Breviconia Conroy-Dalton & Huys, 2000

Breviconia Conroy-Dalton & Huys, 2000 View in CoL

Breviconia australis ( George, 1998) View in CoL c

Breviconia echinata ( Brady, 1918) d

Laophontodes T. Scott, 1894 View in CoL

Laophontodes hamatus ( Thomson, 1883) View in CoL g Laophontodes typicus (T. Scott, 1894) View in CoL h Laophontodes bicornis A. Scott, 1896 View in CoL

Laophontodes propinquus Brady, 1910 i Laophontodes whitsoni T. Scott, 1912 View in CoL

Laophontodes antarcticus Brady, 1918 View in CoL j Laophontodes brevis Nicholls, 1944 k

Laophontodes ornatus Krishnaswamy, 1957 View in CoL Laophontodes macclintocki Schizas & Shirley, 1994 View in CoL Laophontodes spongiosus Schizas & Shirley, 1994 View in CoL Laophontodes mourois Arroyo, George, Benito & Maldonado, 2003 View in CoL

Laophontodes horstgeorgei George & Gheerardyn, 2015 View in CoL Laophontodes sabinegeorgeae George & Gheerardyn, 2015 View in CoL

Laophontodes gertraudae George, 2018 View in CoL Laophontodes monsmaris George, 2018 View in CoL Laophontodes sarsi George, 2018 View in CoL

Laophontodes scottorum George, 2018 Laophontodes georgei sp. nov. l

Paralaophontodes Lang, 1965 View in CoL

Paralaophontodes echinatus ( Willey, 1930) View in CoL m Paralaophontodes armatus View in CoL ( Lang, 1936c) n Paralophontodes robustus ( BoŽić, 1964) o Paralaophontodes hedgpethi ( Lang, 1965) View in CoL n Paralaophontodes psammophilus ( Soyer, 1975) View in CoL n Paralaophontodes elegans Baldari & Cottarelli, 1986 View in CoL Paralaophontodes exopoditus Mielke, 1981 View in CoL Paralaophontodes anjae George, 2017 View in CoL

Tapholaophontodes Soyer, 1975 View in CoL

Tapholaophontodes rollandi Soyer, 1975 View in CoL Tapholaophontodes remotus Cottarelli & Baldari, 1987 View in CoL

Algensiella Cottarelli & Baldari, 1987 View in CoL

Algensiella boitanii Cottarelli & Baldari, 1987 View in CoL Algensiella laurenceae ( Bodiou & Colomines, 1988) View in CoL p

Probosciphontodes Fiers, 1988 View in CoL

Probosciphontodes ptenopostica Fiers, 1988 View in CoL Probosciphontodes stellata Fiers, 1988 View in CoL

Juxtaramia Conroy-Dalton & Huys, 2000 View in CoL Lobopleura Conroy-Dalton, 2004 View in CoL

Juxtaramia polaris Conroy-Dalton & Huys, 2000 View in CoL Lobopleura ambiducti Conroy-Dalton, 2004 View in CoL Lobopleura expansa ( Sars, 1908) View in CoL q

Uptionyx Conroy-Dalton & Huys, 2000 View in CoL Lobopleura multispinata View in CoL (Kornev & Chertoprud,

Uptionyx verenae Conroy-Dalton & Huys, 2000 View in CoL 2008) comb. nov.

a Based on copepodid V stage and relegated to species inquirenda by Gómez & Conroy-Dalton (2002).

b Transferred from Echinopsyllus Sars, 1909 by George (1998).

c Transferred from Arthropsyllus Sars, 1909 by Conroy-Dalton & Huys (2000).

d Transferred from Laophontodes T. Scott, 1894 and placed species inquirenda in Breviconia Conroy-Dalton & Huys, 2000 by Conroy-Dalton & Huys (2000). e Transferred from Polyascophorus George, 1998 by Conroy-Dalton (2001).

