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A new genus of family Akanthophoreidae and new species of genus Parakanthophoreus Larsen & Araújo-Silva, 2014 (Crustacea: Tanaidacea: Tanaidomorpha) from the North Atlantic

A new genus of family Akanthophoreidae and new species of genus Parakanthophoreus Larsen &... Based on tanaidacean material collected primarily during the IceAGE cruises in the North Atlantic, three new species from the family Akanthophoreidae are described; one is classified in the genus Parakanthophoreus, and the other two represent a new genus—Brixia n. gen. The main characters that distinguish Brixia n. gen. from other akanthophoreids is the lack of fully developed pleopods in adult females and a seven-articled antenna. The diagnosis of the family Akanthophoreidae is amended. Cheliped ornamentation of Parakanthophoreus catharina n. sp. is illustrated using SEM pictures. . . . . . Keywords IceAGE Icelandic waters Brixia n. gen. BIOICE BIOFAR AFEN Introduction in Akanthophoreinae were moved later to other families (Larsen and Wilson 2002), e.g., Araphura Bird & Holdich, Sieg (1986a) proposed the tanaidacean subfamily 1984 and Araphuroides Sieg, 1986 to Tanaellidae, and Akanthophoreinae (Anarthruridae) to accommodate Filitanais Kudinova-Pasternak, 1973 to Colletteidae; other paratanaoids characterized by, e.g., a four-articled antennule, genera were placed in the family incertae sedis, e.g., six-articled antenna, mandibular molar usually pointed, at Andrognathia Sieg, 1983, Leptognathioides Bird & Holdich, least the carpus of pereopod-1 with spines, pleopods well- 1984, and Robustochelia Kudinova-Pasternak, 1983. developed, reduced or absent, and uropods with one or two The Akanthophoreinae was rediagnosed and elevated to rami. The broad diagnosis resulted in a prolific composition of family level by Błażewicz-Paszkowycz and Bamber (2011) this subfamily and in a short time it appeared to be a non- with only the genus Akanthophoreus Sieg, 1986 included. monophyletic taxon. Most of the genera originally included Later, Błażewicz-Paszkowycz et al. (2013) added Chauliopleona Dojiri & Sieg, 1997 to this family, but without giving a justification for this reclassification. The most recent This article is registered in ZooBank under: urn:lsid:zoobank.org:pub: revision of this family was by Larsen and Araújo-Silva (2014), 6D7D4138-0170-44BD-BCD9-48B9983B88E1 who, with a phylogenetic approach, accommodated within This article is part of the Topical Collection on Biodiversity of Icelandic Akanthophoreidae a few other genera: Mimicarhaphura Sieg, Waters by Karin Meißner, Saskia Brix, Ken M. Halanych, and Anna 1986; Paraleptognathia Kudinova-Pasternak, 1981; Jazdzewska. Stenotanais Bird & Holdich, 1984; Tumidochelia Knight, Communicated by S. Brix Larsen & Heard, 2003; and the newly described Parakanthophoreus Larsen & Araújo-Silva, 2014. Most re- * Piotr Jóźwiak cently, Bird (2015) added Saurotipleona to this list of [email protected] akanthophoreids. One of the new taxa described below, a Parakanthophoreus species, has a strong resemblance to the Laboratory of Polar Biology and Oceanobiology, University of Łódź, genus Paraleptognathia Kudinova-Pasternak, 1981, which has Banacha 12/16, 90-237 Łódź, Poland been systematically confusing since its first diagnosis (Bird EcoAnalysts, Inc., Moscow, ID 83843, USA 2007;Błażewicz-Paszkowycz and Bamber 2011;Larsenand Independent, Kāpiti, Waikanae, New Zealand Araújo-Silva 2014). 898 Mar Biodiv (2018) 48:897–914 Larsen (2005) pointed out that in deep-sea tanaidomorphan The material is deposited in the Zoological Museum, Hamburg taxa females may lack pleopods. It was previously recorded and the National Museums of Scotland (NMS), Glasgow. for, e.g., all genera in the family Agathotanaidae; Caudalonga Larsen, 2005; Cetiopyge Larsen & Heard, 2002 and Collettea Lang, 1973 from the Colletteidae, and Anarthrura Sars, 1882 Systematics from the Anarthruridae. Until now, the Akanthophoreidae was represented exclusively by taxa with well-developed pleopods Suborder Tanaidomorpha Sieg, 1980 but a new genus described below is with reduced pleopods or Superfamily Paratanaoidea Lang, 1949 completely without pleopods. Family Akanthophoreidae Sieg, 1986 In this paper, we deal with a series of akanthophoreids Diagnosis see Larsen and Araújo-Silva (2014) but with collected in the waters off Iceland during the IceAGE additions: all pereopod meri and carpi with slender spines. (Icelandic Animals Genetic and Ecology; Brix et al. 2014) Pleopods well-developed, reduced or absent. initiative, which allows us to erect a new genus with two Remarks. The diagnosis of the family is modified primarily new species and describe a Paraleptognathia-like species of regarding the pleopods, where they may be absent or rudimen- Parakanthophoreus. tary in the new genus described below. Among the original members of the family, formerly the Tribe Akanthophoreinae within the family Anarthruridae Material and methods Lang, 1971, Paraleptognathia was distinguished by a combi- nation of: a five-articled antennule, with two aesthetascs on the Samples were collected during two IceAGE cruises. The first terminal article, a maxillule with eight spines, an epignath took place in August/September of 2011 on board the R/V tipped with a seta, the cheliped with a strong carpal shield Meteor and the second in August 2013 on board the R/V (and all cheliped articles covered with setules), and a pleopod Poseidon. During those initiatives, various sampling gears exopod of two articles. The unique five-articled antennules in were deployed, e.g., Van Veen grab, Shipek grab, epibenthic the type species of the genus, Paraleptognathia typica sledge, or box corer (Brix et al. 2014). Reference is also made Kudinova-Pasternak, 1981 (Kudinova-Pasternak 1981) were to material collected during the BIOICE, BIOFAR, and AFEN later discussed by Guerrero-Kommritz (2004) and were treated projects from the same, and adjacent, areas (e.g., Bird 2001, as a deviation of the standard state or a unique character of the 2015)(Fig. 1). For environmental data, please refer to Bird species, especially since that P. bacescui Kudinova-Pasternak, (2015), Brix et al. (2011), and Table 1 herein. 1985 has standard four-articled antennules (Kudinova- Specimens were dissected using chemically sharpened Pasternak 1985). This character is known to be a feature of tungsten needles, mounted in glycerin on slides and sealed immature or Bpreparatory^ males in several paratanaoid groups with nail varnish. Drawings were initially prepared using a (Bird 2007,Błażewicz-Paszkowycz et al. 2011). For over a microscope combined with a camera lucida and redrawn on decade, Paraleptognathia was known only from two species, a digital tablet as proposed by Coleman (2003). butDojiriand Sieg (1997) described P. bisetulosa and moved The body length to width ratio was calculated using mea- Akanthophoreus gracilis (Krøyer, 1842) to Paraleptognathia. surements from the tip of the carapace to the end of the Guerrero-Kommritz (2004) synonymized Akanthophoreus pleotelson, and of the widest part of carapace. The length with Paraleptognathia, resulting in a total of 17 species classified and width of articles were measured along their central axes. within this genus. This was not accepted by Bird (2007)and it Dehydration for SE microscopy was carried out sequential- was again split into Akanthophoreus, Parakanthophoreus,and ly in the tissue processor (Leica EMTP) with 95% and abso- Paraleptognathia by Larsen and Araújo-Silva (2014). Although lute ethanol. Specimens were then transferred to a critical the morphology of one of the new akanthophoreid species found point dryer (Leica EMCPD 300) with parameters set as fol- in the Northeast Atlantic during IceAGE cruises corresponds with lows: mixing 50%, CO in–slow with 2 min delay, exchange the main characters defining Paraleptognathia suggested by Bird speed–5with 20 cycles, gas out heat and speed–fast. Dried (2007) and Larsen and Araújo-Silva (2014), namely a large carpal specimens were transferred on coverslips and coated with 6- shield and cheliped articles with Bornamentation,^ it is considered nm-thick layer of gold (Leica EM ACE200). SEM pictures that Paraleptognathia is too ill-defined and poorly supported by were made on Phenom ProX. type material to warrant the addition of new species. The morphological terminology follows that proposed by Brixia n. gen. Błażewicz-Paszkowycz et al. (2013). The term Bpalm^ is used Registered in ZooBank under: urn:lsid:zoobank.org: for the proximal part of the cheliped propodus, i.e., without act:094408C2-6DD8-4161-9ACCEE0F743E505A the fixed finger. It is delimited by the line running through Diagnosis. Pleonite-5 without ventral spur. Pleotelson with- dactylus insertion and reaching the propodus ventral margin out lateral spurs, as long as broad or longer. Antenna with seven at a right angle. distinct articles. Cheliped with distinct carpal shield; carpus and Mar Biodiv (2018) 48:897–914 899 Fig. 1 Map with distribution of described species: yellow circle— Brixia aurora n. sp.; red circle— B. borealis n. sp.; green circles— Parakanthophoreus catharina n. sp. propodus without ornamentation. Pleopods rudimentary or 989-1, 61° 42.26′ N 19° 32.21′ W-61° 42.17′ N 19° 32.03′ W, absent. Uropod basis and endopod segment without spur. depth 1919.9–1925.6 m, EBS, 31 August 2011. Type species. Brixia aurora n. sp., by original designation. Etymology. From the Latin aurora, meaning sunrise. The Etymology. The genus is dedicated to Saskia Brix, the name refers to the Aurora Borealis that delighted and leader of the IceAGE project. entertained the IceAGE crew and participants during the col- Gender. Feminine. lection of the material. Species included. B. aurora n. sp., B. borealis n. sp. Diagnosis. Cephalothorax 1.7 times longer than wide. Remarks. Most of the characters in the amended diagnosis Pleotelson elongate, 1.4 times longer than wide, as long as last of Akanthophoreidae by Larsen and Araújo-Silva (2014)are three pleonites. Antennule article-1 3.7 times longer than wide, observable in Brixia n. gen. Nevertheless, seven distinct arti- longer than distal three articles combined length. Ischium of cles in the antenna, and rudimentary or absent pleopods may be pereopods 1–3 and 6 with one seta (pereopods 4–5 missing). unique characters for the family; the pleotelson is also longer Uropod basis four times longer than wide and as long as endopod than in most, if not all, Akanthophoreus, Chauliopleona, segment-1; exopod 0.6 times as long as endopod segment-1. Parakanthophoreus,and Paraleptognathia species, where it Description of female. Body (Fig. 2A, B) slender, 10.5 is shorter than broad. The antennal character derives from times longer than wide, 3.3 mm long. Cephalothorax two articles (4 and 5) with complete division, homologous to subrectangular, tapering anteriorly, 1.7 times longer than article-4 in those antennae exhibiting partial or indistinct artic- wide, withslighttriangular rostrum;carapacewithsingle seta ulation (Bfusion line^ sensu Larsen et al. 2006) within this at each anterolateral corner; eyelobes poorly marked. element as expressed in other akanthophoreids. The pleopod Pereonites parallel-sided or weakly convex; pereonite-1 rect- character (lack or rudimentary) is somewhat compromised by angular, shortest, 0.4 times as long as cephalothorax, 1.5 times the stated absence of these in Parakanthophoreus fastuosus longer than wide; pereonites 2, 4, and 5 subequal in length, 1.5 (Guerrero-Kommritz, 2004: 56), although this is not remarked times longer than pereonite-1, as long as wide, with parallel on or included in the generic diagnosis (op.cit.: 8–10). lateral margins; pereonite-6 just shorter, 1.4 times longer than Moreover, the results of molecular phylogeny (Błażewicz and pereonite-1, as wide as long. Pleon as long as combined length Jennings unpublished data) unequivocally support Brixia hav- of three last pereonites; pleonites subequal in length; each ing an affinity to, but separateness from, other akanthophoreids pleonite 2.4 times wider than long, without distinct lateral such as Parakanthophoreus or Chauliopleona. setae. Pleotelson elongate (relative to most akanthophoreids), Brixia aurora n. sp. 1.4 times longer than wide, as long as last three pleonites, Registered in ZooBank under: urn:lsid:zoobank.org: slightly tapering distally, bearing two setae mid-posteriorly. act:42A284B1-1D31-4375-A208-6F1D8E17057C Antennule (Fig. 2A, C) about 0.75 times length of cepha- (Figs. 2, 3) lothorax; article-1 3.7 times longer than wide, longer than Material examined: holotype non-ovigerous female distal three articles together, with two outer distal groups of (3.5 mm long) dissected on slides (ZMH K-52920), st. 853/ penicillate setae and one simple seta that reaches distal margin 900 Mar Biodiv (2018) 48:897–914 Table 1 Station data for non IceAge samples referred to in text Survey Sample Gear Date Locality Latitude (°N) Longitude Depth (m) B. Temp. (°C) Temp. (°C) Sediment AFEN 1996 53819#3 MC 25/07/1996 Faroe-Shetland Channel 61.59 − 1.832 1094 Fine sand AFEN 1996 53828#1 MC 05/08/1996 Faroe-Shetland Channel 61.4355 − 2.6175 1350 Very fine sand AFEN 1996 53833#1 MC 05/08/1996 Faroe-Shetland Channel 61.603 − 2.443 1513 Coarse silt AFEN 1998 54509#2 MC 25/05/1998 Faroe-Shetland Channel 62.473 0.382 901 Muddy sand AFEN 1998 54521#1/3 MC 25/05/1998 Faroe-Shetland Channel 61.732 − 1.618 1014 Gravelly muddy sand AFEN 1998 54522#1/2 MC 25/05/1998 Faroe-Shetland Channel 61.74 − 1.6625 1172 Muddy sand AFEN 1998 54524#2 MC 25/05/1998 Faroe-Shetland Channel 61.766 − 1.814 1393 Muddy sand BIOFAR 167 RP 08/05/1988 Norwegian Basin 62.781 − 3.52 1032 − 0.85 − 0.95 BIOFAR 168 RP 08/05/1988 Faroe-Norway Rise 62.693 − 3.62 899 − 0.66 − 0.95 Silt BIOFAR 169 RP 08/05/1988 Faroe-Norway Rise 62.621 − 3.54 808 − 0.6 − 0.7 BIOFAR 170 RP 08/05/1988 Faroe-Norway Rise 62.532 − 3.518 699 − 0.6 − 0.63 BIOFAR 172 RP 09/05/1988 Faroe-Norway Rise 62.318 − 3.914 507 1 1 Sand BIOFAR 478 DS 21/07/1989 Faroe-Shetland Channel 61.061 − 4.732 973 − 0.8 Soft BIOFAR 769 DS 06/10/1990 Faroe-Norway Rise 62.332 − 3.1 645 − 0.6 Silt, gravel, cobbles, stones BIOICE 2010 RP 20/07/1991 Iceland-Faroe Rise [North] 65.3047 − 11.2358 629 1 − 0.36 BIOICE 2011 RP 20/07/1991 Iceland-Faroe Rise [North] 65.3481 − 11.1711 761.5 1 − 0.42 BIOICE 2030 RP 23/07/1991 Iceland Plateau 67.0039 − 13.2628 830 0 − 0.55 BIOICE 2077 RP 03/07/1992 Iceland Plateau 67.675 − 17.173 1048 − 1 − 0.5 Silty sand BIOICE 2088 RP 04/07/1992 Iceland Plateau 67.239 − 17.857 617 − 1 − 0.4 Sand, silt mixed with foraminifera BIOICE 2124 RP 07/07/1992 Kolbeinsey Ridge 67.186 − 19.564 347 0 0.2 BIOICE 2136 RP 08/07/1992 Kolbeinsey Ridge 66.726 − 18.954 417 0 0.6 Soft BIOICE 2315 Sn 02/05/1993 Iceland-Faroe Rise [North] 64.1 − 9.05 991 2 Sandy silt BIOICE 2317 RP 02/05/1993 Iceland-Faroe Rise [North] 64.117 − 9.05 996 2 BIOICE 2318 Sn 02/05/1993 Iceland-Faroe Rise [North] 64.033 − 9.617 772 2 Sandy silt BIOICE 2323 RP 03/05/1993 Iceland-Faroe Rise [North] 63.917 − 10.083 623 2 BIOICE 2364 RP 08/05/1993 Iceland-Faroe Rise [North] 64.583 − 10.05 605 2 Sandy silt BIOICE 2638 RP 13/07/1994 Iceland Plateau 67.8113 − 15.503 1008 − 1 − 0.6 Silt BIOICE 2743 Sn 30/07/1995 Greenland-Iceland Rise 67.7492 − 20.4735 730 0 BIOICE 2749 Sn 31/07/1995 Greenland-Iceland Rise 68.0188 − 20.6547 956 0 − 0.6 Sandy gravelly silt BIOICE 2762 RP 01/08/1995 Iceland Plateau 67.9247 − 17.7043 1129 −1 − 0.5 BIOICE 2765 RP 10/08/1995 Iceland Plateau 68.0659 − 17.3233 1224.5 − 1 − 0.52 BIOICE 3023 RP 08/07/1997 Iceland-Faroe Rise [South] 63.4666 − 12.2153 427 3 4.66 BIOICE 3108 RP 21/08/1999 Greenland-Iceland Rise 67.1864 − 20.4537 326 1 − 0.21 BIOICE 3115 RP 22/08/1999 Kolbeinsey Ridge 67.4212 − 19.47925 604 −1 − 0.57 Mar Biodiv (2018) 48:897–914 901 Table 1 (continued) Survey Sample Gear Date Locality Latitude (°N) Longitude Depth (m) B. Temp. (°C) Temp. (°C) Sediment BIOICE 3121 DS 22/08/1999 Kolbeinsey Ridge 68.0423 − 18.0618 732 − 1 − 0.28 Silt BIOICE 3198 RP 08/07/2001 Iceland-Faroe Rise [North] 65.3005 − 8.3169 935 1 − 0.63 BIOICE 3225 RP 12/07/2001 Norwegian Basin 68.3461 − 8.1532 1994 0 − 0.86 DTI 2000 55282#1 MC 05/08/2000 Faroe-Shetland Channel 61.6135 − 1.9803 1305 DTI 2000 55283#1 MC 05/08/2000 Faroe-Shetland Channel 61.5901 − 1.9739 1262 DTI 2000 55287#1,2 MC 05/08/2000 Faroe-Shetland Channel 61.6535 − 1.8793 1290 DTI 2000 55299#1 MC 09/08/2000 Faroe-Shetland Channel 61.6028 − 1.924 1236 DTI 2000 55304#1 MC 09/08/2000 Faroe-Shetland Channel 61.6081 − 1.7733 1026 DTI 2000 55304#2 MC 09/08/2000 Faroe-Shetland Channel 61.6081 − 1.7733 1026 DTI 2000 55308#1 MC 10/08/2000 Faroe-Shetland Channel 61.648 − 1.8417 1234 DTI 2000 55310#2,3 MC 11/08/2000 Faroe-Shetland Channel 60.6632 − 4.4018 1095 DTI 2000 55322#1 MC 16/08/2000 Faroe-Shetland Channel 62.5917 0.1463 1068 DTI 2000 55324#1 MC 16/08/2000 Faroe-Shetland Channel 62.4385 0.1235 1046 DTI 2000 55326#1 MC 16/08/2000 Faroe-Shetland Channel 62.4415 0.0935 1144 DTI 2000 55328#1,2 MC 16/08/2000 Faroe-Shetland Channel 62.2401 0.1841 727 DTI 2000 55332#1,2 MC 17/08/2000 Faroe-Shetland Channel 62.207 0.381 640 DTI 2000 55336#1 MC 17/08/2000 Faroe-Shetland Channel 62.3015 0.1808 890 DTI 2000 55340#1 MC 18/08/2000 Faroe-Shetland Channel 62.4161 0.0711 1062 DTI 2000 55344#1 MC 18/08/2000 Faroe-Shetland Channel 62.4165 0.0711 1062 DTI 2000 55344#2 MC 18/08/2000 Faroe-Shetland Channel 62.4163 0.0711 1062 DTI 2000 55346#1 MC 19/08/2000 Faroe-Shetland Channel 62.2431 0.2683 849 DTI 2000 55347#1 MC 19/08/2000 Faroe-Shetland Channel 62.3091 0.2521 881 DTI 2000 55351#1 MC 19/08/2000 Faroe-Shetland Channel 62.3078 0.0973 939 DTI 2000 55352#1 MC 19/08/2000 Faroe-Shetland Channel 62.3847 0.0238 1061 DTI 2000 55354#1 MC 19/08/2000 Faroe-Shetland Channel 62.2915 0.1117 880 DTI 2000 55355#1 MC 19/08/2000 Faroe-Shetland Channel 62.2761 0.1267 829 DTI 2000 55372#2 MC 22/08/2000 North Shetland Slope 62.0458 0.2011 434 DTI 2000 55374#1 MC 22/08/2000 Faroe-Shetland Channel 62.308 0.1751 907 DTI 2000 55375#2 MC 22/08/2000 Faroe-Shetland Channel 62.325 0.1525 958 DTI 2000 55376#1 MC 22/08/2000 Faroe-Shetland Channel 62.3877 0.0985 1033 DTI 2000 55378#1 MC 23/08/2000 Faroe-Shetland Channel 62.3505 0.0927 1010 DTI 2000 55379#1 MC 23/08/2000 Faroe-Shetland Channel 62.3108 0.1323 934 DTI 2000 55386#1 MC 24/08/2000 Faroe-Shetland Channel 62.4005 0.586 791 DTI 2000 55387#1,2 MC 24/08/2000 Faroe-Shetland Channel 62.355 0.5367 781 DTI 2000 55395#1,2 MC 26/08/2000 Faroe-Shetland Channel 61.4198 − 3.116 1408 902 Mar Biodiv (2018) 48:897–914 of article-2; article-2 1.5 times longer than wide, 0.4 times as long as article-1, with outer pair of penicillate setae and single simple distal seta; article-3 1.3 times longer than wide, 0.7 times as long as article-2, with inner distal one simple seta and one penicillate seta; article-4 tapering, little longer than article-2, with five distal setae and one aesthetasc. Antenna (Fig. 2B, D) about 0.75 times as long as anten- nule; article-1 very short, naked; article-2 with two distal se- tae; article-3 as long as wide, with dorsodistal finely denticu- late seta; article-4 0.8 times as long as article-5 and as long as article-6, with ventrodistal penicillate seta; article-5 1.2 times longer than article-6, with two long ventrodistal, finely den- ticulate setae; article-6 more slender than article-5, with one distal seta; article-7 minute, with four distal setae. Labrum not recovered (but visible in whole view, Fig. 2B), conical. Left mandible (Fig. 2E) with six rounded teeth on pars incisiva; lacinia mobilis narrow, crenulate; pars molaris slender, distally bent downward, with rosette of numerous distal spinules. Right mandible (Fig. 2F) pars incisiva with three blunt teeth; pars molaris as in left mandible. Maxillule (Fig. 2G, G′) with nine terminal spines (three with fine den- ticulation, one plumose); palp with two plumose setae. Maxilla (Fig. 2H) larger than half of maxilliped plate, subtriangular. Labium (Fig. 2I) lobes narrow, distally setulose, outer lobe with microtrichia distally. Maxilliped (Fig. 2J) basis 1.8 times longer than wide, with single long seta reaching over distal margin of endite; endite distally with two small setae and two round tubercles; palp article-1 naked; article-2 with one outer distal finely denticulate and one simple and two inner distal finely denticulate setae; article-3 with two longer finely denticulate and one smaller simple inner distal setae; article-4 with subdistal simple seta and five distal finely denticulate setae. Epignath not recovered. Cheliped (Fig. 3A) attached via sclerite, with elongate basis 2.2 times longer than wide, posterior lobe as long as anterior mass, latter with one dorsal seta; merus subtriangular, with single ventral seta; carpus stout, 1.5 times longer than wide, with prominent carpal shield, well-developed rounded dorsoproximal extension, two mid-ventral setae and single dorsoproximal and dorsodistal setae; propodus large, just lon- ger but narrower than carpus, palm about as long as wide, dorsal and ventral margins smooth, with comb of three fine spines on inner surface near dactylus insertion; fixed finger with one seta near dactylus insertion, two setae ventrally and four teeth and three setae on cutting edge; dactylus with slight- ly undulate dorsal margin and two minute spines on cutting margin. Pereopod-1 (Fig. 3B) coxa with simple seta; basis 3.4 times longer than wide, narrow in most proximal part, naked; ischi- um with one ventral seta; merus one-third as long as basis, with one minute seta and robust ventrodistal finely denticulate spine that reaches beyond distal margin of carpus; carpus 1.2 times longer than merus, with numerous microtrichia and fine Table 1 (continued) Survey Sample Gear Date Locality Latitude (°N) Longitude Depth (m) B. Temp. (°C) Temp. (°C) Sediment SEAS4 57003#1&2 MC 01/07/2002 Faroe-Shetland Channel 62.2501 − 0.0663 803 88 μm mean grain size) SEAS4 57008#1–3 MC 01/07/2002 Faroe-Shetland Channel 62.0733 − 0.9671 897 116 μm SEAS4 57010#1&2 MC 01/07/2002 Faroe-Shetland Channel 62.2415 − 0.5503 893 108 μm SEAS4 57080#1 MC 01/07/2002 Faroe-Shetland Channel 62.1855 − 0.2955 700 181 μm SEAS4 57088#1 MC 01/07/2002 Faroe-Shetland Channel 62.1505 − 0.3171 650 227 μm Longitudes are °W, except positive values °E. B. Temp.-annual mean bottom temperature; Temp.–sampled bottom temperature Gear etc.: DS detritus sledge, MC megacorer, RP epibenthic sledge, Sn Sneli dredge Mar Biodiv (2018) 48:897–914 903 Fig. 2 Brixia aurora n. sp. female holotype (ZMH K-52920) A habitus dorsal view; B cephalothorax ventral view; C antennule; D antenna; E left mandible; F right mandible; G maxillule; G′ maxillule palp; H maxilla; I labium; J maxilliped. Scale bars 1 mm for A, 0.1 mm for C–J distal setulation, with single dorsodistal robust finely denticu- carpus, with microtrichia on ventral margin, with two late spine and two unequal (small and long) ventrodistal finely ventrodistal finely denticulate spines and two short dorsodistal denticulate spines; propodus 1.4 times longer than carpus, setae; dactylus denticulate on ventral margin, 1.4 times longer with microtrichia on ventral margin, with single ventrodistal than unguis, together 1.2 times longer than propodus. finely denticulate spine and dorsodistal fine seta; dactylus just Pleopods absent. shorter than unguis, together 0.9 times as long as propodus. Uropod (Fig. 3F) basis naked, as long as endopod segment- Pereopods 2 and 3 (Fig. 3C, D) similar to pereopod-1, but 1; exopod of two segments, 0.6 times as long as endopod pereopod-3 dactylus with accessory setae (possibly unob- segment-1; segment-2 with at least one short distal seta; served in pereopods 1–2). endopod of two segments; endopod segment-1 1.2 times as Pereopods 4 and 5 missing from dissected specimen. long as segment-2, that is armed with one seta distally Pereopod-6 (Fig. 3E) basis twice as long as wide, narrow (setation incomplete on specimen). distally, naked; ischium with one ventral seta; merus one third Remarks. Brixia aurora n. sp. differs from its congener (see as long as basis, with two ventrodistal finely denticulate spines below) by having a relatively long pleotelson—1.4 times lon- (one broken), that reach midway along carpus, and ventral ger than wide, a more slender antennule article-1, slight cren- microtrichia; carpus 1.7 times longer than merus, with ulation on the dorsal margin of the cheliped dactylus, only one microtrichia on ventral margin, and with three distal finely seta on the ischium of pereopod-6 (and possibly on pereopods denticulate spines (one broken) reaching one third along 4–5), no pleopods, and elongate uropod endopod segment-2, propodus and fine dorsodistal seta; propodus little shorter than that is 0.9 times as long as segment-1. 904 Mar Biodiv (2018) 48:897–914 Fig. 3 Brixia aurora n. sp. female holotype A cheliped; B pereopod- 1; C pereopod-2; D pereopod-3; E pereopod-6; F uropod. Scale bars 0.1 mm Distribution: Single record from the Iceland Basin, 1919.9– subrectangular, tapering towards anterior, 1.5 times longer 1925.6 m; see Fig. 1. than wide, with slight triangular rostrum, carapace with fine Brixia borealis n. sp. seta at each anterolateral corner, eyelobes poorly marked. Registered in ZooBank under: urn:lsid:zoobank.org: Pereonites sub-hexagonal, wider over pereopod insertions; act:0139C2E7-E1F7-4F7F-9FE7-8B556EE082F4 pereonite-1 tapering distally, shortest, 0.5 times as long as (Figs. 4, 5) cephalothorax, 1.3 times longer than wide; pereonites 2, 3 Material examined: holotype non-ovigerous female (3.3 mm and 4 subequal in length, 1.7 times longer than pereonite-1, long) dissected on slides (ZMH K-52921), st. 1149, 67° 50.57' 1.2 times longer than wide; pereonite-5 1.2 times longer than N 23° 41.85' W, depth 1246 m, AGT, 15 September 2011. pereonite-6; pereonite-6 just longer than pereonite-1, as wide Etymology. From the Latin borealis, northernorNorth. as long. Pleon as long as combined length of pereonites 5, 6 Diagnosis. Cephalothorax 1.5 times longer than wide. and two-thirds of pereonite-4; pleonites subequal in length, Pleotelson just longer than broad, shorter than last two pleonites. each pleonite 1.6 times wider than long, without lateral setae. Antennule article-1 2.7 times longer than wide, clearly shorter Pleotelson 1.1 times as long as wide, and shorter than last than distal three articles combined length. Pereopod-6 ischium two pleonites, slightly tapering distally, with pair of distal with two setae. Pleopods rudimentary. Uropod basis 1.5 times setae. longer than wide and 0.7 times as long as endopod segment-1; Antennule (Fig. 4A, C) about 0.9 times as long as cepha- exopod less than half as long as endopod segment-1. lothorax, article-1 2.7 times longer than wide, shorter than Description of female. Body (Fig. 4A, B) slender, 11.5 distal three articles together, with outer distal group of peni- times longer than wide, 3.3 mm long. Cephalothorax cillate setae and one simple distal seta shorter than article-2; Mar Biodiv (2018) 48:897–914 905 Fig. 4 Brixia borealis n. sp. female holotype (ZMH K-52921) A habitus dorsal view; B habitus lateral view; C antennule; D antenna; E labrum; F left mandible; G right mandible pars incisiva; H maxillule; H′ maxillule palp; I labium. Scale bars 1 mm for A, B, 0.1 mm for C–I article-2 2.3 times longer than wide, 0.7 times as long as denticulation, one plumose); palp (Fig. 4H′) with two plumose article-1, with outer single simple distal seta; article-3 0.7 setae. Maxilla (Fig. 5A) larger than half of maxilliped plate, times as long as wide, and 0.7 times as long as article-4, with subtriangular, distally setulose. Labium (Fig. 4I) lobes narrow, two small distal setae; article-4 tapering, 1.5 times longer than distally setulose, inner lobes with two distinct spines; outer article-3, with four