European Journal of Taxonomy 212: 1-31

http://dx.doi.org/10.5852/ejt.2016.212

ISSN 2118-9773

www. europeanj ournaloftaxonomy. eu

2016 • Holynska M. & Dimante-Deimantovica I.

This work is licensed under a Creative Commons Attribution 3.0 License.

Research article

urn:lsid:zoobank.org:pub:F69CE441-CF59-4CEl-BC52-782A79DD3626

Redescription and taxonomic notes on Cyclops bohater Kozminski, 1933

and Cyclops lacustris G.O. Sars, 1863 (Arthropoda, Crustacea), with

an identification key to the Cyclops species of Fenno-Scandinavia

Maria HOLYNSKA 1 * & Inta DIMANTE-DEIMANTOVICA 2

Museum and Institute of Zoology Polish Academy of Sciences, Wilcza 64, 00679 Warszawa, Poland.

2 Norwegian Institute for Nature Research, Gaustadalleen 21, NO-0349 Oslo, Norway.

"Corresponding author: [email protected]

2 E-mail: inta deimantovica@gmail com

1 urn:lsid:zoobank.org:author:DEFCAC09-2B6A-4030-87D9-A317F189763C

2 urn:lsid:zoobank.org:author:A52AB433-E7AE-4886-A534-6354DF9CE104

Abstract. The studies of Cyclops O.F. Muller, 1776 have more than two centuries of tradition in Fenno-

Scandinavia, yet, basic taxonomic questions remain unresolved and the limits of the distributional area

are very poorly understood in the majority of the taxa (11+) occurring in the region. Based on the types

and other materials we redescribe two species, C. lacustris G.O. Sars, 1863 and C. bohater Kozminski,

1933, the latter being reported here for the first time from Fenno-Scandinavia; lectotypes of both

species have been designated herein. Setulation characters of the antennal coxobasis- and maxillulary

palp setae and PI medial spine, as well as the spine formula of the terminal exopodal segments of the

swimming legs (3433) in the adult female distinguish C. lacustris and C. bohater from their congeners.

An identification key is provided to the Cyclops species of Fenno-Scandinavia, including information

about both the female and male morphology. Although the limited material does not allow us to define

the species range limits, our data nevertheless support a Northern European distribution in C. lacustris ,

and Northern and Central European distribution in C. bohater , while the records from the southern

Palearctic need further verification.

Key words. Cyclopidae, morphology, zoogeography, freshwater, Europe.

Holynska M. & Dimante-Deimantovica 1.2016. Redescription and taxonomic notes on Cyclops bohater Kozminski,

1933 and Cyclops lacustris G.O. Sars, 1863 (Arthropoda, Crustacea), with an identification key to the Cyclops

species of Fenno-Scandinavia. European Journal of Taxonomy 212: 1-31. http://dx.doi.org/10.5852/ejt.2016.212

Introduction

Cyclops O.F. Muller, 1776, frequently being a dominant component of the zooplankton in the cold

temperate and Arctic lakes, has been the focus of various fields of biology (Kozminski 1927, 1933;

Wierzbicka 1934; Findberg 1957; Einsle 1975, 1985, 1996a; Elgmork & Halvorsen 1998; Krajicek

et al. in press). Nevertheless, in several representatives of the genus very basic taxonomical issues

have hitherto remained unresolved, and the geographic distribution of most species is poorly known.

European Journal of Taxonomy 212: 1-31 (2016)

Table 1. Cyclops species of Fenno-Scandinavia (Herbst 1951; Lindberg 1957; Einsle 1968, 1996a;

Silfverberg 1999; Boxshall & Defaye 2013; this paper). + = species present; ? = expected in Denmark,

as the species occurs in northern Germany and Scandinavia; (1) = this paper; (2) = unpublished data of

MH; * = Cyclops abyssorum divulsus Lindberg, 1956 is here considered as a junior synonym of Cyclops

divergens Lindberg, 1936; ** = the records may refer to more than one species.

