Vol.:(0123456789) 1 3
Polar Biology
https://doi.org/10.1007/s00300-018-2438-7
ORIGINAL PAPER
Species identity and genetic structure of nemerteans of the “Lineus
ruber–viridis” complex (Muller, 1774) from Arctic waters
Irina A. Cherneva
1
· Alexei V. Chernyshev
2,3
· Irina A. Ekimova
1,3
· Neonila E. Polyakova
2
·
Dimitry M. Schepetov
1,4
· Sergei V. Turanov
2
· Tatiana V. Neretina
1
· Elena M. Chaban
5
· Vladimir V. Malakhov
1,3
Received: 13 July 2018 / Revised: 16 November 2018 / Accepted: 19 November 2018
© Springer-Verlag GmbH Germany, part of Springer Nature 2018
Abstract
Lineus species are the most abundant nemerteans that inhabit boreal regions of the Atlantic and Arctic Oceans and are the
focus of many zoological studies. However, its taxonomy and species composition in Arctic areas remain poorly known. In
this study, we performed an updated genetic analysis of nemerteans from the “Lineus ruber–viridis” complex, clarifed its
species composition in the White Sea, and identifed their diagnostic features. Molecular data based on mtDNA analysis
indicate the presence of L. ruber, L. viridis, and L. clandestinus in the studied area, which was consistent with morphological
data. Short descriptions of external morphological features are provided. External morphologies of European and White Sea
Lineus species have some peculiarities: L. clandestinus from the White Sea lacks the iridescent ventral fold, and L. ruber
has an unusual “green” form. The recovered genetic structure difered among the three species, which may have resulted
from diferent dispersal scenarios during the last glacial maximum. The low level of nucleotide diversity combined with the
typical star-like structure of the network indicated that an ancestral northern L. ruber population underwent a bottleneck
event in the recent historical past.
Keywords Lineus · Nemertea · Integrative taxonomy · Field identifcation · Arctic · Population genetics
Introduction
Lineus ruber (Müller 1774) and L. viridis (Müller 1774)
are the most abundant nemertean species that inhabit
boreal regions of the Atlantic Ocean and some Arctic seas
(White, Barents, Kara and Chukchi Seas; see Chernyshev
and Maslakova 2001, 2011). Thus, the “Lineus ruber–vir-
idis” species complex is a convenient model for morpho-
logical and developmental studies as specimens are easy to
acquire for scientists in diferent regions. Previously many
zoological studies examined their epidermal ultrastructure
(Gontcharoff and Lechenault 1966; Storch and Welsch
1972), proboscis (Gontcharof 1957; Ling 1971), sensory
organs (Ling 1970; Storch and Moritz 1971), foregut (Ling
and Willmer 1973), nephridia (Bartolomaeus 1985), and
other morphological features. Developmental studies of
both species (Schmidt 1930, 1941, 1964; Gontcharof 1960;
Alluchon-Gerard and Gontcharof 1970) are often cited in
many textbooks (Schmidt 1953; Gontcharof 1961; Ivanova-
Kazas 1975; Friedrich 1979). Several detailed investigations
also examined their regeneration capabilities (Gontcharof
1951; Bierne 1962; Zhuravleva et al. 1970). Lineus ruber
and L. viridis were described by O.F. Müller (1774) more
than 200 years ago. However, the original description did
not provide sufcient information on the morphology and
sampling locations of these species (Gibson 1995). Subse-
quently, many zoologists referred to these nemerteans as a
Electronic supplementary material The online version of this
article (https://doi.org/10.1007/s00300-018-2438-7) contains
supplementary material, which is available to authorized users.
* Alexei V. Chernyshev
nemertea1969@gmail.com
1
Lomonosov Moscow State University, Leninskie gory 1/12,
Moscow, Russia 119234
2
National Scientifc Center of Marine Biology, Far Eastern
Branch RAS, Vladivostok, Russia 690041
3
Far Eastern Federal University, Sukhanova str., 8,
Vladivostok, Russia 690950
4
Koltzov Institute of Developmental Biology RAS, Vavilov
Str. 26, Moscow, Russia 119334
5
Zoological Institute RAS, Universitetskaya Emb. 1,
St. Petersburg, Russia 199 034