distal setae and one aesthetasc. Blobe^ with microtrichia. Antenna (Fig. 4D) about 0.75 times as long as antennule; Maxilliped (Fig. 5B, B′) basis 1.8 times longer than wide, article-1 very short, naked; article-2 1.4 times longer than with single, long seta reaching over distal margin of endite; article-3, with one dorsodistal seta; article-3 as long as wide, endite distally with two rounded tubercles and one, fine seta; with dorsodistal seta; article-4 0.7 times as long as article-5 palp article-1 naked; article-2 with one outer distal and three and 0.8 times as long as article-6, with penicillate seta distally; inner distal setae (at least one of them finely denticulate); article-5 almost as long as article-6, with two simple and two article-3 with two short simple and two finely denticulate in- penicillate distal setae; article-6 more slender than article-5, ner setae; article-4 with subdistal seta and five finely denticu- with one distal seta; article-7 minute, with four distal setae. late distal setae. Epignath not recovered. Labrum (Fig. 4E) hood shaped, relatively elongate, finely Cheliped (Fig. 5C) attached via elongate sclerite reaching setose. Left mandible (Fig. 4F) pars incisiva with three teeth; to posterior of cephalothorax inserted to dorsum of basis; basis lacinia mobilis narrow, weakly crenulate dorsally; pars elongate, 2.3 times longer than wide, posterior lobe as long as molaris slender, distally bent downwards, with rosette of nu- anterior mass; merus subtriangular, with fine ventral seta; car- merous short and two long distal spinules. Right mandible pus stout, almost as long as wide, with well-developed round- (Fig. 4G) pars incisiva with several pointed teeth. Maxillule ed dorsoproximal extension and prominent carpal shield, with two fine mid-ventral setae and single dorsoproximal and (Fig. 4H) with nine terminal spines (three with fine 906 Mar Biodiv (2018) 48:897–914 Fig. 5 Brixia borealis n. sp. female holotype A maxilla; B maxillipedal palp and endite; B′ maxillipedal basis; C cheliped; D pereopod-1; E pereopod-2; F pereopod-4; G pereopod-6; H uropod. Scale bars 0.1 mm dorsodistal setae (one broken); propodus large, longer than ventrodistal finely denticulate robust spines; propodus 1.4 wide but shorter than carpus, palm about as long as wide, times longer than carpus, with microtrichia on ventral margin dorsal and ventral margins smooth, with comb of three fine and with single ventrodistal finely denticulate spine; dactylus spines on inner surface near dactylus insertion; fixed finger 0.8 times as long as unguis, ventrally denticulate, together 0.9 with one seta near dactylus insertion, two setae ventrally and times as long as propodus. three teeth and three setae on cutting edge; dactylus dorsal Pereopod-2 (Fig. 5E) similar to pereopod-1, but carpus margin smooth, with two minute spines on cutting margin. with additional ventrodistal spine. Pereopod-1 (Fig. 5D) coxa with simple seta (not shown on Pereopod-3 (not figured) as pereopod-2. figure); basis 2.6 times longer than wide, narrow in most distal Pereopod-4 (Fig. 5F) basis twice as long as wide, narrow in part, with long penicillate, dorsal seta; ischium with one ven- most distal part, apparently with two dorsal penicillate setae; tral seta; merus one-third as long as basis, with one robust ischium with two ventral setae; merus one third as long as basis, ventrodistal finely denticulate spine that reaches beyond distal with two ventrodistal finely denticulate spines, which reach margin of carpus; carpus 1.2 times longer than merus, with beyond midway on carpus; carpus 1.6 times longer than merus, numerous microtrichia, with single dorsodistal and with microtrichia on ventral margin, and with three distal finely Mar Biodiv (2018) 48:897–914 907 denticulate spines, the longest spine about 0.6 times as long as 53833#1; one specimen, AFEN 1998 54509#2; one specimen, propodus and dorsodistal seta; propodus as long as carpus, with AFEN 1998 54521#1,3; one specimen, AFEN 1998 54522#1,2; microtrichia on ventral margin, with two ventrodistal finely two specimens, AFEN 1998 54524#2; one specimen, BIOICE denticulate spines and one short dorsodistal seta; dactylus as 2010; 11 specimens, BIOICE 2011; four non-ovigerous fe- long as propodus, denticulate on ventral margin; unguis about males, BIOICE 2030; one non-ovigerous female, one ovigerous as long as half of dactylus, with ventral denticulation. female, BIOICE 2077; two non-ovigerous females, BIOICE Pereopod-5 (not figured) as pereopod-4. 2088; two manca-3, four non-ovigerous females, BIOICE Pereopod-6 (Fig. 5G) similar to pereopod-4, but propodus 2315; six non-ovigerous females, two ovigerous females, one with two dorsodistal setae. post-ovigerous female, BIOICE 2317; one manca-3, 12 non- Pleopods (Fig. 4B) biramous, rudimentary, without setae. ovigerous females, four ovigerous females, one preparatory Uropod (Fig. 5H) basis 0.7 times as long as endopod seg- male (uncertain?), BIOICE 2318; one ovigerous female, ment-1, naked; exopod 0.4 times as long as endopod segment- BIOICE 2323; one non-ovigerous female, one ovigerous fe- 1; segment-1 with distal seta; segment-2 with one long and male, two post-ovigerous females, BIOICE 2364; one non- one short distal setae; endopod segment-1, five times as long ovigerous female, one ovigerous female, BIOICE 2638; one as wide, twice as long as segment-2, with one penicillate seta preparatory male (?), BIOICE 2743; one non-ovigerous female, distally; segment-2 with one subdistal and two distal setae. BIOICE 2749; three non-ovigerous females, one non-ovigerous Remarks. In contrast to B. aurora,in B. borealis n. sp., the female, BIOICE 2762; one specimen, BIOICE 2765; one pleotelson is shorter, (but still as long as or just longer than broad), the cheliped dactylus is apparently smooth, the ischi- um of pereopods 4–6 has two setae, and the uropod endopod segment-2 is about half as long as segment-1. Distribution: Single record from the Denmark Strait, 1246 m; see Fig. 1. Parakanthophoreus Larsen & Araújo-Silva, 2014 Parakanthophoreus catharina n. sp. Registered in ZooBank under: urn:lsid:zoobank.org: act:62B32816-9167-4CB9-82A9-C69AD931AE52 (Figs. 6–10) Akanthophoreus inermis (Hansen, 1913): Bird (2001): 32–33. Material examined: holotype non-ovigerous female (2.8 mm long) (NMS.Z.2018.26.1), DTI2000 st. 55283#1, 61° 35.06′ N 1° 58.43′ W, depth 1262 m, Megacore, 5 August 2000. Allotype preparatory male (2.5 mm long) (NMS.Z.2018.26.2), partly dissected on microslide (NMS.Z.2018.26.3), DTI2000 st. 55374#1, 62° 18.48′ N0° 10.51′ E, depth 907 m, Megacore, 22 August 2000. Paratype female dissected on slides (ZMH K-52922), st. 853/ 1132-1, 67° 38.39′ N 26° 45.46' W-67° 38.28′ N 26° 45.72′ W, depth 316.6–316.8 m, EBS, 14 September 2011. Other material taken during IceAGE project: one non-ovigerous female (2.4 mm long), one manca (1.3 mm long), st. 853/1116-1, 67° 12.82′N26° 16.31′ W, depth 683.1 m, GKG, 14 September 2011; one non- ovigerous female (1.6 mm long), one manca (1.3 mm), st. 853/ 1129-1, 67° 38.77′ N 26° 44.78′ W, depth 320.6 m, GKG, 14 September 2011; four non-ovigerous females (1.4, 1.6, 2.3, and 2.3 mm long), one manca (1.2 mm long), st. 853/1141-1, 67° 50.22′ N 23° 42.11′ W, depth 1241.6 m, GKG, 15 September 2011; one female with marsupium, one non-ovigerous female (1.7 mm long), one manca (0.9 mm long), st. 873-4, 61° 46.52′ N 03° 52.51′ W, depth 833.8 m, Van Veen grab, 28 July 2013. Fig. 6 Parakanthophoreus catharina n. sp. (NMS.Z.2018.26.1) habitus Supplementary material and records examined—see Table 1 A female holotype dorsal view; B male allotype (NMS.Z.2018.26.2) for station data: one specimen, AFEN 1996 53819#3; one spec- dorsal view; C male antennule (NMS.Z.2018.26.3). Scale bars 1 mm imen, AFEN 1996 53828#1; one specimen, AFEN 1996 for A, B and 0.1 for C 908 Mar Biodiv (2018) 48:897–914 specimen, BIOICE 3023; one non-ovigerous female, BIOICE 55352#1; one non-ovigerous female, DTI2000 55354#1; two 3108; four specimens, two non-ovigerous females, two prepara- non-ovigerous females, DTI2000 55355#1; one non-ovigerous tory males (?), BIOICE 3115; two specimens, BIOICE 3121; female, DTI2000 55374#1; two non-ovigerous females, one specimen, BIOICE 3198; one specimen, BIOICE 3225; DTI2000 55375#3; one non-ovigerous female, DTI2000 three non-ovigerous females, four specimens, BIOFAR 167; 55376#1; one non-ovigerous female, DTI2000 55378#1; one six specimens, BIOFAR 168; one specimen, BIOFAR 169; non-ovigerous female, DTI2000 55379#1; one non-ovigerous two specimens, BIOFAR 170; seven specimens, BIOFAR female, DTI2000 55395#1; two non-ovigerous females, SEAS4 172; seven specimens, BIOFAR 478; six specimens, BIOFAR 57003#1&2; two non-ovigerous females, two post-ovigerous 769; four non-ovigerous females, DTI2000 55283#1; three non- females, SEAS4 57008#1-3; one specimen, SEAS4 57080#1; ovigerous females, DTI2000 55322#1; three non-ovigerous fe- one non-ovigerous female, SEAS4 57088#1. males, DTI2000 55324#; one non-ovigerous female, DTI2000 Etymology. Catharina (Lat.) (= Katarzyna in Polish): the 55336#1; three non-ovigerous females, DTI2000 55344#1; one species is dedicated to the senior author’s wife, Katarzyna preparatory male (?), DTI2000 55344#2; one non-ovigerous Pawicka-Jóźwiak. female, DTI 2000 55347#1; one preparatory male (?), Diagnosis. Body about ten times longer than wide. DTI2000 55351#1; two non-ovigerous females, DTI2000 Pereonite-1 as long as wide. Maxilliped basis with setae. Fig. 7 Parakanthophoreus catharina n. sp. female (ZMH K-52922) A antennule; B antenna; C labrum; D left mandible; E right mandible; F maxillule; G maxilla; H labium; I maxilliped; J epignath. A–D, F–H and J female paratype; E and I preparatory male allotype (NMS.Z.2018.26.3), DTI2000 Stn 55374#1. Scale bars 0.1 mm for B–D, F, H and J and 0.25 for A, E and I Mar Biodiv (2018) 48:897–914 909 Maxilliped endite with pair of distal setae. Cheliped carpus, than pleonites 4–5, with rounded apex. Cephalothorax and propodus and dactylus without crenulation but carpus with tergites of pereonites and pleonites with dense pitting. deep shield; cheliped carpus with single ventral seta; fixed Antennule (Figs 6A, 7A) about 0.75 times as long as ceph- finger ventral setae unequal in thickness. Pereopods 1–3car- alothorax; article-1, twice as long as wide, with one simple pus with dorsodistal seta. Pereopods 4–6 ischium with pair of and two penicillate setae on outer distal margin; article-2 setae and dactylus with denticulate ventral margins. Pereopod- about half as long as article-1, with one short inner distal seta 6 propodus with three dorsodistal finely denticulate spines. and one long outer distal seta; article-3 half as long as article- Uropod exopod 0.7 times as long as endopod segment-1. 2, with one short inner distal seta, one longer outer distal seta Description of non-ovigerous female. Body (Fig. 6A) slen- and some distal spinules; article-4 1.5 times longer than arti- der, 10.1 times longer than wide, 2.8 mm long (range 1.24– cle-3, with two simple setae distally; article-5 minute, with 2.98 mm, n = 70 G. Bird material). Cephalothorax 15% of total four simple setae distally. Articles with dense pitting. body length, about 1.5 times longer than wide, carapace with Antenna (Figs 7B, 9A) article-1 very short, naked; article-2 fine seta at each anterolateral corner, eyelobes poorly marked. just longer than wide, with two dorsodistal and distolateral Pereonites parallel-sided, length/width ratio: 1, 1.1, 1.4, 1.4, setae; article-3 0.7 times as long as wide, with one simple 1.1, and 0.9. Pleon 18% of total body length, pleonites distal seta; article-4 3.8 times longer than wide, with fusion subequal in length, epimera with single seta. Pleotelson longer line, one penicillate seta near fusion line, one penicillate seta Fig. 8 Parakanthophoreus catharina n. sp. female paratype (ZMH K-52922) A cheliped; B pereopod-1; C pereopod-2; D pereopod-3; E pereopod-4; F pereopod-5; G pereopod-6; H pleopod basis and endopod; H' pleopod exopod; setae all plu- mose but not shown for clarity; I uropod. Scale bars 0.1 mm 910 Mar Biodiv (2018) 48:897–914 Fig. 9 Parakanthophoreus catharina n. sp. cuticle ornamentation using SE microscope A antenna article-4; B cheliped basis and sclerite; C pereopod-1; D uropod. Scale bars 0.05 mm for A and C, 0.02 for B and D subdistally (partly broken), one minute and four simple setae Cheliped (Figs 8A, 9B, 10A–D) attached via large distally; article-5 just longer than half of article-4, with two posterodorsal sclerite; posterior lobe and anterior mass subequal simple setae distally; article-6 minute, with five simple setae in size, naked; merus with one simple ventral seta; carpus with distally. Articles with dense pitting. well-developed rounded dorsoproximal extension and deep Labrum (Fig. 7C) trapezoidal in lateral view, distally setose. shield, one simple midlength seta ventrally and one simple seta Left mandible (Fig. 7D) pars incisiva with five blunt denticles; dorsodistally; propodus about half as long as wide, palm about as lacinia mobilis weakly crenulate dorsally; pars molaris long as wide, dorsal and ventral margins smooth, with simple narrowing distally, with rosette of numerous distal spinules. seta near dactylus insertion, and one longer and three short finely Right mandible (Fig. 7E) pars incisiva with three denticles dis- denticulate spines on inner margin, inner and outer margins with tally; pars molaris as in left mandible. Maxillule (Fig. 7F) endite microtrichia, outer margin additionally with protrusions; fixed with some inner microtrichia, subdistally with setae, with eight finger with two denticles and three simple setae on cutting edge, terminal spines (at least three finely denticulate); palp not ob- and two setae unequal in thickness ventrally; dactylus as long as served. Maxilla (Fig. 7G) triangular, inner and distal margin fixed finger, naked. Articles with dense pitting (Figs 9B, 