Denmark

Norway

Sweden

Finland

C. abyssorum G.O. Sars, 1863

C. bohater Kozminski, 1933

+ (1) (Largen)

C. divergens Lindberg, 1936*

+ (Bornholm)

+ (2) (Lund)

C. furcifer Claus, 1857**

C. insignis Claus, 1857

C. kikuchii Smirnov, 1932

+ (2) (Lund)

C. kolensis Li 1 ljeborg, 1901

C. lacustris G.O. Sars, 1863

C. scutifer G.O. Sars, 1863

C. strenuus Fischer, 1851

C. vicinus Uljanin, 1875

Morphometric characters based on the adult female morphology and used until the eighties of the

last century in Cyclops taxonomy are not so helpful in the identification of the male and larvae. Also,

certain morphometric and body shape traits (e.g., body length, relative width of the cephalothorax,

pedigers 4 and 5, P4 enp3, and relative length of the caudal rami and caudal setae) show seasonal and/

or environment-dependent variation within species (Einsle 1969, 1975; Elgmork & Halvorsen 1998),

or in other cases they fail to separate evolutionary lineages that diverged in ecological, cytogenetic

and enzyme electrophoretic characteristics (Einsle 1996b). Morphology nonetheless is a powerful tool

in species delineation when the characters are extended to the surface microstructures of the limbs

and somites, as it has been demonstrated in all recently revised cyclopid genera (papers pertaining to

Cyclops : Einsle 1985; Brandi & Lavicka 2002; Holynska & Dahms 2004; Holynska 2008).

The predominantly Palearctic Cyclops (31 putative species) is represented by at least 11 species in

Fenno-Scandinavia (Table 1).

The aim of our paper is to redescribe two species occurring in Fenno-Scandinavia, C. lacustris G.O. Sars,

1863 and Cyclops bohater Kozminski, 1933, based on original material from the type localities and other

specimens from different parts of the species ranges, and to provide a (partial) revision of the geographic

distributional data. Updated morphological information, supplemented with an identification key to

all reported and expected species of Cyclops in Fenno-Scandinavia, may significantly facilitate the

limnological studies in the region, and is also needed for a comprehensive phylogenetic reconstruction

of the evolutionary history of the genus.

Material and methods

Qualitative and morphometric characters were examined in the types and other specimens of C. bohater

(adult $ and and copepodid IV instar) and C. lacustris (adult $ and S, and copepodid IV-V instars)

collected in Northern and Central Europe. The collecting sites are listed in the species descriptions. Most

of the specimens were observed and measured in glycerine, and for a few microscope slides (Kiefer

Collection, Karlsruhe; Sars Collection, Oslo) the mounting medium was unknown to us. Telescoping

somites were measured separately and summed for total body length (Kozminski 1936). Width of a

HOFYNSKAM. & DIMANTE-DEIMANTOVICA I., Redescription of Cyclops bohater and C. lacustris

segment was measured across its widest part. Drawings were made using a camera lucida attached to

Olympus BX 50 or Nikon Ci-L compound microscopes.

Morphological abbreviations (caudal setae are coded according to Huys & Boxshall 1991)

Abd

A e

Caudal seta I

Caudal seta II

Caudal seta III

Caudal seta IV

Caudal seta V

Caudal seta VI

Caudal seta VII

CIV-CV

Enpl-3

Exp1-3

Mxl

P1-P4

Ante nn ule

Ante nn a

Abdomen

Aesthetasc

Anterolateral accessory

Anterolateral

Posterolateral

Outer terminal

Inner terminal

Terminal accessory

Dorsal

Fourth to fifth copepodid instar

First to third endopodal segment

First to third exopodal segment

Maxillule

First to fourth swimming leg

Institutional abbreviations:

IDD

MIZ

NHMUK

SMNK

ZMFU

ZMO

Results

private collection of Inta Dimante-Deimantovica

Museum and Institute of Zoology PAS, Warsaw

Natural History Museum, Fondon

Staatliches Museum fur Naturkunde, Karlsruhe

Zoological Museum, Fund University

Zoological Museum, Oslo

Taxonomy

Phylum Arthropoda von Siebold, 1848

Subphylum Crustacea, Brunnich, 1772

Subclass Copepoda Milne-Edwards, 1840

Order Cyclopoida Burmeister, 1835

Family Cyclopidae Rafinesque, 1815

Genus Cyclops O.F. Muller, 1776

Cyclops bohater Kozminski, 1933

Figs 1-6

Cyclops bohater Kozminski, 1933: 105-106, 111-113, 123-124, 135, 138-139, figs 5-6, pi. 3, tab. 3.

