Zoosyst. Evol. 93 (2) 2017, 211–241 | DOI 10.3897/zse.93.11314 museum für naturkunde Phylogeny of the taxon Paralaophontodes Lang (Copepoda, Harpacticoida, Laophontodinae), including species descriptions, chorological remarks, and a key to species Kai Horst George1 1 Senckenberg am Meer, Abt. Deutsches Zentrum für Marine Biodiversitätsforschung DZMB, Südstrand 44, D-26382 Wilhelmshaven, Germany http://zoobank.org/32051770-28D6-4A10-8321-BF82758AA0D6 Corresponding author: Kai Horst George ([email protected]) Abstract Received 24 November 2016 A detailed phylogenetic analysis of the taxon Paralaophontodes Lang (Copepoda, Har- Accepted 8 March 2017 pacticoida, Laophontodinae Lang) based on morphological characters is presented. The Published 22 March 2017 monophylum Paralaophontodes is supported by 16 unambiguous autapomorphies such as the presence of characteristic dorsal processes on cephalothorax and body somites, Academic editor: a 5-segmented male antennule, the loss of the syncoxal seta on the maxilliped, and the Michael Ohl endopodal strengthening of the first swimming leg. The corresponding extensive phy- logenetic evaluation includes the description of Paralaophontodes anjae sp. n. from a Key Words beach on Chiloé Island (Chile), the re-description of Laophontodes armatus Lang, and the re-establishment of Paralaophontodes robustus (Bŏzić), the displacement of Laop- Crustacea hontodes armatus, L. hedgpethi Lang and L. psammophilus Soyer to Paralaophontodes, Meiofauna a discussion on relationships within that taxon, remarks on its geographical distribution, Ancorabolidae and a key to the species. systematics taxonomy geographical distribution Introduction Laophontodinae, relocating L. echinata into the new ge- nus as Paralaophontodes echinatus (Willey, 1930) while Lang (1936) erroneously synonymized Laophonte ech- retaining L. armatus within Laophontodes T. Scott, 1894. inata Willey, 1930 (Laophontidae T. Scott, 1905) with Lang (1965) transferred Laophontodes robustus Bŏzić, Laophontodes armatus Lang, 1936 (Laophontodinae 1964 from the Island La Reunión, Indian Ocean, to his Lang, 1944 in the paraphylum Ancorabolidae Sars, newly created genus Paralaophontodes. However, Lang 1909), owing to peculiar shared derived features such (1965) doubted the distinct specific status of P. robustus, as two lateral extensions and moderately long hairy el- pointing to its strong similarity with P. echinatus. Wells ements mid-dorsally on the cephalothorax, and a par- and Rao (1987) formally synonymized P. robustus with P. ticular dorsal armature of the body somites (Lang 1936). echinatus. At present the taxon Paralaophontodes contains Obvious differences between L. echinata and L. armatus three species, namely P. echinatus, P. elegans Baldari and were according to Lang (1936) due to misinterpretations Cottarelli, 1986, and P. exopoditus Mielke, 1981. by Willey (1930). Particularly noticeable differences in- When describing Laophontodes hedgpethi Lang, 1965, clude: the P1 exp being 2-segmented in L. echinata and Lang (1965) noted a strong similarity of that species with 3-segmented in L. armatus, and; the P1 enp-2 carrying L. armatus and Paralaophontodes echinatus. However, 1 terminal claw in L. echinata, but a terminal claw and he did not carry out the consequent step by unifying all an additional long seta in L. armatus. Twenty-nine years corresponding species, which remain therefore in the later Lang (1965) himself recognized his error and estab- laophontodin genera Laophontodes and Paralaophon- lished the taxon Paralaophontodes Lang, 1965 within todes, respectively. The objective of this contribution is Copyright Kai Horst George. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 212 George, K.H.: The taxon Paralaophontodes reconsidered to update the systematics of Paralaophontodes by the disposal by Mrs Karin Sindemark Kronestedt, Swedish Mu- inclusion of all respective species hitherto assigned to seum of Natural History, Department of Invertebrate Zool- Laophontodes. To support this synonymisation, a detailed ogy, Stockholm, Sweden. It consists of eight individuals: phylogenetic analysis based on all available morpholog- six females and two males are complete and of moderate ical characters is presented. In that context, a re-descrip- condition but covered with compact detritus. Moreover, tion of Laophontodes armatus is given, along with the de- four fragments are included: SMNH 2158(a): one female scription of a new Paralaophontodes species, P. anjae sp. cephalothorax and thorax (until swimming leg P3); SMNH n., from Chiloé Island (Chile). Moreover, P. robustus is 2158(b): one female thorax (P2–P4); SMNH 2158(c): two