10). setose. Labium (Fig. 7H) simple, slender, distally with spinules. Pereopod-1 (Figs 8B, 9C) coxa with simple seta (not Maxilliped (Fig. 7I) basis with one distal seta, half as long shown on figure); basis 1.9 times longer than wide, naked; as endite; endite with distal tubercle, pair of distal setae and ischium with one seta; merus about as long as carpus, with outer distal setules; palp article-1 naked; article-2 with one ventral microtrichia, distal seta, and ventrodistal finely dentic- simple outer seta and one plumose and two simple setae on ulate spine; carpus with one complex-tip seta and finely den- inner distal margin; article-3 with two simple short, and two ticulate spine dorsodistally, one finely denticulate spine long setae on inner margin; article-4 with one subdistal seta ventrodistally and numerous microtrichia along article; and five distal setae. Epignath (Fig. 7J) slender, tapering dis- propodus 1.4 times longer than carpus, with ventrodistal spine tally to the pointed tip. and ventral microtrichia; dactylus and unguis combined 0.6 Mar Biodiv (2018) 48:897–914 911 Fig. 10 Parakanthophoreus catharina n. sp. cheliped ornamentation of using SE microscope A outer surface; B propodus outer surface; C chela inner surface; D carpus outer surface. Scale bars, respectively, 0.1, 0.02, 0.05, and 0.01 mm times as long as propodus; unguis 1.7 times longer than Pleopod (Fig. 8H, H') basis shorter than ramus, with dactylus. Articles of all pereopods with dense pitting. ventrodistal seta; endopod with 12 setae along ventral and distal Pereopod-2 (Fig. 8C) similar to pereopod-1, but basis with margins; exopod with eight distal setae. All setae plumose. long penicillate seta dorsally and carpus with additional Uropod (Figs 8I, 9D) basis 0.8 times as long as exopod, ventrodistal finely denticulate spine. naked; exopod of two segments, 0.7 times as long as endopod Pereopod-3 (Fig. 8D) similar to pereopod-2, but basis without segment-1, segments subequal; segment-1 with short seta; penicillate seta (possibly lost) and dactylus with proximal seta. segment-2 with one short and one long setae; endopod of Pereopod-4 (Fig. 8E) basis 2.4 times longer than wide, two equal segments; segment-1 with one simple and three narrower than those of pereopods 1–3, with one penicillate penicillate setae distally; segment-2 with one long midlength midlength seta ventrally; ischium with two setae; merus one seta and one penicillate and four simple setae distally. third of basis length, with two finely denticulate spines and Manca-3. Length 1.03–1.05 mm (n =3). microtrichia ventrodistally; carpus 1.3 times longer than merus, Ovigerous female. As female (described above) but more with one complex-tip seta and one finely denticulate spine dorsoventrally compressed, with oostegites; length 1.93– dorsodistally, two finely denticulate spines ventrodistally and 2.51 mm (n =10). some ventral microtrichia; propodus just longer than carpus, Post-ovigerous female. As female (described above) but with one finely denticulate spine dorsodistally, two finely den- dorsoventrally compressed, lacking oostegites, but with scars ticulate spines ventrodistally and ventral microtrichia; dactylus or remnants thereof; length 2.15–2.35 mm (n =5). and unguis combined 1.3 times longer than propodus, both Preparatory male. As female (described above) but with denticulate ventrally; unguis about half as long as dactylus. proportionately larger antennules, with stouter articles 1–2; Pereopod-5 (Fig. 8F) as pereopod-4. length 1.79–3.00 mm (n = 6). Body (Fig. 6B) slender, 10.3 Pereopod-6 (Fig. 8G) as pereopod-4, but propodus with times longer than wide, 2.8 mm long. Cephalothorax 14% of three dorsodistal finely denticulate setae. total body length, about 1.5 times longer than wide, carapace 912 Mar Biodiv (2018) 48:897–914 Fig. 11 Parakanthophoreus inermis near-topotypical BIOICE Stn 2629 A female habitus; B right cheliped; C pereopod-1; D pereopod-6. Scale bars 1 mm for A, 0.25 mm for B–D with fine seta at each anterolateral corner, eyelobes poorly (about ten times longer than wide), maxilliped basis with seta, marked. Pereonites parallel-sided, length/width ratio: 0.8, maxilliped endites with a pair of distal setae, cheliped carpus, 1.1, 1.1, 1.0, 0.9, and 0.6. Pleon 18% of total body length, propodus, and dactylus without crenulation, but carpus with deep pleonites subequal in length, epimera with single seta. shield, pereopods 4–6 ischium with pair of setae and dactylus Pleotelson as long as pleonites 3–5 and half of pleonite-2, with with denticulate ventral margins, uropod exopod distinctly longer rounded apex, terminated with two pairs of setae. than half of endopod segment-1. Another useful characteristic Antennule (Fig. 6B, C) about 0.75 times as long as cepha- distinguishing P. catharina from its congeners, but also from lothorax; article-1, twice as long as wide, with one simple and other akanthophoreids, is the setation of the cheliped fixed finger, five penicillate setae on outer distal margin; article-2 0.6 times where the two ventral setae are of unequal thickness. as long as article-1, with one long and three penicillate outer Of the described species of Parakanthophoreus known from distal setae; article-3 0.7 times as long as article-2, with one the general area of the IceAGE project—P. alba (Hansen, 1913), short and one longer inner distal setae; article-4 just shorter than P. brachiatus (Hansen, 1913), P. inermis (Hansen, 1913), article-2, with four simple setae distally and one aesthetasc. P. longiremis (Lilljeborg, 1864), P. multiserratus (Hansen, 1913), Remarks. Parakanthophoreus catharina n. sp. differs from and P. vikingra (Błażewicz-Paszkowycz & Bamber, 2011)—the other members of the genus by a combination of: elongated body most likely species to be confused with P. catharina is P. inermis, Mar Biodiv (2018) 48:897–914 913 Ethical approval All applicable international, national, and/or institution- since it can be sympatric, and most of the non-IceAGE records of al guidelines for the care and use of animals were followed by the authors. P. catharina were originally assigned, provisionally, to that spe- cies. A review and comparison of these species (except Field study Permits and approval of field or observational studies are P. vikingra) was given by Guerrero-Kommritz (2004), while not applicable for authors. Hansen (1913) is still a good reference for species comparisons. Open Access This article is distributed under the terms of the Creative Parakanthophoreus inermis was characterized by Hansen Commons Attribution 4.0 International License (http:// (1913) by having a relatively short antennule article-1 compared creativecommons.org/licenses/by/4.0/), which permits unrestricted use, to other similar species, as well as a deep cheliped carpal shield distribution, and reproduction in any medium, provided you give appro- priate credit to the original author(s) and the source, provide a link to the and a short claw on pereopods 1–3, all of which are shared by Creative Commons license, and indicate if changes were made. P. catharina but not by the other Parakanthophoreus species mentioned above. Redescriptions of P. inermis by Guerrero- Kommritz (2004)and Sieg (1986b) were unhelpful in clarifying References the nature of this species because of several factors such as un- clear attribution of material, poor illustrations, or use of non- Bird GJ (2001) Tanaidacea of the Atlantic Margin: the AFEN surveys of topotypical specimens. Indeed, the taxon dealt with by Sieg 1996 and 1998. Unpublished report to the Atlantic Frontier may be conspecific with, or at least clearly related to, Environmental Network and UK Offshore Oil Association, P. catharina rather than P. inermis and it is possible that some February 2001 Bird GJ (2007) Family incertae cedis [sic]. In: Larsen K, Shimomura M of Hansen’s material may too be of both species. A revision of (eds) Tanaidacea (Crustacea: Peracarida) from Japan III. The deep P. inermis is in progress (Bird, in prep.). trenches: the Kurile-Kamchatka Trench and Japan Trench. Magnolia Examination of near-topotypical specimens of Press, Auckland, pp 121–149 Parakanthophoreus inermis (Fig. 11) indicates that it can be sep- Bird GJ (2015) Tanaidacea (Crustacea: Peracarida) of the Northeast arated from P. catharina by, inter alia, its shorter cephalothorax, Atlantic: Chauliopleona Dojiri and Sieg, 1997 and Saurotipleona n. gen. from the ‘Atlantic Margin’.JNatHist49:1507–1547. https:// shorter pereonites (especially pereonite-1, Fig. 11A)–none longer doi.org/10.1080/00222933.2015.1005715 than wide, weaker (but still relatively deep) cheliped carpal shield Błażewicz-Paszkowycz M, Bamber R (2011) Tanaidomorph Tanaidacea (Fig. 11B), lack of a distodorsal seta on the carpus of pereopods 1– (Crustacea: Peracarida) from mud-volcano and seep sites on the 3(Fig. 11C; very conspicuous in P. catharina), and only two Norwegian margin. Zootaxa 3061:1–35 Błażewicz-Paszkowycz M, Bamber RN, Cunha MR (2011) New dorsodistal spines on the propodus of pereopod-6 (Fig. 11D; three tanaidomorph Tanaidacea (Crustacea: Peracarida) from submarine in P. catharina); the cuticular ornamentation is also less pro- mud-volcanoes in the Gulf of Cadiz (North-East Atlantic). Zootaxa nounced, without the pitting seen in P. catharina.Other 2769:1–53 akanthophoreids in the Iceland-Faroe region with large cheliped Błażewicz-Paszkowycz M, Bamber RN, Jóźwiak P (2013) Tanaidaceans (Crustacea: Peracarida) from the SoJaBio joint expedition in slope carpal shields include Chauliopleona hastata (Hansen, 1913) and and deeper waters in the Sea of Japan. Deep-Sea Res II 111:325– Saurotipleona julii Bird, 2015 but these can be distinguished, at 332. https://doi.org/10.1016/j.dsr2.2014.08.021 least, by their ventral spur on pleonite-5. Brix S, Bauernfeind W, Brenke N, Blazewicz M, Borges V, Buldt K, Distribution: BIOICE records; 23, from the Greenland- Cannon J, Díaz Agras G, Fiege D, Fiorentino D, Haraldsdóttir S, Iceland Rise, Iceland Plateau, Iceland-Faroes Rise (all but Hoffmann S, Holst S, Hüttmann F, Jeskulke K, Jennings R, Kocot K, Khodami S, Lucas Rodriguez Y, Martinez Arbizu P, Meißner K, one from northern flank), Kolbeinsey Ridge, and Norwegian Mikkelsen N, Miller M, Murray A, Neumann H, Ostmann A, Riehl Basin, 326–1994 m, in temperatures < 1 °C. Other records, 53 T, Schnurr S, Savavarsson J, Yasuhara M (2011) Cruise Report records from the Faroe-Shetland Channel, Faroe-Norway M85/3 IceAGE1 2011 with RV Meteor. Reykjavik, Cuxhaven Rise, North Shetland Slope, and Norwegian Basin, 434– Brix S,MeißnerK,StranskyB,Halanych KM, Jennings RM, Kocot KM, Svavarsson J (2014) The IceAGE project a follow up of BIOICE. Pol 1513 m (Table 1; Bird 2001 and ined.) (see Fig. 1). Polar Res 35:141–150. https://doi.org/10.2478/popore-2014-0010 Coleman CO (2003) Digital inking: how to make perfect line drawings on Acknowledgements Authors would like to thank Saskia Brix for making computers. Org Divers Evol 14:1–14 the material available for study and the crew of R/V Meteor and R/V Dojiri M, Sieg J (1997) The Tanaidacea. In: Blake JA, Scott PH (eds) Poseidon as well as the scientists who participated in IceAGE Program. Taxonomic atlas of the benthic fauna of the Santa Maria Basin and We are also grateful for two anonymous reviewers for their valuable Western Santa Barbara Channel. Volume 11. The Crustacea Part 2. comments. Studies were financed by Polish National Science Centre The Isopoda, Cumacea and Tanaidacea. Santa Barbara Museum of grant UMO-2014/13/B/NZ8/04702. Natural History, Santa Barbara, CA, pp 181–278 Guerrero-Kommritz J (2004) A revision of the genus Paraleptognathia Funding This study was funded by the Polish National Science Centre Kudinova-Pasternak, 1981 (Crustacea: Tanaidacea) and description grant (UMO-2014/13/B/NZ8/04702). of four new species. Zootaxa 481:1–63 Hansen HJ (1913) Crustacea, Malacostraca. II. IV. The order Tanaidacea. Compliance with ethical standards Danish Ingolf Expedit 3:1–145 Kudinova-Pasternak RK (1981) Tanaidacea. In: Kusnecov AP, Mironov AN Conflict of interest The authorsdeclare thattheyhave noconflictofinterest. (eds) Bentos podvodnych gor Markus-Nekker i smeznych rajonov 914 Mar Biodiv (2018) 48:897–914 tichogo okeana [Benthos of the Submarine Mountains Markus Necker Larsen K, Wilson GDF (2002) Tanaidacean phylogeny, the first step: the superfamily Paratanaidoidea. J Zool Syst Evol Res 40:205–222 and adjacent Pacific regions]. Institut Okeanologii, Moscow, pp 94–112 Kudinova-Pasternak RK (1985) Tanaidacea (Crustacea, Malacostraca) Sieg J (1986a) Tanaidacea (Crustacea) von der Antarktis und collected on the summit and at foot of Great-Meteor Seamount. T Subantarktis. II. Tanaidacea gesammelt von Dr. JW. Wägele I Ocean 120:52–64 wahrend der Deutschen Antarktis Expedition 1983. Zoologischen Larsen K (2005) Deep-sea Tanaidacea (Peracarida) from the Gulf of Museum der Universität Kiel 2:1–80 Mexico. Brill, Leiden Sieg J (1986b) Crustacea Tanaidacea of the Antarctic and the Larsen K, Araújo-Silva CL (2014) The ANDEEP Tanaidacea (Crustacea: Subantarctic. 1. On material collected at Tierra del Fuego, Isla de Peracarida) revisited III: the family Akanthophoreidae. Zootaxa los Estados, and the west coast of the Antarctic Peninsula. In: 3796:237–264. https://doi.org/10.11646/zootaxa.3796.2.2 Korniker LS (ed.) Biology of the Antarctic Seas 18. Volume 45 in Larsen K, Błażewicz-Paszkowycz M, Cunha MR (2006) Tanaidacean the Antarctic Research Series. American Geophysical Union, (Crustacea: Peracarida) fauna from chemically reduced habitats— Washington, D.C., pp 1–180 the lucky strike hydrothermal vent system, mid-Atlantic [sic] ridge. Zootaxa 1187:1–36 http://www.deepdyve.com/assets/images/DeepDyve-Logo-lg.png Marine Biodiversity Springer Journals

A new genus of family Akanthophoreidae and new species of genus Parakanthophoreus Larsen & Araújo-Silva, 2014 (Crustacea: Tanaidacea: Tanaidomorpha) from the North Atlantic