Cyclops bohater- Findberg 1957: 96-97, fig. 55. —Einsle 1975: 100-102, 147, fig. 3; 1993: 136-137,

fig. 80; 1996a: 28-31, fig. 18. — ? Stankovic&Ternjej 2007: 189-199, figs 2-12, tab. 2.—Deimantovica

2010 (part) (records from Fake Brigene only).

non Cyclops bohaterponorensis Naidenow & Pandurski, 1992: 27-30, fig. 1, tabs 1-2; synonymized by

Pandurski (1997) as C. abyssorum divergens Findberg, 1936.

non Cyclops bohater - Deimantovica 2010: 216-222, figs 2-5, tabs 1, 3M.

European Journal of Taxonomy 212 : 1-31 ( 2016 )

Type material

Lectotype (designated here)

POLAND: 5, Wigry lakes, —132 m a.s.l., Lake Okrggle (~0.141cm 2 surface area) close to the southern bay

of Lake Wigry proper, 54°01' N, 23°01' E, 1 Mar. 1927, leg. and det. Z. Koz in in ski (NHMUK 2016. 38).

Fig. 1. Cyclops bohater Kozminski, 1933, A. Habitus, dorsal. B. Pediger 5 and genital double¬

somite, ventral, arrow points to lateral “wing-like” protrusion of pediger 5. C. Anal somite and caudal

rami, dorsal. D-G. Antennule, ventral. D. Segments 1-6 (I-XIV). E. Segments 7-14 (XV-XXIII).

F. Segments 12-14 (XXI-XXIII). G. Segments 15-17 (XXIV-XXVIII). Except for A (NHMUK 2016.

39) all drawings show the lectotype (NHMUK 2016. 38). Scale bars: A = 1 mm; B-G = 100 pm.

HOLYNSKAM. & DIMANTE-DEIMANTOVICA I., Redescription of Cyclops bohater and C. lacustris

Paralectotypes

POLAND: 1 § in alcohol (NHMUK 2016. 39) and 1 $ dissected (NHMUK 2016. 40), same locality

and collecting date as lectotype; 1 $ dissected on two slides (SMNK: 4396-4397) and 1 undissected $

Fig. 2. Cyclops bohater Kozminski, 1933, §. A-B. Surface ornamentation of antennal coxobasis.

A. Frontal, arrow points to insertion site of long spinules missing in C. bohater. B. Caudal, arrow points

to a group of spinules sometimes present. C. Labrum, epistoma and rostrum. D. Mandible, frontal,

arrow points to insertion site of spinules missing in C. bohater. E. Maxillulary palp, arrows point to long

setules of the setae, and spinulose ornamentation of the palp. All drawings show the lectotype (NHMUK

2016.38). Scale bars: A-E = 100 pm.

European Journal of Taxonomy 212 : 1-31 ( 2016 )

(SMNK: 4398), “Wigryseen”, 54°01' N, 23°06' E, 31 Jan. 1928, leg. et det. Kozminski, “paratype”, prep.

Fr. Kiefer 13 Dec. 1939.

Other material examined

AUSTRIA: 1 §, Lake Worth (Carinthia), 46°38' N, 14°09' E, 439 m a.s.l., Knut Lindberg’s Collection,

nr. 25, “Autriche AUT.l” (ZMLU).

LATVIA: 1 CIV $ toto (MIZ 2/2016/1) and 9 CIV parts (A2, Mxl, PI), Latgale region, Augshzeme,

Lake Brigene, -148 m a.s.l., 55°45’ N, 26°33’ E, 12 May 2008, leg. I. Deimantovica (MIZ 2/2016/2-10).

Fig. 3. Cyclops bohater Kozminski, 1933, §. A. Maxilla, caudal. B. Maxilliped, frontal. C. Leg 1,

frontal, arrows point to the long setules of the medial spine/seta of basipodite, and long spinules on the

basipodite between the insertions of exopodite and endopodite. D-E. Leg 4: EXP, P4 exp 3; ENP, P4

enp3. All drawings show the lectotype (NHMUK 2016. 38). Scale bars: A-E = 100 pm.

HOFYNSKAM. & DIMANTE-DEIMANTOVICA I., Redescription of Cyclops bohater and C. lacustris

SWEDEN: 1 $ (MIZ 2/2016/11), Fargen, 59°35'N, 18°3 E E, ~35 km NE of Stockholm, 3 Aug. 2011,leg.