re-established, new records of known Paralaophontodes female urosomes. As neither a holotype nor any paratypes species are documented, and a key to the known species have been designated, the above named specimens are of Paralaophontodes is provided. henceforth declared as syntypes. Additional material was collected in the Magellan Straits, Chile; one female (coll. no. SMF 37111/1) was sampled by the author on 27.05.1996 Material and methods from Bahía Catalina in the north of Punta Arenas (Chile) (53°7.008’S, 70°52.323’W, 0m depth), and one female and Paralaophontodes anjae sp. n.: one single female was one male (coll. nos. SMF 37112/1 and SMF 37113/1–7, collected by the author on 27.03.1994 in Puente Quilo respectively) were kindly provided by Dr Matthias Gorny (Gulf of Quetalmahue), Chiloé Island, Chile. The mate- (Santiago de Chile, Chile) from sampling at Punta Yar- rial was sampled at low tide at station #Q4 directly at tou, Canal Whiteside, Tierra del Fuego (53°53.723’S, the waterline (Fig. 1). The substrate consists of medi- 70°09.132’W, 15m depth) on 15.02.2003. The Patagonian um-sized sand, the measured temperature was 12°C, and specimens are kept in the collection of Senckenberg For- the salinity was 32.3. Material was immediately fixed schungsinstitut und Naturmuseum Frankfurt, Germany. with 4% non-buffered formalin and later on sorted under Species identification and drawings were made with a binocular in the laboratories of the Instituto de Biología the use of a camera lucida on a Leica DMR compound Marina “Dr Jürgen Winter” at the Universidad Austral de microscope equipped with differential interference con- Chile, Valdivia, Chile. trast. General terminology follows Lang (1948) and Huys and Boxshall (1991). Terminology referring to phyloge- Additional Paralaophontodes material netic aspects follows Ax (1984); the terms “telson” and Paralaophontodes echinatus (Willey, 1930): two females “furca” are adopted from Schminke (1976). and one male were collected by Mr Johannes Dürbaum Abbreviations used in the text: A1 = antennule, A2 = (Jülich, Germany) from an intertidal mangrove-mudflat antenna, aes = aesthetasc, benp = baseoendopod, cphth = at Bering Point village, south-east of Andros Island, Ba- cephalothorax, DP I–DP IX = dorsal processes I–IX, enp hamas, in July 1993 (cf. Seifried and Dürbaum 2000) = endopod, enp-1–enp-3 = endopodal segments 1–3, exp and kindly put to the author’s disposal. The material is = exopod, exp-1–exp-3 = exopodal segments 1–3, FR = kept in the collection of Senckenberg Forschungsinstitut furcal rami, GF = genital field, P1–P6 = swimming legs und Naturmuseum Frankfurt, Germany, coll. nos. SMF 1–6, R = rostrum, T = telson. 37104/1, SMF 37105/1, and SMF 37106/1. One male and one CI copepodid was sampled by the author on June 29th, 2008 at the port of Golfito (Pacific coast of Costa Rica). Results The material is kept in the collection of Senckenberg For- schungsinstitut und Naturmuseum Frankfurt, Germany, Phylum Arthropoda coll. nos. SMF 37107/1 (male), SMF 37108/1 (CI). Subphylum Crustacea Brünnich, 1772 Paralaophontodes exopoditus Mielke, 1981: one female Superclass Multicrustacea Regier et al., 2010 collected by Dr Gritta Veit-Köhler (Wilhelmshaven, Ger- Class Hexanauplia Oakley et al., 2013 many) at the coastline of Dahab (28°29.0’N, 34°30.0’E, Subclass Copepoda Milne-Edwards, 1840 Gulf of Akaba, Egypt) in summer 1995. The specimen was Order Harpacticoida Sars, 1903 kindly put to the author’s disposal and is kept in the collec- Family “Ancorabolidae” Sars, 1909 tion of Senckenberg Forschungsinstitut und Naturmuseum Subfamily Laophontodinae Lang, 1944 Frankfurt, Germany, coll. no. SMF 37109/1. One male Genus Paralaophontodes Lang, 1944 collected by Prof. Dr Horst Kurt Schminke (Oldenburg, Germany) at station PNG M4, Ednago Island, near Kavi- Type species. P. echinatus (Willey, 1930). Additional spe- eng (2°35’S, 150°5’E, New Ireland, Papua New Guinea, cies: P. anjae sp. n., P. armatus (Lang, 1936) comb. n., P. cf. Seifried 2003), on 11.11.1984, and kindly provided to elegans Baldari and Cottarelli, 1986, P. exopoditus Mielke, the author. The individual is kept in in the collection of 1981, P. hedgpethi (Lang, 1965) comb. n., P. psammophi- Senckenberg Forschungsinstitut und Naturmuseum Frank- lus (Soyer, 1974) comb. n., P. robustus (Bŏzić, 1964). furt, Germany, coll. no. SMF 37110/1. Laophontodes armatus Lang, 1936: The type material Amended generic diagnosis. Laophontodinae. Body (coll. no. Type SMNH 2158) was kindly put to the author’s slender and cylindrical. Cphth with triangular extensions zse.pensoft.net Zoosyst. Evol. 93 (2) 2017, 211–241 213 medio- and postero-laterally