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Springer Journals
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Copyright © 2018 by The Author(s)
Subject
Life Sciences; Biodiversity; Freshwater & Marine Ecology; Plant Systematics/Taxonomy/Biogeography; Animal Systematics/Taxonomy/Biogeography
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1867-1616
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1867-1624
DOI
10.1007/s12526-018-0866-9
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Abstract

Based on tanaidacean material collected primarily during the IceAGE cruises in the North Atlantic, three new species from the family Akanthophoreidae are described; one is classified in the genus Parakanthophoreus, and the other two represent a new genus—Brixia n. gen. The main characters that distinguish Brixia n. gen. from other akanthophoreids is the lack of fully developed pleopods in adult females and a seven-articled antenna. The diagnosis of the family Akanthophoreidae is amended. Cheliped ornamentation of Parakanthophoreus catharina n. sp. is illustrated using SEM pictures. . . . . . Keywords IceAGE Icelandic waters Brixia n. gen. BIOICE BIOFAR AFEN Introduction in Akanthophoreinae were moved later to other families (Larsen and Wilson 2002), e.g., Araphura Bird & Holdich, Sieg (1986a) proposed the tanaidacean subfamily 1984 and Araphuroides Sieg, 1986 to Tanaellidae, and Akanthophoreinae (Anarthruridae) to accommodate Filitanais Kudinova-Pasternak, 1973 to Colletteidae; other paratanaoids characterized by, e.g., a four-articled antennule, genera were placed in the family incertae sedis, e.g., six-articled antenna, mandibular molar usually pointed, at Andrognathia Sieg, 1983, Leptognathioides Bird & Holdich, least the carpus of pereopod-1 with spines, pleopods well- 1984, and Robustochelia Kudinova-Pasternak, 1983. developed, reduced or absent, and uropods with one or two The Akanthophoreinae was rediagnosed and elevated to rami. The broad diagnosis resulted in a prolific composition of family level by Błażewicz-Paszkowycz and Bamber (2011) this subfamily and in a short time it appeared to be a non- with only the genus Akanthophoreus Sieg, 1986 included. monophyletic taxon. Most of the genera originally included Later, Błażewicz-Paszkowycz et al. (2013) added Chauliopleona Dojiri & Sieg, 1997 to this family, but without giving a justification for this reclassification. The most recent This article is registered in ZooBank under: urn:lsid:zoobank.org:pub: revision of this family was by Larsen and Araújo-Silva (2014), 6D7D4138-0170-44BD-BCD9-48B9983B88E1 who, with a phylogenetic approach, accommodated within This article is part of the Topical Collection on Biodiversity of Icelandic Akanthophoreidae a few other genera: Mimicarhaphura Sieg, Waters by Karin Meißner, Saskia Brix, Ken M. Halanych, and Anna 1986; Paraleptognathia Kudinova-Pasternak, 1981; Jazdzewska. Stenotanais Bird & Holdich, 1984; Tumidochelia Knight, Communicated by S. Brix Larsen & Heard, 2003; and the newly described Parakanthophoreus Larsen & Araújo-Silva, 2014. Most re- * Piotr Jóźwiak cently, Bird (2015) added Saurotipleona to this list of [email protected] akanthophoreids. One of the new taxa described below, a Parakanthophoreus species, has a strong resemblance to the Laboratory of Polar Biology and Oceanobiology, University of Łódź, genus Paraleptognathia Kudinova-Pasternak, 1981, which has Banacha 12/16, 90-237 Łódź, Poland been systematically confusing since its first diagnosis (Bird EcoAnalysts, Inc., Moscow, ID 83843, USA 2007;Błażewicz-Paszkowycz and Bamber 2011;Larsenand Independent, Kāpiti, Waikanae, New Zealand Araújo-Silva 2014). 898 Mar Biodiv (2018) 48:897–914 Larsen (2005) pointed out that in deep-sea tanaidomorphan The material is deposited in the Zoological Museum, Hamburg taxa females may lack pleopods. It was previously recorded and the National Museums of Scotland (NMS), Glasgow. for, e.g., all genera in the family Agathotanaidae; Caudalonga Larsen, 2005; Cetiopyge Larsen & Heard, 2002 and Collettea Lang, 1973 from the Colletteidae, and Anarthrura Sars, 1882 Systematics from the Anarthruridae. Until now, the Akanthophoreidae was represented exclusively by taxa with well-developed pleopods Suborder Tanaidomorpha Sieg, 1980 but a new genus described below is with reduced pleopods or Superfamily Paratanaoidea Lang, 1949 completely without pleopods. Family Akanthophoreidae Sieg, 1986 In this paper, we deal with a series of akanthophoreids Diagnosis see Larsen and Araújo-Silva (2014) but with collected in the waters off Iceland during the IceAGE additions: all pereopod meri and carpi with slender spines. (Icelandic Animals Genetic and Ecology; Brix et al. 2014) Pleopods well-developed, reduced or absent. initiative, which allows us to erect a new genus with two Remarks. The diagnosis of the family is modified primarily new species and describe a Paraleptognathia-like species of regarding the pleopods, where they may be absent or rudimen- Parakanthophoreus. tary in the new genus described below. Among the original members of the family, formerly the Tribe Akanthophoreinae within the family Anarthruridae Material and methods Lang, 1971, Paraleptognathia was distinguished by a combi- nation of: a five-articled antennule, with two aesthetascs on the Samples were collected during two IceAGE cruises. The first terminal article, a maxillule with eight spines, an epignath took place in August/September of 2011 on board the R/V tipped with a seta, the cheliped with a strong carpal shield Meteor and the second in August 2013 on board the R/V (and all cheliped articles covered with setules), and a pleopod Poseidon. During those initiatives, various sampling gears exopod of two articles. The unique five-articled antennules in were deployed, e.g., Van Veen grab, Shipek grab, epibenthic the type species of the genus, Paraleptognathia typica sledge, or box corer (Brix et al. 2014). Reference is also made Kudinova-Pasternak, 1981 (Kudinova-Pasternak 1981) were to material collected during the BIOICE, BIOFAR, and AFEN later discussed by Guerrero-Kommritz (2004) and were treated projects from the same, and adjacent, areas (e.g., Bird 2001, as a deviation of the standard state or a unique character of the 2015)(Fig. 1). For environmental data, please refer to Bird species, especially since that P. bacescui Kudinova-Pasternak, (2015), Brix et al. (2011), and Table 1 herein. 1985 has standard four-articled antennules (Kudinova- Specimens were dissected using chemically sharpened Pasternak 1985). This character is known to be a feature of tungsten needles, mounted in glycerin on slides and sealed immature or Bpreparatory^ males in several paratanaoid groups with nail varnish. Drawings were initially prepared using a (Bird 2007,Błażewicz-Paszkowycz et al. 2011). For over a microscope combined with a camera lucida and redrawn on decade, Paraleptognathia was known only from two species, a digital tablet as proposed by Coleman (2003). butDojiriand Sieg (1997) described P. bisetulosa and moved The body length to width ratio was calculated using mea- Akanthophoreus gracilis (Krøyer, 1842) to Paraleptognathia. surements from the tip of the carapace to the end of the Guerrero-Kommritz (2004) synonymized Akanthophoreus pleotelson, and of the widest part of carapace. The length with Paraleptognathia, resulting in a total of 17 species classified and width of articles were measured along their central axes. within this genus. This was not accepted by Bird (2007)and it Dehydration for SE microscopy was carried out sequential- was again split into Akanthophoreus, Parakanthophoreus,and ly in the tissue processor (Leica EMTP) with 95% and abso- Paraleptognathia by Larsen and Araújo-Silva (2014). Although lute ethanol. Specimens were then transferred to a critical the morphology of one of the new akanthophoreid species found point dryer (Leica EMCPD 300) with parameters set as fol- in the Northeast Atlantic during IceAGE cruises corresponds with lows: mixing 50%, CO in–slow with 2 min delay, exchange the main characters defining Paraleptognathia suggested by Bird speed–5with 20 cycles, gas out heat and speed–fast. Dried (2007) and Larsen and Araújo-Silva (2014), namely a large carpal specimens were transferred on coverslips and coated with 6- shield and cheliped articles with Bornamentation,^ it is considered nm-thick layer of gold (Leica EM ACE200). SEM pictures that Paraleptognathia is too ill-defined and poorly supported by were made on Phenom ProX. type material to warrant the addition of new species. The morphological terminology follows that proposed by Brixia n. gen. Błażewicz-Paszkowycz et al. (2013). The term Bpalm^ is used Registered in ZooBank under: urn:lsid:zoobank.org: for the proximal part of the cheliped propodus, i.e., without act:094408C2-6DD8-4161-9ACCEE0F743E505A the fixed finger. It is delimited by the line running through Diagnosis. Pleonite-5 without ventral spur. Pleotelson with- dactylus insertion and reaching the propodus ventral margin out lateral spurs, as long as broad or longer. Antenna with seven at a right angle. distinct articles. Cheliped with distinct carpal shield; carpus and Mar Biodiv (2018) 48:897–914 899 Fig. 1 Map with distribution of described species: yellow circle— Brixia aurora n. sp.; red circle— B. borealis n. sp.; green circles— Parakanthophoreus catharina n. sp. propodus without ornamentation. Pleopods rudimentary or 989-1, 61° 42.26′ N 19° 32.21′ W-61° 42.17′ N 19° 32.03′ W, absent. Uropod basis and endopod segment without spur. depth 1919.9–1925.6 m, EBS, 31 August 2011. Type species. Brixia aurora n. sp., by original designation. Etymology. From the Latin aurora, meaning sunrise. The Etymology. The genus is dedicated to Saskia Brix, the name refers to the Aurora Borealis that delighted and leader of the IceAGE project. entertained the IceAGE crew and participants during the col- Gender. Feminine. lection of the material. Species included. B. aurora n. sp., B. borealis n. sp. Diagnosis. Cephalothorax 1.7 times longer than wide. Remarks. Most of the characters in the amended diagnosis Pleotelson elongate, 1.4 times longer than wide, as long as last of Akanthophoreidae by Larsen and Araújo-Silva (2014)are three pleonites. Antennule article-1 3.7 times longer than wide, observable in Brixia n. gen. Nevertheless, seven distinct arti- longer than distal three articles combined length. Ischium of cles in the antenna, and rudimentary or absent pleopods may be pereopods 1–3 and 6 with one seta (pereopods 4–5 missing). unique characters for the family; the pleotelson is also longer Uropod basis four times longer than wide and as long as endopod than in most, if not all, Akanthophoreus, Chauliopleona, segment-1; exopod 0.6 times as long as endopod segment-1. Parakanthophoreus,and Paraleptognathia species, where it Description of female. Body (Fig. 2A, B) slender, 10.5 is shorter than broad. The antennal character derives from times longer than wide, 3.3 mm long. Cephalothorax two articles (4 and 5) with complete division, homologous to subrectangular, tapering anteriorly, 1.7 times longer than article-4 in those antennae exhibiting partial or indistinct artic- wide, withslighttriangular rostrum;carapacewithsingle seta ulation (Bfusion line^ sensu Larsen et al. 2006) within this at each anterolateral corner; eyelobes poorly marked. element as expressed in other akanthophoreids. The pleopod Pereonites parallel-sided or weakly convex; pereonite-1 rect- character (lack or rudimentary) is somewhat compromised by angular, shortest, 0.4 times as long as cephalothorax, 1.5 times the stated absence of these in Parakanthophoreus fastuosus longer than wide; pereonites 2, 4, and 5 subequal in length, 1.5 (Guerrero-Kommritz, 2004: 56), although this is not remarked times longer than pereonite-1, as long as wide, with parallel on or included in the generic diagnosis (op.cit.: 8–10). lateral margins; pereonite-6 just shorter, 1.4 times longer than Moreover, the results of molecular phylogeny (Błażewicz and pereonite-1, as wide as long. Pleon as long as combined length Jennings unpublished data) unequivocally support Brixia hav- of three last pereonites; pleonites subequal in length; each ing an affinity to, but separateness from, other akanthophoreids pleonite 2.4 times wider than long, without distinct lateral such as Parakanthophoreus or Chauliopleona. setae. Pleotelson elongate (relative to most akanthophoreids), Brixia aurora n. sp. 1.4 times longer than wide, as long as last three pleonites, Registered in ZooBank under: urn:lsid:zoobank.org: slightly tapering distally, bearing two setae mid-posteriorly. act:42A284B1-1D31-4375-A208-6F1D8E17057C Antennule (Fig. 2A, C) about 0.75 times length of cepha- (Figs. 2, 3) lothorax; article-1 3.7 times longer than wide, longer than Material examined: holotype non-ovigerous female distal three articles together, with two outer distal groups of (3.5 mm long) dissected on slides (ZMH K-52920), st. 853/ penicillate setae and one simple seta that reaches distal margin 900 Mar Biodiv (2018) 48:897–914 Table 1 Station data for non IceAge samples referred to in text Survey Sample Gear Date Locality Latitude (°N) Longitude Depth (m) B. Temp. (°C) Temp. (°C) Sediment AFEN 1996 53819#3 MC 25/07/1996 Faroe-Shetland Channel 61.59 − 1.832 1094 Fine sand AFEN 1996 53828#1 MC 05/08/1996 Faroe-Shetland Channel 61.4355 − 2.6175 1350 Very fine sand AFEN 1996 53833#1 MC 05/08/1996 Faroe-Shetland Channel 61.603 − 2.443 1513 Coarse silt AFEN 1998 54509#2 MC 25/05/1998 Faroe-Shetland Channel 62.473 0.382 901 Muddy sand AFEN 1998 54521#1/3 MC 25/05/1998 Faroe-Shetland Channel 61.732 − 1.618 1014 Gravelly muddy sand AFEN 1998 54522#1/2 MC 25/05/1998 Faroe-Shetland Channel 61.74 − 1.6625 1172 Muddy sand AFEN 1998 54524#2 MC 25/05/1998 Faroe-Shetland Channel 61.766 − 1.814 1393 Muddy sand BIOFAR 167 RP 08/05/1988 Norwegian Basin 62.781 − 3.52 1032 − 