Oslo University; one $ dissected (MIZ 2/2016/12) and 1 undissected $ in alcohol (IDD), 3 S<5 dissected,

same locality, 8 Nov. 2012, leg. I. Dimante-Deimantovica & B. Walseng, (MIZ 2/2016/13-15).

Description

Female

Body length 1960-2500 pm (lectotype, 2370 pm) (Fig. 1A). Cephalothorax length/width 1.05-1.14

(lectotype, 1.10), cephalothorax width/genital double-somite width 2.6-2.9 (lectotype, 2.9), prosome

length/urosome length 1.5-1.8 (lectotype, 1.8). Pediger 2 (Fig. 1A) bearing distinct posterolateral lobes

and conspicuously wider than succeeding somites. Pediger 5 laterally protruding, “wing-like” (Fig. IB,

arrowed).

Genital double-somite (Fig. IB) slightly shorter than, or as long as wide. Seminal receptacle as typical

in genus, transverse ridge present next to copulatory pore. Posterior margin of anal somite (Fig. 1C)

with continuous row of spinules, anal sinus without surface ornamentation, anal operculum weakly

developed. Caudal rami (Fig. 1C) 5.0-6.5 times as long as wide, medial margin bearing hairs. Midline

crest running almost full length of ramus on dorsal surface. Few spinules present on lateral margin

-D, E

-F

Fig. 4. Cyclops bohater Kozminski, 1933, <$■ A. Habitus, dorsal. B. Prosome with distinctly wide

pediger 2. C. Pediger 5, genital segment, and first abdominal segment, ventral. D=E. Antennal coxobasis.

D. Frontal. E. Caudal. F. Fabrum. Except for B (Fake Fargen, Sweden - MIZ 2/2016/15) all drawings

show a paralectotype (NHMUK 2016. 40). Scale bars: A-B = 500 pm; C-F = 100 pm.

European Journal of Taxonomy 212: 1-31 (2016)

in anterior third, indicating presumptive insertion site of the ancestral anterolateral accessory seta (I).

Spinules also present at insertion of caudal setae II and III. Seta II inserted at distance of 0.21-0.23

ramus length measured from posterior end. Relative length of caudal setae VII, VI, V, IV, and III:

Fig. 5. Cyclops bohater Kozminski, 1933, <$■ A. Maxillulary palp. B. Maxilliped syncoxopodite, frontal.

C. Leg 1, frontal. D. Leg 2, slender spine of first exopodal segment and robust spines of exopodal

segments 2 and 3. E. Exopodal armature of leg 3. F. Leg 4 coxopodite, basipodite and intercoxal sclerite,

caudal. All drawings show a paralectotype (NHMLJK 2016. 40). Scale bars =100 pm.

HOLYNSKAM. & DIMANTE-DEIMANTOVICA I., Redescription of Cyclops bohater and C. lacustris

1.0-1.2 (dorsal seta broken in lectotype), 1.7-2.2 (lectotype, 2.2), 3.7-4.4 (lectotype, 4.0), 2.8-3.6

(lectotype, 3.5), 1.0. Seta VI (medialmost) 1.1-1.3 (lectotype, 1.2) times as long as caudal rami, seta V

(longest) 0.66-0.80 (lectotype, 0.72) times as long as urosome. Caudal setae homonomously setulose,

setae IV and V with breaking plane.

Antennule (Fig. 1D-G) 17-segmented (I-V, VI-VII, VIII, IX-XI, XII-XIII, XIV, XV-XVI, XVII,

XVIII, XIX, XX, XXI, XXII, XXIII, XXIV, XXV, XXVI-XXVIII) and reaching middle of pediger

2 to middle of pediger 3. Seta formula as common in plurisegmented Cyclopinae: 8, 4, 2, 6, 4, 1+sp,

2, 1, 1, 0, 1, 1+ae, 0, 1, 2, 2+ae, 7+ae. Aesthetasc on segment 12 (Fig. 1E-F ) reaching distal margin

of segment 13 to middle of segment 14, aesthetasc length 2.2-2.8% of body length (n = 3; lectotype,

2.24%). Aesthetasc on segment 16 not reaching middle of segment 17, aesthetasc length 2.1-2.7% of

body length (n = 3; lectotype, 2.1%). Spinules only present on ventral surface of segment 1. Last three

segments of ante nn ule with hyaline membrane.