f Transferred from Ceratonotus Sars, 1909 by Conroy-Dalton (2003a).

g Transferred from Merope Thomson, 1883 by Gurney (1927).

h T. Scott (1894: 249) designated the new species ‘ Laophontodes typicus ’ as the type (by monotypy) for the new subgenus Laophontodes in the genus Laophonte Philippi, 1840 . However, in the description of the accompanying plate (p. 270) he cited the species as ‘ Laophontodes typicus , gen. et sp. nov. ’ attributing instead generic rank to Laophontodes . According to ICZN Recommendation 69A.10, when designating the type species for a genus (or subgenus), all other things being equal (i.e. a choice cannot be made on the basis of Recommendations 69A.1–9), an author should give preference to the species (or spelling) cited first in the work, page or line. Huys (2009) followed this position precedence and pointed out that the author and date of the type species of the genus Laophontodes should therefore be enclosed in parentheses, reflecting the changed combination currently adopted. The first author to use Laophontodes explicitly as a generic name was A. Scott (1896: 148).

i Brady’s (1910) description is grossly incomplete and probably deficient. No information is available on P2–P3 except that the endopods are one-segmented. Given that the species cannot be identified unequivocally on the basis of the original description, it is here relegated to species inquirenda in the genus. It is known from very few specimens from Observatory Bay in the Kerguelen Islands and has never been found again since its discovery. j Relegated to species incertae sedis by Lang (1936b); according to Gheerardyn & Lee (2012), it should probably be retained in Laophontodes (as shown by the elongate caudal rami and certain characteristics of antennule, P1 and P5), but no firm conclusion can be made because of the lack of detail in the original description.

k Lang (1965) claimed that Laophontodes brevis agreed well with the last copepodid of La. bicornis found in the Gullmarfjord ( Sweden) and that both species were probably conspecific; he did not, however, formally synonymize them, although some authors subsequently listed them as such ( Bodin, 1997) or no longer regarded La. brevis as a valid species ( Gee & Fleeger, 1986; Schizas & Shirley, 1994; Wells, 2007; George & Gheerardyn, 2015; George, 2018). The description of the latter is indeed based on a copepodid V, but given the confusion surrounding the identity of Nicholls’s (1944) material of the co-occurring La. bicornis (cf. George & Gheerardyn, 2015) it seems prudent to maintain La. brevis as a species inquirenda in the genus. l George (2018: 34) proposed the new name Laophontodes norvegicus for the species originally redescribed by Sars (1908: 270) under the name La. typicus (T. Scott, 1894) . In the absence of extant material, the author refrained from explicitly fixing a holotype, but his course of action effectively makes the new name unavailable (ICZN Art. 16.4). The ovigerous female specimen illustrated by Sars (1908: pl. CLXXXVII) is here designated as the holotype of Laophontodes georgei (ICZN Art. 72.5.6). The new species can be differentiated by the characters mentioned and illustrated by Sars (1908: 270–271, pl. CXXXVII) (ICZN Art. 13.1) and summarized by George (2018: 37–38). It is named after Dr Kai Horst George in recognition of his contributions to the systematics and phylogeny of ancorabolid harpacticoids. LSID: urn:lsid:zoobank.org:act:011872B0-D001-40D8-95B5-94B86A11E3E7 .

m Transferred from Laophonte Philippi, 1840 by Lang (1965). BoŽić (1964) proposed the new replacement name Laophontodes willeyi for Laophontodes echinatus ( Willey, 1930) , a junior secondary homonym at the time of La. echinatus Brady, 1918 . Given that both species are no longer considered congeneric (having been transferred to Paralaophontodes Lang, 1965 and Breviconia , respectively) a substitute name is not necessary, and the junior secondary synonym is to be reinstated (ICZN Art. 59.4).