0.85 − 0.95 BIOFAR 168 RP 08/05/1988 Faroe-Norway Rise 62.693 − 3.62 899 − 0.66 − 0.95 Silt BIOFAR 169 RP 08/05/1988 Faroe-Norway Rise 62.621 − 3.54 808 − 0.6 − 0.7 BIOFAR 170 RP 08/05/1988 Faroe-Norway Rise 62.532 − 3.518 699 − 0.6 − 0.63 BIOFAR 172 RP 09/05/1988 Faroe-Norway Rise 62.318 − 3.914 507 1 1 Sand BIOFAR 478 DS 21/07/1989 Faroe-Shetland Channel 61.061 − 4.732 973 − 0.8 Soft BIOFAR 769 DS 06/10/1990 Faroe-Norway Rise 62.332 − 3.1 645 − 0.6 Silt, gravel, cobbles, stones BIOICE 2010 RP 20/07/1991 Iceland-Faroe Rise [North] 65.3047 − 11.2358 629 1 − 0.36 BIOICE 2011 RP 20/07/1991 Iceland-Faroe Rise [North] 65.3481 − 11.1711 761.5 1 − 0.42 BIOICE 2030 RP 23/07/1991 Iceland Plateau 67.0039 − 13.2628 830 0 − 0.55 BIOICE 2077 RP 03/07/1992 Iceland Plateau 67.675 − 17.173 1048 − 1 − 0.5 Silty sand BIOICE 2088 RP 04/07/1992 Iceland Plateau 67.239 − 17.857 617 − 1 − 0.4 Sand, silt mixed with foraminifera BIOICE 2124 RP 07/07/1992 Kolbeinsey Ridge 67.186 − 19.564 347 0 0.2 BIOICE 2136 RP 08/07/1992 Kolbeinsey Ridge 66.726 − 18.954 417 0 0.6 Soft BIOICE 2315 Sn 02/05/1993 Iceland-Faroe Rise [North] 64.1 − 9.05 991 2 Sandy silt BIOICE 2317 RP 02/05/1993 Iceland-Faroe Rise [North] 64.117 − 9.05 996 2 BIOICE 2318 Sn 02/05/1993 Iceland-Faroe Rise [North] 64.033 − 9.617 772 2 Sandy silt BIOICE 2323 RP 03/05/1993 Iceland-Faroe Rise [North] 63.917 − 10.083 623 2 BIOICE 2364 RP 08/05/1993 Iceland-Faroe Rise [North] 64.583 − 10.05 605 2 Sandy silt BIOICE 2638 RP 13/07/1994 Iceland Plateau 67.8113 − 15.503 1008 − 1 − 0.6 Silt BIOICE 2743 Sn 30/07/1995 Greenland-Iceland Rise 67.7492 − 20.4735 730 0 BIOICE 2749 Sn 31/07/1995 Greenland-Iceland Rise 68.0188 − 20.6547 956 0 − 0.6 Sandy gravelly silt BIOICE 2762 RP 01/08/1995 Iceland Plateau 67.9247 − 17.7043 1129 −1 − 0.5 BIOICE 2765 RP 10/08/1995 Iceland Plateau 68.0659 − 17.3233 1224.5 − 1 − 0.52 BIOICE 3023 RP 08/07/1997 Iceland-Faroe Rise [South] 63.4666 − 12.2153 427 3 4.66 BIOICE 3108 RP 21/08/1999 Greenland-Iceland Rise 67.1864 − 20.4537 326 1 − 0.21 BIOICE 3115 RP 22/08/1999 Kolbeinsey Ridge 67.4212 − 19.47925 604 −1 − 0.57 Mar Biodiv (2018) 48:897–914 901 Table 1 (continued) Survey Sample Gear Date Locality Latitude (°N) Longitude Depth (m) B. Temp. (°C) Temp. (°C) Sediment BIOICE 3121 DS 22/08/1999 Kolbeinsey Ridge 68.0423 − 18.0618 732 − 1 − 0.28 Silt BIOICE 3198 RP 08/07/2001 Iceland-Faroe Rise [North] 65.3005 − 8.3169 935 1 − 0.63 BIOICE 3225 RP 12/07/2001 Norwegian Basin 68.3461 − 8.1532 1994 0 − 0.86 DTI 2000 55282#1 MC 05/08/2000 Faroe-Shetland Channel 61.6135 − 1.9803 1305 DTI 2000 55283#1 MC 05/08/2000 Faroe-Shetland Channel 61.5901 − 1.9739 1262 DTI 2000 55287#1,2 MC 05/08/2000 Faroe-Shetland Channel 61.6535 − 1.8793 1290 DTI 2000 55299#1 MC 09/08/2000 Faroe-Shetland Channel 61.6028 − 1.924 1236 DTI 2000 55304#1 MC 09/08/2000 Faroe-Shetland Channel 61.6081 − 1.7733 1026 DTI 2000 55304#2 MC 09/08/2000 Faroe-Shetland Channel 61.6081 − 1.7733 1026 DTI 2000 55308#1 MC 10/08/2000 Faroe-Shetland Channel 61.648 − 1.8417 1234 DTI 2000 55310#2,3 MC 11/08/2000 Faroe-Shetland Channel 60.6632 − 4.4018 1095 DTI 2000 55322#1 MC 16/08/2000 Faroe-Shetland Channel 62.5917 0.1463 1068 DTI 2000 55324#1 MC 16/08/2000 Faroe-Shetland Channel 62.4385 0.1235 1046 DTI 2000 55326#1 MC 16/08/2000 Faroe-Shetland Channel 62.4415 0.0935 1144 DTI 2000 55328#1,2 MC 16/08/2000 Faroe-Shetland Channel 62.2401 0.1841 727 DTI 2000 55332#1,2 MC 17/08/2000 Faroe-Shetland Channel 62.207 0.381 640 DTI 2000 55336#1 MC 17/08/2000 Faroe-Shetland Channel 62.3015 0.1808 890 DTI 2000 55340#1 MC 18/08/2000 Faroe-Shetland Channel 62.4161 0.0711 1062 DTI 2000 55344#1 MC 18/08/2000 Faroe-Shetland Channel 62.4165 0.0711 1062 DTI 2000 55344#2 MC 18/08/2000 Faroe-Shetland Channel 62.4163 0.0711 1062 DTI 2000 55346#1 MC 19/08/2000 Faroe-Shetland Channel 62.2431 0.2683 849 DTI 2000 55347#1 MC 19/08/2000 Faroe-Shetland Channel 62.3091 0.2521 881 DTI 2000 55351#1 MC 19/08/2000 Faroe-Shetland Channel 62.3078 0.0973 939 DTI 2000 55352#1 MC 19/08/2000 Faroe-Shetland Channel 62.3847 0.0238 1061 DTI 2000 55354#1 MC 19/08/2000 Faroe-Shetland Channel 62.2915 0.1117 880 DTI 2000 55355#1 MC 19/08/2000 Faroe-Shetland Channel 62.2761 0.1267 829 DTI 2000 55372#2 MC 22/08/2000 North Shetland Slope 62.0458 0.2011 434 DTI 2000 55374#1 MC 22/08/2000 Faroe-Shetland Channel 62.308 0.1751 907 DTI 2000 55375#2 MC 22/08/2000 Faroe-Shetland Channel 62.325 0.1525 958 DTI 2000 55376#1 MC 22/08/2000 Faroe-Shetland Channel 62.3877 0.0985 1033 DTI 2000 55378#1 MC 23/08/2000 Faroe-Shetland Channel 62.3505 0.0927 1010 DTI 2000 55379#1 MC 23/08/2000 Faroe-Shetland Channel 62.3108 0.1323 934 DTI 2000 55386#1 MC 24/08/2000 Faroe-Shetland Channel 62.4005 0.586 791 DTI 2000 55387#1,2 MC 24/08/2000 Faroe-Shetland Channel 62.355 0.5367 781 DTI 2000 55395#1,2 MC 26/08/2000 Faroe-Shetland Channel 61.4198 − 3.116 1408 902 Mar Biodiv (2018) 48:897–914 of article-2; article-2 1.5 times longer than wide, 0.4 times as long as article-1, with outer pair of penicillate setae and single simple distal seta; article-3 1.3 times longer than wide, 0.7 times as long as article-2, with inner distal one simple seta and one penicillate seta; article-4 tapering, little longer than article-2, with five distal setae and one aesthetasc. Antenna (Fig. 2B, D) about 0.75 times as long as anten- nule; article-1 very short, naked; article-2 with two distal se- tae; article-3 as long as wide, with dorsodistal finely denticu- late seta; article-4 0.8 times as long as article-5 and as long as article-6, with ventrodistal penicillate seta; article-5 1.2 times longer than article-6, with two long ventrodistal, finely den- ticulate setae; article-6 more slender than article-5, with one distal seta; article-7 minute, with four distal setae. Labrum not recovered (but visible in whole view, Fig. 2B), conical. Left mandible (Fig. 2E) with six rounded teeth on pars incisiva; lacinia mobilis narrow, crenulate; pars molaris slender, distally bent downward, with rosette of numerous distal spinules. Right mandible (Fig. 2F) pars incisiva with three blunt teeth; pars molaris as in left mandible. Maxillule (Fig. 2G, G′) with nine terminal spines (three with fine den- ticulation, one plumose); palp with two plumose setae. Maxilla (Fig. 2H) larger than half of maxilliped plate, subtriangular. Labium (Fig. 2I) lobes narrow, distally setulose, outer lobe with microtrichia distally. Maxilliped (Fig. 2J) basis 1.8 times longer than wide, with single long seta reaching over distal margin of endite; endite distally with two small setae and two round tubercles; palp article-1 naked; article-2 with one outer distal finely denticulate and one simple and two inner distal finely denticulate setae; article-3 with two longer finely denticulate and one smaller simple inner distal setae; article-4 with subdistal simple seta and five distal finely denticulate setae. Epignath not recovered. Cheliped (Fig. 3A) attached via sclerite, with elongate basis 2.2 times longer than wide, posterior lobe as long as anterior mass, latter with one dorsal seta; merus subtriangular, with single ventral seta; carpus stout, 1.5 times longer than wide, with prominent carpal shield, well-developed rounded dorsoproximal extension, two mid-ventral setae and single dorsoproximal and dorsodistal setae; propodus large, just lon- ger but narrower than carpus, palm about as long as wide, dorsal and ventral margins smooth, with comb of three fine spines on inner surface near dactylus insertion; fixed finger with one seta near dactylus insertion, two setae ventrally and four teeth and three setae on cutting edge; dactylus with slight- ly undulate dorsal margin and two minute spines on cutting margin. Pereopod-1 (Fig. 3B) coxa with simple seta; basis 3.4 times longer than wide, narrow in most proximal part, naked; ischi- um with one ventral seta; merus one-third as long as basis, with one minute seta and robust ventrodistal finely denticulate spine that reaches beyond distal margin of carpus; carpus 1.2 times longer than merus, with numerous microtrichia and fine Table 1 (continued) Survey Sample Gear Date Locality Latitude (°N) Longitude Depth (m) B. Temp. (°C) Temp. (°C) Sediment SEAS4 57003#1&2 MC 01/07/2002 Faroe-Shetland Channel 62.2501 − 0.0663 803 88 μm mean grain size) SEAS4 57008#1–3 MC 01/07/2002 Faroe-Shetland Channel 62.0733 − 0.9671 897 116 μm SEAS4 57010#1&2 MC 01/07/2002 Faroe-Shetland Channel 62.2415 − 0.5503 893 108 μm SEAS4 57080#1 MC 01/07/2002 Faroe-Shetland Channel 62.1855 − 0.2955 700 181 μm SEAS4 57088#1 MC 01/07/2002 Faroe-Shetland Channel 62.1505 − 0.3171 650 227 μm Longitudes are °W, except positive values °E. B. Temp.-annual mean bottom temperature; Temp.–sampled bottom temperature Gear etc.: DS detritus sledge, MC megacorer, RP epibenthic sledge, Sn Sneli dredge Mar Biodiv (2018) 48:897–914 903 Fig. 2 Brixia aurora n. sp. female holotype (ZMH K-52920) A habitus dorsal view; B cephalothorax ventral view; C antennule; D antenna; E left mandible; F right mandible; G maxillule; G′ maxillule palp; H maxilla; I labium; J maxilliped. Scale bars 1 mm for A, 0.1 mm for C–J distal setulation, with single dorsodistal robust finely denticu- carpus, with microtrichia on ventral margin, with two late spine and two unequal (small and long) ventrodistal finely ventrodistal finely denticulate spines and two short dorsodistal denticulate spines; propodus 1.4 times longer than carpus, setae; dactylus denticulate on ventral margin, 1.4 times longer with microtrichia on ventral margin, with single ventrodistal than unguis, together 1.2 times longer than propodus. finely denticulate spine and dorsodistal fine seta; dactylus just Pleopods absent. shorter than unguis, together 0.9 times as long as propodus. Uropod (Fig. 3F) basis naked, as long as endopod segment- Pereopods 2 and 3 (Fig. 3C, D) similar to pereopod-1, but 1; exopod of two segments, 0.6 times as long as endopod pereopod-3 dactylus with accessory setae (possibly unob- segment-1; segment-2 with at least one short distal seta; served in pereopods 1–2). endopod of two segments; endopod segment-1 1.2 times as Pereopods 4 and 5 missing from dissected specimen. long as segment-2, that is armed with one seta distally Pereopod-6 (Fig. 3E) basis twice as long as wide, narrow (setation incomplete on specimen). distally, naked; ischium with one ventral seta; merus one third Remarks. Brixia aurora n. sp. differs from its congener (see as long as basis, with two ventrodistal finely denticulate spines below) by having a relatively long pleotelson—1.4 times lon- (one broken), that reach midway along carpus, and ventral ger than wide, a more slender antennule article-1, slight cren- microtrichia; carpus 1.7 times longer than merus, with ulation on the dorsal margin of the cheliped dactylus, only one microtrichia on ventral margin, and with three distal finely seta on the ischium of pereopod-6 (and possibly on pereopods denticulate spines (one broken) reaching one third along 4–5), no pleopods, and elongate uropod endopod segment-2, propodus and fine dorsodistal seta; propodus little shorter than that is 0.9 times as long as segment-1. 904 Mar Biodiv (2018) 48:897–914 Fig. 3 Brixia aurora n. sp. female holotype A cheliped; B pereopod- 1; C pereopod-2; D pereopod-3; E pereopod-6; F uropod. Scale bars 0.1 mm Distribution: Single record from the Iceland Basin, 1919.9– subrectangular, tapering towards anterior, 1.5 times longer 1925.6 m; see Fig. 1. than wide, with slight triangular rostrum, carapace with fine Brixia borealis n. sp. seta at each anterolateral corner, eyelobes poorly marked. Registered in ZooBank under: urn:lsid:zoobank.org: Pereonites sub-hexagonal, wider over pereopod insertions; act:0139C2E7-E1F7-4F7F-9FE7-8B556EE082F4 pereonite-1 tapering distally, shortest, 0.5 times as long as (Figs. 4, 5) cephalothorax, 1.3 times longer than wide; pereonites 2, 3 Material examined: holotype non-ovigerous female (3.3 mm and 4 subequal in length, 1.7 times longer than pereonite-1, long) dissected on slides (ZMH K-52921), st. 1149, 67° 50.57' 1.2 times longer than wide; pereonite-5 1.2 times longer than N 23° 41.85' W, depth 1246 m, AGT, 15 September 2011. pereonite-6; pereonite-6 just longer than pereonite-1, as wide Etymology. From the Latin borealis, northernorNorth. as long. Pleon as long as combined length of pereonites 5, 6 Diagnosis. Cephalothorax 1.5 times longer than wide. and two-thirds of pereonite-4; pleonites subequal in length, Pleotelson just longer than broad, shorter than last two pleonites. each pleonite 1.6 times wider than long, without lateral setae. Antennule article-1 2.7 times longer than wide, clearly shorter Pleotelson 1.1 times as long as wide, and shorter than last than distal three articles combined length. Pereopod-6 ischium two pleonites, slightly tapering distally, with pair of distal with two setae. Pleopods rudimentary. Uropod basis 1.5 times setae. longer than wide and 0.7 times as long as endopod segment-1; Antennule (Fig. 4A, C) about 0.9 times as long as cepha- exopod less than half as long as endopod segment-1. lothorax, article-1 2.7 times longer than wide, shorter than Description of female. Body (Fig. 4A, B) slender, 11.5 distal three articles together, with outer distal group of peni- times longer than wide, 3.3 mm long. Cephalothorax cillate setae and one simple distal seta shorter than article-2; Mar Biodiv (2018) 48:897–914 905 Fig. 4 Brixia borealis n. sp. female holotype (ZMH K-52921) A habitus dorsal view; B habitus lateral view; C antennule; D antenna; E labrum; F left mandible; G right mandible pars incisiva; H maxillule; H′ maxillule palp; I labium. Scale bars 1 mm for A, B, 0.1 mm for C–I article-2 2.3 times longer than wide, 0.7 times as long as denticulation, one plumose); palp (Fig. 4H′) with two plumose article-1, with outer single simple distal seta; article-3 0.7 setae. Maxilla (Fig. 5A) larger than half of maxilliped plate, times as long as wide, and 0.7 times as long as article-4, with