Antenna composed of coxobasis and three-segmented endopodite, and bearing 3,1,9 (8 in Lake Largen,

Sweden - MIZ 2/2016/11) and 7 setae, respectively. Inner medial seta of coxobasis (Fig. 2A-B) with

short setules, outer medial seta naked. Exopodite seta reaching beyond enp3 and bearing setules

conspicuously longer in proximal than in distal section of seta. Frontal surface of coxobasis (Fig. 2A)

with few spinules in -4/10 of segment and longer spinules more proximally next to lateral margin, and

distinctly long spinules in oblique/longitudinal row near middle line in proximal third of segment. Caudal

surface of coxobasis (Fig. 2B) with double (single in Lake Largen, Sweden, - MIZ 2/2016/12) row of

long spinules near long proximal spinules on lateral margin, elongate spinules (7-13) in longitudinal

row(s) near lateral margin distally to double row, and oblique field of small spinules proximally to

insertions of medial setae. Additional groups, such as small spinules medially to proximal double row

(Fig. 2B, arrowed) and row of tiny spinules along presumptive proximal margin of basipodite (Lake

Worth, Austria), sometimes present.

Labrum (Fig. 2C) bearing teeth on arcuate distal margin, lateral lobes with small spinules. Distal hairs

long and arranged in two groups. Epistoma and vertical cleft (Fig. 2C) naked, except for few hair-like

elements on one side only next to epistoma (observed in lectotype, but missing in § from Lake Worth,

Austria). Mandibular palp (Fig. 2D) with two long and one short seta, no surface ornamentation next and

proximal to palp. Maxillule: setation of precoxal arthrite as typical in plurisegmented Cyclopinae. Palp

with spinules (Fig. 2E, arrowed), and bearing long proximal seta, three setae on lateral lobe and three

apical setae; proximal seta and at least one of lateral lobe setae with distinctly long setules (Fig. 2E,

arrowed). Maxilla (Fig. 3A): precoxopodite and coxopodite fused on frontal surface but separated on

caudal surface; syncoxopodite, basipodite and two-segmented endopodite with 5, 2, 2 and 3 setae,

respectively. Claw-like attenuation of basipodite with spinules on both concave (inner) and convex

(outer) margin. Basipodite seta inserted in front of claw-like attenuation, bearing long setules on both

concave and convex edge in proximal half, and tiny spinules on convex margin in distal half of seta.

Maxilliped (Fig. 3B) composed of syncoxopodite, basipodite and two-segmented endopodite with 3,

2, 1 and 3 setae, respectively. On caudal surface of basipodite spinules arranged in two groups. First

endopodal segment and basipodite bearing hair-like spinules on frontal surface. Syncoxopodite frontally

adorned with medium-sized or small spinules midway between median and distalmost setae, and tiny

spinules more proximally next to finger-like membraneous element.

P1-P4 setation formula shown in Table 2.

Medial spine of PI basipodite with heteronomous setulation: setules long in proximal 4/10 (Fig. 3C,

arrowed), and short spinule-like more distally. Medial spine reaching slightly beyond distal margin of

enp2 to proximal A of enp3. Long spinules (Fig. 3C, arrowed) arranged in arc on frontal surface of PI

European Journal of Taxonomy 212: 1-31 (2016)

Table 2. Armature of legs 1-4 in adult Cyclops bohater Kozminski, 1933. Spines are denoted by Roman,

setae by Arabic numerals. The armature on the lateral margin of any segment is given first, followed by

the elements on the apical and medial margins.

Coxopodite

Basipodite

Exopodite

Endopodite

Leg 1

0-1

1-1

1-1; I—1; H-I, 2-3

0-1; 0-2; l-I,l-3

Leg 2

0-1

1-0

I—1; I—1; III—I, 1-4

0-1; 0-2; 1-1, 1-3

Leg 3

0-1

1-0

I—1; I—1; II-I, 1^1

0-1; 0-2; 1—I, 1-3

Leg 4

0-1

1-0

1-1; I—1; II-I, l^t

0-1; 0-2; 1-II-2

basipodite between insertions of exo- and endopodite. Intercoxal sclerites naked on frontal and caudal

surfaces in P1-P3, and sparsely pilose on caudal surface in P4. Obtuse protuberances of P4 intercoxal

sclerite low, hardly reaching beyond distal margin of segment, or extending well beyond it (Fig. 3D).