n Transferred from Laophontodes by George (2017). Soyer (1976: 146) listed ‘ Laophontodes kerguelenensis ’ from the Rivière norvégienne, which is the type locality of Laophontodes psammophilus Soyer, 1976 . It is conceivable that he referred to the latter species, because it was mentioned as one of the two ancorabolid representatives obtained in the Kerguelen samples ( Soyer, 1976: 149).

o Transferred from Laophontodes by Lang (1965). Synonymized with P.echinatus by Wells & Rao (1987) but reinstated as a valid species by George (2017). p Transferred from Tapholaophontodes Soyer, 1975 by Wells (2007).

q Transferred from Laophontodes by Conroy-Dalton (2004).

r Transferred from Laophontodes and placed species inquirenda in Calypsophontodes Gheerardyn & Lee, 2012 by Gheerardyn & Lee (2012).

s Transferred from Laophontodes by Gheerardyn & Lee (2012).

discovered in previously unexplored habitats, such as hydrothermal vents ( Conroy-Dalton & Huys, 2000), seamounts ( George, 2006b, 2018; George & Plum, 2009; Gheerardyn & George, 2010; Schulz & George, 2010) and submarine caves ( George & Tiltack, 2009), or recognized on the basis of revisionary systematic studies ( Conroy-Dalton & Huys, 2000; Conroy-Dalton, 2003a; George & Gheerardyn, 2015; George, 2017, 2018).

Although some recent studies have advocated a monophyletic Ancorabolidae ( George, 2006c), there now exists a general consensus that the relationship between the two subfamilies, Ancorabolinae and Laophontodinae , is not well understood and that at least one of them (e.g. Gheerardyn & George, 2010), if not the entire family ( George & Müller, 2013), is paraphyletic. Although the phylogeny of the Ancorabolinae appears largely to be resolved with the recognition of a basal divide into an Ancorabolus group ( Conroy-Dalton & Huys, 2000) and a Ceratonotus group ( Conroy-Dalton, 2001), this outcome was disputed to some extent in subsequent work and, unfortunately, created unwarranted animosity (e.g. George, 2006a, b, c). Little progress has been made towards elucidating the phylogenetic interrelationships in the Laophontodinae , but some recently discovered apomorphic character states appear to lend support to its monophyletic status ( Gheerardyn & George, 2010; Gheerardyn & Lee, 2012).

Despite their global distribution, representatives of the family Ancorabolidae (subfamily Laophontodinae ) have been recorded on only four occasions in East Asia. Zhang & Li (1976) reported Laophontodes hamatus ( Thomson, 1883) from seaweed washings collected in Yongxing Island (= Woody Island) and the Xisha (= Paracel) Islands in the South China Sea. Baldari & Cottarelli (1986) described Paralaophontodes elegans Baldari & Cottarelli, 1986 from a sandy beach on Mindoro Island, Philippines. Recently, Kim (2013) recorded Laophontodes bicornis A. Scott, 1896 from Seogwipo on Jeju Island, South Korea, where he recovered a small number of specimens from washings of wood infested by limnoriid isopods and teredinid bivalves and from coralline sand collected at moderate depths. In a later report, Kim (2014) listed Paralaophontodes psammophila ( Soyer, 1975) and Algensiella boitanii Cottarelli & Baldari, 1987 from the same locality and habitat as La. bicornis , but both records are highly dubious and should be discarded (see below - Authenticity of Laophontodes bicornis in Korea). Members of the Ancorabolinae have remained unrecorded from this part of the world so far. Here, we describe two new deep-water representatives of the latter subfamily, both belonging to the Ceratonotus group and collected from fine sandy deposits at 105 m depth off the south coast of the Korean peninsula. This contribution also: (1) addresses the taxonomic position of two genera of uncertain affinity, Patagoniaella Pallares, 1968 and Ancorabolina George, 2006c ; (2) reviews the status of certain species in Laophontodes T. Scott, 1894 ; and (3) provides updated keys to the species of Laophontodes and to the genera of both Ancorabolinae and Laophontodinae .