subtriangular, distally setulose. Labium (Fig. 4I) lobes narrow, two small distal setae; article-4 tapering, 1.5 times longer than distally setulose, inner lobes with two distinct spines; outer article-3, with four distal setae and one aesthetasc. Blobe^ with microtrichia. Antenna (Fig. 4D) about 0.75 times as long as antennule; Maxilliped (Fig. 5B, B′) basis 1.8 times longer than wide, article-1 very short, naked; article-2 1.4 times longer than with single, long seta reaching over distal margin of endite; article-3, with one dorsodistal seta; article-3 as long as wide, endite distally with two rounded tubercles and one, fine seta; with dorsodistal seta; article-4 0.7 times as long as article-5 palp article-1 naked; article-2 with one outer distal and three and 0.8 times as long as article-6, with penicillate seta distally; inner distal setae (at least one of them finely denticulate); article-5 almost as long as article-6, with two simple and two article-3 with two short simple and two finely denticulate in- penicillate distal setae; article-6 more slender than article-5, ner setae; article-4 with subdistal seta and five finely denticu- with one distal seta; article-7 minute, with four distal setae. late distal setae. Epignath not recovered. Labrum (Fig. 4E) hood shaped, relatively elongate, finely Cheliped (Fig. 5C) attached via elongate sclerite reaching setose. Left mandible (Fig. 4F) pars incisiva with three teeth; to posterior of cephalothorax inserted to dorsum of basis; basis lacinia mobilis narrow, weakly crenulate dorsally; pars elongate, 2.3 times longer than wide, posterior lobe as long as molaris slender, distally bent downwards, with rosette of nu- anterior mass; merus subtriangular, with fine ventral seta; car- merous short and two long distal spinules. Right mandible pus stout, almost as long as wide, with well-developed round- (Fig. 4G) pars incisiva with several pointed teeth. Maxillule ed dorsoproximal extension and prominent carpal shield, with two fine mid-ventral setae and single dorsoproximal and (Fig. 4H) with nine terminal spines (three with fine 906 Mar Biodiv (2018) 48:897–914 Fig. 5 Brixia borealis n. sp. female holotype A maxilla; B maxillipedal palp and endite; B′ maxillipedal basis; C cheliped; D pereopod-1; E pereopod-2; F pereopod-4; G pereopod-6; H uropod. Scale bars 0.1 mm dorsodistal setae (one broken); propodus large, longer than ventrodistal finely denticulate robust spines; propodus 1.4 wide but shorter than carpus, palm about as long as wide, times longer than carpus, with microtrichia on ventral margin dorsal and ventral margins smooth, with comb of three fine and with single ventrodistal finely denticulate spine; dactylus spines on inner surface near dactylus insertion; fixed finger 0.8 times as long as unguis, ventrally denticulate, together 0.9 with one seta near dactylus insertion, two setae ventrally and times as long as propodus. three teeth and three setae on cutting edge; dactylus dorsal Pereopod-2 (Fig. 5E) similar to pereopod-1, but carpus margin smooth, with two minute spines on cutting margin. with additional ventrodistal spine. Pereopod-1 (Fig. 5D) coxa with simple seta (not shown on Pereopod-3 (not figured) as pereopod-2. figure); basis 2.6 times longer than wide, narrow in most distal Pereopod-4 (Fig. 5F) basis twice as long as wide, narrow in part, with long penicillate, dorsal seta; ischium with one ven- most distal part, apparently with two dorsal penicillate setae; tral seta; merus one-third as long as basis, with one robust ischium with two ventral setae; merus one third as long as basis, ventrodistal finely denticulate spine that reaches beyond distal with two ventrodistal finely denticulate spines, which reach margin of carpus; carpus 1.2 times longer than merus, with beyond midway on carpus; carpus 1.6 times longer than merus, numerous microtrichia, with single dorsodistal and with microtrichia on ventral margin, and with three distal finely Mar Biodiv (2018) 48:897–914 907 denticulate spines, the longest spine about 0.6 times as long as 53833#1; one specimen, AFEN 1998 54509#2; one specimen, propodus and dorsodistal seta; propodus as long as carpus, with AFEN 1998 54521#1,3; one specimen, AFEN 1998 54522#1,2; microtrichia on ventral margin, with two ventrodistal finely two specimens, AFEN 1998 54524#2; one specimen, BIOICE denticulate spines and one short dorsodistal seta; dactylus as 2010; 11 specimens, BIOICE 2011; four non-ovigerous fe- long as propodus, denticulate on ventral margin; unguis about males, BIOICE 2030; one non-ovigerous female, one ovigerous as long as half of dactylus, with ventral denticulation. female, BIOICE 2077; two non-ovigerous females, BIOICE Pereopod-5 (not figured) as pereopod-4. 2088; two manca-3, four non-ovigerous females, BIOICE Pereopod-6 (Fig. 5G) similar to pereopod-4, but propodus 2315; six non-ovigerous females, two ovigerous females, one with two dorsodistal setae. post-ovigerous female, BIOICE 2317; one manca-3, 12 non- Pleopods (Fig. 4B) biramous, rudimentary, without setae. ovigerous females, four ovigerous females, one preparatory Uropod (Fig. 5H) basis 0.7 times as long as endopod seg- male (uncertain?), BIOICE 2318; one ovigerous female, ment-1, naked; exopod 0.4 times as long as endopod segment- BIOICE 2323; one non-ovigerous female, one ovigerous fe- 1; segment-1 with distal seta; segment-2 with one long and male, two post-ovigerous females, BIOICE 2364; one non- one short distal setae; endopod segment-1, five times as long ovigerous female, one ovigerous female, BIOICE 2638; one as wide, twice as long as segment-2, with one penicillate seta preparatory male (?), BIOICE 2743; one non-ovigerous female, distally; segment-2 with one subdistal and two distal setae. BIOICE 2749; three non-ovigerous females, one non-ovigerous Remarks. In contrast to B. aurora,in B. borealis n. sp., the female, BIOICE 2762; one specimen, BIOICE 2765; one pleotelson is shorter, (but still as long as or just longer than broad), the cheliped dactylus is apparently smooth, the ischi- um of pereopods 4–6 has two setae, and the uropod endopod segment-2 is about half as long as segment-1. Distribution: Single record from the Denmark Strait, 1246 m; see Fig. 1. Parakanthophoreus Larsen & Araújo-Silva, 2014 Parakanthophoreus catharina n. sp. Registered in ZooBank under: urn:lsid:zoobank.org: act:62B32816-9167-4CB9-82A9-C69AD931AE52 (Figs. 6–10) Akanthophoreus inermis (Hansen, 1913): Bird (2001): 32–33. Material examined: holotype non-ovigerous female (2.8 mm long) (NMS.Z.2018.26.1), DTI2000 st. 55283#1, 61° 35.06′ N 1° 58.43′ W, depth 1262 m, Megacore, 5 August 2000. Allotype preparatory male (2.5 mm long) (NMS.Z.2018.26.2), partly dissected on microslide (NMS.Z.2018.26.3), DTI2000 st. 55374#1, 62° 18.48′ N0° 10.51′ E, depth 907 m, Megacore, 22 August 2000. Paratype female dissected on slides (ZMH K-52922), st. 853/ 1132-1, 67° 38.39′ N 26° 45.46' W-67° 38.28′ N 26° 45.72′ W, depth 316.6–316.8 m, EBS, 14 September 2011. Other material taken during IceAGE project: one non-ovigerous female (2.4 mm long), one manca (1.3 mm long), st. 853/1116-1, 67° 12.82′N26° 16.31′ W, depth 683.1 m, GKG, 14 September 2011; one non- ovigerous female (1.6 mm long), one manca (1.3 mm), st. 853/ 1129-1, 67° 38.77′ N 26° 44.78′ W, depth 320.6 m, GKG, 14 September 2011; four non-ovigerous females (1.4, 1.6, 2.3, and 2.3 mm long), one manca (1.2 mm long), st. 853/1141-1, 67° 50.22′ N 23° 42.11′ W, depth 1241.6 m, GKG, 15 September 2011; one female with marsupium, one non-ovigerous female (1.7 mm long), one manca (0.9 mm long), st. 873-4, 61° 46.52′ N 03° 52.51′ W, depth 833.8 m, Van Veen grab, 28 July 2013. Fig. 6 Parakanthophoreus catharina n. sp. (NMS.Z.2018.26.1) habitus Supplementary material and records examined—see Table 1 A female holotype dorsal view; B male allotype (NMS.Z.2018.26.2) for station data: one specimen, AFEN 1996 53819#3; one spec- dorsal view; C male antennule (NMS.Z.2018.26.3). Scale bars 1 mm imen, AFEN 1996 53828#1; one specimen, AFEN 1996 for A, B and 0.1 for C 908 Mar Biodiv (2018) 48:897–914 specimen, BIOICE 3023; one non-ovigerous female, BIOICE 55352#1; one non-ovigerous female, DTI2000 55354#1; two 3108; four specimens, two non-ovigerous females, two prepara- non-ovigerous females, DTI2000 55355#1; one non-ovigerous tory males (?), BIOICE 3115; two specimens, BIOICE 3121; female, DTI2000 55374#1; two non-ovigerous females, one specimen, BIOICE 3198; one specimen, BIOICE 3225; DTI2000 55375#3; one non-ovigerous female, DTI2000 three non-ovigerous females, four specimens, BIOFAR 167; 55376#1; one non-ovigerous female, DTI2000 55378#1; one six specimens, BIOFAR 168; one specimen, BIOFAR 169; non-ovigerous female, DTI2000 55379#1; one non-ovigerous two specimens, BIOFAR 170; seven specimens, BIOFAR female, DTI2000 55395#1; two non-ovigerous females, SEAS4 172; seven specimens, BIOFAR 478; six specimens, BIOFAR 57003#1&2; two non-ovigerous females, two post-ovigerous 769; four non-ovigerous females, DTI2000 55283#1; three non- females, SEAS4 57008#1-3; one specimen, SEAS4 57080#1; ovigerous females, DTI2000 55322#1; three non-ovigerous fe- one non-ovigerous female, SEAS4 57088#1. males, DTI2000 55324#; one non-ovigerous female, DTI2000 Etymology. Catharina (Lat.) (= Katarzyna in Polish): the 55336#1; three non-ovigerous females, DTI2000 55344#1; one species is dedicated to the senior author’s wife, Katarzyna preparatory male (?), DTI2000 55344#2; one non-ovigerous Pawicka-Jóźwiak. female, DTI 2000 55347#1; one preparatory male (?), Diagnosis. Body about ten times longer than wide. DTI2000 55351#1; two non-ovigerous females, DTI2000 Pereonite-1 as long as wide. Maxilliped basis with setae. Fig. 7 Parakanthophoreus catharina n. sp. female (ZMH K-52922) A antennule; B antenna; C labrum; D left mandible; E right mandible; F maxillule; G maxilla; H labium; I maxilliped; J epignath. A–D, F–H and J female paratype; E and I preparatory male allotype (NMS.Z.2018.26.3), DTI2000 Stn 55374#1. Scale bars 0.1 mm for B–D, F, H and J and 0.25 for A, E and I Mar Biodiv (2018) 48:897–914 909 Maxilliped endite with pair of distal setae. Cheliped carpus, than pleonites 4–5, with rounded apex. Cephalothorax and propodus and dactylus without crenulation but carpus with tergites of pereonites and pleonites with dense pitting. deep shield; cheliped carpus with single ventral seta; fixed Antennule (Figs 6A, 7A) about 0.75 times as long as ceph- finger ventral setae unequal in thickness. Pereopods 1–3car- alothorax; article-1, twice as long as wide, with one simple pus with dorsodistal seta. Pereopods 4–6 ischium with pair of and two penicillate setae on outer distal margin; article-2 setae and dactylus with denticulate ventral margins. Pereopod- about half as long as article-1, with one short inner distal seta 6 propodus with three dorsodistal finely denticulate spines. and one long outer distal seta; article-3 half as long as article- Uropod exopod 0.7 times as long as endopod segment-1. 2, with one short inner distal seta, one longer outer distal seta Description of non-ovigerous female. Body (Fig. 6A) slen- and some distal spinules; article-4 1.5 times longer than arti- der, 10.1 times longer than wide, 2.8 mm long (range 1.24– cle-3, with two simple setae distally; article-5 minute, with 2.98 mm, n = 70 G. Bird material). Cephalothorax 15% of total four simple setae distally. Articles with dense pitting. body length, about 1.5 times longer than wide, carapace with Antenna (Figs 7B, 9A) article-1 very short, naked; article-2 fine seta at each anterolateral corner, eyelobes poorly marked. just longer than wide, with two dorsodistal and distolateral Pereonites parallel-sided, length/width ratio: 1, 1.1, 1.4, 1.4, setae; article-3 0.7 times as long as wide, with one simple 1.1, and 0.9. Pleon 18% of total body length, pleonites distal seta; article-4 3.8 times longer than wide, with fusion subequal in length, epimera with single seta. Pleotelson longer line, one penicillate seta near fusion line, one penicillate seta Fig. 8 Parakanthophoreus catharina n. sp. female paratype (ZMH K-52922) A cheliped; B pereopod-1; C pereopod-2; D pereopod-3; E pereopod-4; F pereopod-5; G pereopod-6; H pleopod basis and endopod; H' pleopod exopod; setae all plu- mose but not shown for clarity; I uropod. Scale bars 0.1 mm 910 Mar Biodiv (2018) 48:897–914 Fig. 9 Parakanthophoreus catharina n. sp. cuticle ornamentation using SE microscope A antenna article-4; B cheliped basis and sclerite; C pereopod-1; D uropod. Scale bars 0.05 mm for A and C, 0.02 for B and D subdistally (partly broken), one minute and four simple setae Cheliped (Figs 8A, 9B, 10A–D) attached via large distally; article-5 just longer than half of article-4, with two posterodorsal sclerite; posterior lobe and anterior mass subequal simple setae distally; article-6 minute, with five simple setae in size, naked; merus with one simple ventral seta; carpus with distally. Articles with dense pitting. well-developed rounded dorsoproximal extension and deep Labrum (Fig. 7C) trapezoidal in lateral view, distally setose. shield, one simple midlength seta ventrally and one simple seta Left mandible (Fig. 7D) pars incisiva with five blunt denticles; dorsodistally; propodus about half as long as wide, palm about as lacinia mobilis weakly crenulate dorsally; pars molaris long as wide, dorsal and ventral margins smooth, with simple narrowing distally, with rosette of numerous distal spinules. seta near dactylus insertion, and one longer and three short finely Right mandible (Fig. 7E) pars incisiva with three denticles dis- denticulate spines on inner margin, inner and outer margins with tally; pars molaris as in left mandible. Maxillule (Fig. 7F) endite microtrichia, outer margin additionally with protrusions; fixed with some inner microtrichia, subdistally with setae, with eight finger with two denticles and three simple setae on cutting edge, terminal spines (at least three finely