insolitus and three setulose spiniform and several tiny spiniform setae in V. rotermundiae),
pereopods 1–6 number and type of seta; uropod basal article about 1.8 times as long as endopod article 1 (1.2 times
in V. glandurus, 1.5 times and in V. rotermundiae); uropod exopod half size of endopod article 1 (1.5 times as long
as endopod article 1 in V. insolitus, 0.8 in V. rotermundiae and 0.4 in V. glandurus).
The first record of a Venusticrus male was made by Gardiner (1975: 154) under the species V. insolitus.
However, the author reported the male as non copulatory and just described the main characters without providing
illustrations. The second record of a Venusticrus male was by Weigmann & Guerrero-Kommritz (2009: 28), when
describing V. rotermundiae. The authors reported the specimen as damaged and lacking the chelipeds and
accordingly did not illustrate or describe the specimen fully. To redeem this, a diagnosis and illustrations of a
Venusticrus male are presented here. In addition, Weigmann & Guerrero-Kommritz (2009: 28), cited a "rudimental
oostegites on pereopods 1 to 4" that was not found in the allotype of V. thor. Finally, a row of fine simple setae
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(about 8–10) was observed proximal to mid-length on the outer margin of the uropodal endopod article 1 (see Figs.
1I and 6C). This character was previously found in N. mesostenoceps and N. hadalis and is well illustrated by
Gardiner (1975: 69, fig. 27D; 75, fig. 32B, respectively). It is unclear if this character has been missed in other
previous descriptions, hence should be carefully observed when describing new species of Venusticrus or
Neotanais.
All previous species of Venusticrus (including V. rotermundiae) were collected in the Atlantic Ocean, with the
bathymetric distribution between 4460–5139 m.
Discussion
Gardiner (1975: 158) separated Venusticrus from the other three neotanaid genera Carololangia, Herpotanais and
Neotanais, by some unusual characters such as the row of setae on the chelipedal merus, the cheliped bearing a
large, thin, brown-bordered dorsal crest, the pleotelson acorn-shaped in dorsal view, the uropods attached mid-
length on ventral margin on the pleotelson, as well as other characters. Later, after a revision of genus Neotanais,
Larsen (1999) transferred V. insolitus to Venusticrus due to its ventral pereopod and uropod attachments, setation of
the cheliped carpus, and slim cheliped basis. In the present study we found additional characters that strongly
support this transfer to Venusticrus. For instance, a consistent difference was observed between Neotanais,
Herpotanais, Carololangia and Venusticrus regarding the armament at the dactylus of pereopods 4–6. The first
three genera (at least where the species are properly described and illustrated) have a dactylus with a row of
denticles arranged as a ring around the insertion of unguis (Fig. 3D–F). In Venusticrus, the species have the
previously-mentioned dactylus armament consisting of bilobate and finely serrated terminal shields at the insertion
of the unguis (Fig. 6J–L). Until now, this character has not been used for genus separation, but this study indicates
that it might be a potential generic character and should be considered for further morphological analysis, since it is
very conservative and easy to observe. Regarding the ventral uropod attachment, this character is found exclusively
in the species of Venusticrus, possibly being an apomorphic character of this genus, while in Neotanais,
Herpotanais and Carololangia the uropod is laterally attached.
Acknowledgments
This work is part of the senior author´s Ph.D. dissertation (CLAS), funded by CAPES—Coordenação de
Aperfeiçoamento de Pessoal de Nível Superior (Grant number 5428/10–6). In addition, this research was partially
supported by the European Regional Development Fund (ERDF) through the COMPETE—Operational
Competitiveness Programme and national funds through FCT—Foundation for Science and Technology, under the
project “PEst-C/MAR/LA0015/2013” "PEst-C/MAR/LA0015/2014".
The authors are grateful to Professor A. Brandt, University of Hamburg for allowing the use of the material
from the ANDEEP cruises and to Dr. Graham Bird for identifying most of the ANDEEP material to genus level.
Thanks are also extended to Drs Menot (IFREMER), to Rühlemann (BGR), chief scientists of the BIONOD
expedition for inviting the junior author to participate in this cruise and for making the material available for study.
We also thank to the editor Dr. Graham Bird and an anonymous referee who provided very constructive
suggestions.
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