denticulate); palp not ob- and two setae unequal in thickness ventrally; dactylus as long as served. Maxilla (Fig. 7G) triangular, inner and distal margin fixed finger, naked. Articles with dense pitting (Figs 9B, 10). setose. Labium (Fig. 7H) simple, slender, distally with spinules. Pereopod-1 (Figs 8B, 9C) coxa with simple seta (not Maxilliped (Fig. 7I) basis with one distal seta, half as long shown on figure); basis 1.9 times longer than wide, naked; as endite; endite with distal tubercle, pair of distal setae and ischium with one seta; merus about as long as carpus, with outer distal setules; palp article-1 naked; article-2 with one ventral microtrichia, distal seta, and ventrodistal finely dentic- simple outer seta and one plumose and two simple setae on ulate spine; carpus with one complex-tip seta and finely den- inner distal margin; article-3 with two simple short, and two ticulate spine dorsodistally, one finely denticulate spine long setae on inner margin; article-4 with one subdistal seta ventrodistally and numerous microtrichia along article; and five distal setae. Epignath (Fig. 7J) slender, tapering dis- propodus 1.4 times longer than carpus, with ventrodistal spine tally to the pointed tip. and ventral microtrichia; dactylus and unguis combined 0.6 Mar Biodiv (2018) 48:897–914 911 Fig. 10 Parakanthophoreus catharina n. sp. cheliped ornamentation of using SE microscope A outer surface; B propodus outer surface; C chela inner surface; D carpus outer surface. Scale bars, respectively, 0.1, 0.02, 0.05, and 0.01 mm times as long as propodus; unguis 1.7 times longer than Pleopod (Fig. 8H, H') basis shorter than ramus, with dactylus. Articles of all pereopods with dense pitting. ventrodistal seta; endopod with 12 setae along ventral and distal Pereopod-2 (Fig. 8C) similar to pereopod-1, but basis with margins; exopod with eight distal setae. All setae plumose. long penicillate seta dorsally and carpus with additional Uropod (Figs 8I, 9D) basis 0.8 times as long as exopod, ventrodistal finely denticulate spine. naked; exopod of two segments, 0.7 times as long as endopod Pereopod-3 (Fig. 8D) similar to pereopod-2, but basis without segment-1, segments subequal; segment-1 with short seta; penicillate seta (possibly lost) and dactylus with proximal seta. segment-2 with one short and one long setae; endopod of Pereopod-4 (Fig. 8E) basis 2.4 times longer than wide, two equal segments; segment-1 with one simple and three narrower than those of pereopods 1–3, with one penicillate penicillate setae distally; segment-2 with one long midlength midlength seta ventrally; ischium with two setae; merus one seta and one penicillate and four simple setae distally. third of basis length, with two finely denticulate spines and Manca-3. Length 1.03–1.05 mm (n =3). microtrichia ventrodistally; carpus 1.3 times longer than merus, Ovigerous female. As female (described above) but more with one complex-tip seta and one finely denticulate spine dorsoventrally compressed, with oostegites; length 1.93– dorsodistally, two finely denticulate spines ventrodistally and 2.51 mm (n =10). some ventral microtrichia; propodus just longer than carpus, Post-ovigerous female. As female (described above) but with one finely denticulate spine dorsodistally, two finely den- dorsoventrally compressed, lacking oostegites, but with scars ticulate spines ventrodistally and ventral microtrichia; dactylus or remnants thereof; length 2.15–2.35 mm (n =5). and unguis combined 1.3 times longer than propodus, both Preparatory male. As female (described above) but with denticulate ventrally; unguis about half as long as dactylus. proportionately larger antennules, with stouter articles 1–2; Pereopod-5 (Fig. 8F) as pereopod-4. length 1.79–3.00 mm (n = 6). Body (Fig. 6B) slender, 10.3 Pereopod-6 (Fig. 8G) as pereopod-4, but propodus with times longer than wide, 2.8 mm long. Cephalothorax 14% of three dorsodistal finely denticulate setae. total body length, about 1.5 times longer than wide, carapace 912 Mar Biodiv (2018) 48:897–914 Fig. 11 Parakanthophoreus inermis near-topotypical BIOICE Stn 2629 A female habitus; B right cheliped; C pereopod-1; D pereopod-6. Scale bars 1 mm for A, 0.25 mm for B–D with fine seta at each anterolateral corner, eyelobes poorly (about ten times longer than wide), maxilliped basis with seta, marked. Pereonites parallel-sided, length/width ratio: 0.8, maxilliped endites with a pair of distal setae, cheliped carpus, 1.1, 1.1, 1.0, 0.9, and 0.6. Pleon 18% of total body length, propodus, and dactylus without crenulation, but carpus with deep pleonites subequal in length, epimera with single seta. shield, pereopods 4–6 ischium with pair of setae and dactylus Pleotelson as long as pleonites 3–5 and half of pleonite-2, with with denticulate ventral margins, uropod exopod distinctly longer rounded apex, terminated with two pairs of setae. than half of endopod segment-1. Another useful characteristic Antennule (Fig. 6B, C) about 0.75 times as long as cepha- distinguishing P. catharina from its congeners, but also from lothorax; article-1, twice as long as wide, with one simple and other akanthophoreids, is the setation of the cheliped fixed finger, five penicillate setae on outer distal margin; article-2 0.6 times where the two ventral setae are of unequal thickness. as long as article-1, with one long and three penicillate outer Of the described species of Parakanthophoreus known from distal setae; article-3 0.7 times as long as article-2, with one the general area of the IceAGE project—P. alba (Hansen, 1913), short and one longer inner distal setae; article-4 just shorter than P. brachiatus (Hansen, 1913), P. inermis (Hansen, 1913), article-2, with four simple setae distally and one aesthetasc. P. longiremis (Lilljeborg, 1864), P. multiserratus (Hansen, 1913), Remarks. Parakanthophoreus catharina n. sp. differs from and P. vikingra (Błażewicz-Paszkowycz & Bamber, 2011)—the other members of the genus by a combination of: elongated body most likely species to be confused with P. catharina is P. inermis, Mar Biodiv (2018) 48:897–914 913 Ethical approval All applicable international, national, and/or institution- since it can be sympatric, and most of the non-IceAGE records of al guidelines for the care and use of animals were followed by the authors. P. catharina were originally assigned, provisionally, to that spe- cies. A review and comparison of these species (except Field study Permits and approval of field or observational studies are P. vikingra) was given by Guerrero-Kommritz (2004), while not applicable for authors. Hansen (1913) is still a good reference for species comparisons. Open Access This article is distributed under the terms of the Creative Parakanthophoreus inermis was characterized by Hansen Commons Attribution 4.0 International License (http:// (1913) by having a relatively short antennule article-1 compared creativecommons.org/licenses/by/4.0/), which permits unrestricted use, to other similar species, as well as a deep cheliped carpal shield distribution, and reproduction in any medium, provided you give appro- priate credit to the original author(s) and the source, provide a link to the and a short claw on pereopods 1–3, all of which are shared by Creative Commons license, and indicate if changes were made. P. catharina but not by the other Parakanthophoreus species mentioned above. Redescriptions of P. inermis by Guerrero- Kommritz (2004)and Sieg (1986b) were unhelpful in clarifying References the nature of this species because of several factors such as un- clear attribution of material, poor illustrations, or use of non- Bird GJ (2001) Tanaidacea of the Atlantic Margin: the AFEN surveys of topotypical specimens. Indeed, the taxon dealt with by Sieg 1996 and 1998. Unpublished report to the Atlantic Frontier may be conspecific with, or at least clearly related to, Environmental Network and UK Offshore Oil Association, P. catharina rather than P. inermis and it is possible that some February 2001 Bird GJ (2007) Family incertae cedis [sic]. In: Larsen K, Shimomura M of Hansen’s material may too be of both species. A revision of (eds) Tanaidacea (Crustacea: Peracarida) from Japan III. The deep P. inermis is in progress (Bird, in prep.). trenches: the Kurile-Kamchatka Trench and Japan Trench. Magnolia Examination of near-topotypical specimens of Press, Auckland, pp 121–149 Parakanthophoreus inermis (Fig. 11) indicates that it can be sep- Bird GJ (2015) Tanaidacea (Crustacea: Peracarida) of the Northeast arated from P. catharina by, inter alia, its shorter cephalothorax, Atlantic: Chauliopleona Dojiri and Sieg, 1997 and Saurotipleona n. gen. from the ‘Atlantic Margin’.JNatHist49:1507–1547. https:// shorter pereonites (especially pereonite-1, Fig. 11A)–none longer doi.org/10.1080/00222933.2015.1005715 than wide, weaker (but still relatively deep) cheliped carpal shield Błażewicz-Paszkowycz M, Bamber R (2011) Tanaidomorph Tanaidacea (Fig. 11B), lack of a distodorsal seta on the carpus of pereopods 1– (Crustacea: Peracarida) from mud-volcano and seep sites on the 3(Fig. 11C; very conspicuous in P. catharina), and only two Norwegian margin. Zootaxa 3061:1–35 Błażewicz-Paszkowycz M, Bamber RN, Cunha MR (2011) New dorsodistal spines on the propodus of pereopod-6 (Fig. 11D; three tanaidomorph Tanaidacea (Crustacea: Peracarida) from submarine in P. catharina); the cuticular ornamentation is also less pro- mud-volcanoes in the Gulf of Cadiz (North-East Atlantic). Zootaxa nounced, without the pitting seen in P. catharina.Other 2769:1–53 akanthophoreids in the Iceland-Faroe region with large cheliped Błażewicz-Paszkowycz M, Bamber RN, Jóźwiak P (2013) Tanaidaceans (Crustacea: Peracarida) from the SoJaBio joint expedition in slope carpal shields include Chauliopleona hastata (Hansen, 1913) and and deeper waters in the Sea of Japan. Deep-Sea Res II 111:325– Saurotipleona julii Bird, 2015 but these can be distinguished, at 332. https://doi.org/10.1016/j.dsr2.2014.08.021 least, by their ventral spur on pleonite-5. Brix S, Bauernfeind W, Brenke N, Blazewicz M, Borges V, Buldt K, Distribution: BIOICE records; 23, from the Greenland- Cannon J, Díaz Agras G, Fiege D, Fiorentino D, Haraldsdóttir S, Iceland Rise, Iceland Plateau, Iceland-Faroes Rise (all but Hoffmann S, Holst S, Hüttmann F, Jeskulke K, Jennings R, Kocot K, Khodami S, Lucas Rodriguez Y, Martinez Arbizu P, Meißner K, one from northern flank), Kolbeinsey Ridge, and Norwegian Mikkelsen N, Miller M, Murray A, Neumann H, Ostmann A, Riehl Basin, 326–1994 m, in temperatures < 1 °C. Other records, 53 T, Schnurr S, Savavarsson J, Yasuhara M (2011) Cruise Report records from the Faroe-Shetland Channel, Faroe-Norway M85/3 IceAGE1 2011 with RV Meteor. Reykjavik, Cuxhaven Rise, North Shetland Slope, and Norwegian Basin, 434– Brix S,MeißnerK,StranskyB,Halanych KM, Jennings RM, Kocot KM, Svavarsson J (2014) The IceAGE project a follow up of BIOICE. Pol 1513 m (Table 1; Bird 2001 and ined.) (see Fig. 1). Polar Res 35:141–150. https://doi.org/10.2478/popore-2014-0010 Coleman CO (2003) Digital inking: how to make perfect line drawings on Acknowledgements Authors would like to thank Saskia Brix for making computers. Org Divers Evol 14:1–14 the material available for study and the crew of R/V Meteor and R/V Dojiri M, Sieg J (1997) The Tanaidacea. In: Blake JA, Scott PH (eds) Poseidon as well as the scientists who participated in IceAGE Program. Taxonomic atlas of the benthic fauna of the Santa Maria Basin and We are also grateful for two anonymous reviewers for their valuable Western Santa Barbara Channel. Volume 11. The Crustacea Part 2. comments. Studies were financed by Polish National Science Centre The Isopoda, Cumacea and Tanaidacea. Santa Barbara Museum of grant UMO-2014/13/B/NZ8/04702. Natural History, Santa Barbara, CA, pp 181–278 Guerrero-Kommritz J (2004) A revision of the genus Paraleptognathia Funding This study was funded by the Polish National Science Centre Kudinova-Pasternak, 1981 (Crustacea: Tanaidacea) and description grant (UMO-2014/13/B/NZ8/04702). of four new species. Zootaxa 481:1–63 Hansen HJ (1913) Crustacea, Malacostraca. II. IV. The order Tanaidacea. Compliance with ethical standards Danish Ingolf Expedit 3:1–145 Kudinova-Pasternak RK (1981) Tanaidacea. In: Kusnecov AP, Mironov AN Conflict of interest The authorsdeclare thattheyhave noconflictofinterest. (eds) Bentos podvodnych gor Markus-Nekker i smeznych rajonov 914 Mar Biodiv (2018) 48:897–914 tichogo okeana [Benthos of the Submarine Mountains Markus Necker Larsen K, Wilson GDF (2002) Tanaidacean phylogeny, the first step: the superfamily Paratanaidoidea. J Zool Syst Evol Res 40:205–222 and adjacent Pacific regions]. Institut Okeanologii, Moscow, pp 94–112 Kudinova-Pasternak RK (1985) Tanaidacea (Crustacea, Malacostraca) Sieg J (1986a) Tanaidacea (Crustacea) von der Antarktis und collected on the summit and at foot of Great-Meteor Seamount. T Subantarktis. II. Tanaidacea gesammelt von Dr. JW. Wägele I Ocean 120:52–64 wahrend der Deutschen Antarktis Expedition 1983. Zoologischen Larsen K (2005) Deep-sea Tanaidacea (Peracarida) from the Gulf of Museum der Universität Kiel 2:1–80 Mexico. Brill, Leiden Sieg J (1986b) Crustacea Tanaidacea of the Antarctic and the Larsen K, Araújo-Silva CL (2014) The ANDEEP Tanaidacea (Crustacea: Subantarctic. 1. On material collected at Tierra del Fuego, Isla de Peracarida) revisited III: the family Akanthophoreidae. Zootaxa los Estados, and the west coast of the Antarctic Peninsula. In: 3796:237–264. https://doi.org/10.11646/zootaxa.3796.2.2 Korniker LS (ed.) Biology of the Antarctic Seas 18. Volume 45 in Larsen K, Błażewicz-Paszkowycz M, Cunha MR (2006) Tanaidacean the Antarctic Research Series. American Geophysical Union, (Crustacea: Peracarida) fauna from chemically reduced habitats— Washington, D.C., pp 1–180 the lucky strike hydrothermal vent system, mid-Atlantic [sic] ridge. Zootaxa 1187:1–36

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