Medial expansion of basipodite pilose in P1-P3, and naked (Fig. 3D) or bearing few short apical hairs

(Lake Largen, Sweden - MIZ 2/2016/12; and Lake Worth, Austria) in P4. Caudal surface ornamentation

of P4 coxopodite (spinule groups are coded according to Einsle 1996a, see Fig. 3D) composed of:

undulate line of medium-sized spinules near proximal margin (group A), with 9-16 spinules in lateral

section, and 10-13 spinules in medial section; rows of spinules of unequal size along distal margin, with

3-7 spinules in medial row (group C) and 1-4 spinules in lateral row (group D); and oblique field of

long spinules at laterodistal angle (group E); spinules sometimes present at lateroproximal angle (group

B) (Lake Worth, Austria) and/or lateral margin (group F) (Lake Worth, Austria; paralectotype - SMNK:

4398). Coxopodal seta conspicuously longer (~1.9x) than height of medial expansion of basipodite.

Distal margin of first and second exopodal segment of P4 naked on caudal surface (Fig. 3D), or bearing

spinules. P4 enp3 2.6-2.9 times as long as wide; of apical spines medial one 1.6-2.3 times as long as

lateral, and 0.94-1.10 times as long as segment.

P5 (Fig. IB) segmentation and setation as typical in genus. Distal segment 2.0-2.3 times as long as

wide. Medial spine inserted near half-length of segment (0.45-0.55 segment length measured from

proximal margin), medial spine 0.5-0.9 times as long as distal segment. Lateral seta on proximal

segment relatively long, 0.6-0.9 times as long as apical seta and 1.9-3.0 times as long as distal segment.

Apical seta 3.1-4.0 times as long as distal segment. Spinules present at insertion of median spine and

apical seta, but absent at insertion of lateral seta.

Male

Body length 1525-1685 pm (Fig. 4A), cephalothorax length/width 1.1-1.3, prosome length/urosome

length 1.6-1.8. Pediger 2 without distinct posterolateral lobes, but sometimes significantly wider than

succeeding prosomal segments (Lake Largen, Sweden) (Fig. 4B). Pediger 5 (Fig. 4C) laterally not

protruding. Caudal rami without dorsal crest, 4.2-5.1 times as long as wide, pilose on medial margin.

Short transverse row of spinules present in anterior third at presumptive insertion site of ancestral

anterolateral accessory seta (I), as well as at insertion of caudal seta II and III. Seta II inserted at distance

of 0.24-0.26 ramus length measured from posterior end. Relative length of caudal setae VII, VI, V, IV,

and HI: 1.1-1.2, 2.1-2.2, 3.4-4.0, 2.9-3.3, 1.0. Seta VI 1.6-1.8 times as long as caudal rami, seta V

0.8-0.9 times as long as urosome.

Antennule 17-segmented (verified on dorsal surface): I-V, VI-VII, VIII, IX, X, XI, XII, XIII, XIV, XV,

XVI, XVII, XVIII, XIX-XX, XXI-XXIII, XXIV-XXV, XXVI-XXVIII. Armature formula 8+3ae, 4,

2, 2+ae, 2, 2, 2, 2, 1+sp+ae, 2, 2, 2, 2+ae, 2, 1+ae, 4+ae, 7+ae (setation of segment 13 and segment 17

were verified in males from Lake Largen, Sweden). Modified setae: one and two striated plates present

on segment 14 and 15, respectively; and one cone-like element present on segment 14 and 15 each.

HOLYNSKAM. & DIMANTE-DEIMANTOVICA I., Redescription of Cyclops bohater and C. lacustris

Antenna segmentation and setation as in female, except for enp2, which bears 8 or 7 setae. Spinule

ornamentation of antennal coxobasis similar to that in female (Fig. 4D-E), of medial setae inner one

bearing short setules and outer seta naked. Mouthparts (Figs 4F, 5A-B) similar to those in female, but

Fig. 6. Cyclops bohater Kozminski, 1933, copepodid IV, $. A-C. Antemiule, ventral. A. Segments

1-5 (I-XVT), arrow points to long anterodistal seta of ancestral segment XIV. B. Segments 6-8 (XVII-

XXIV). C. Segments 9-10 (XXV-XXVIII). D. Leg 2, caudal. E. Leg 3, caudal. All drawings show

specimen from Latvia (Lake Brigene - MIZ 2/2016/1). Scale bar = 100 pm.

European Journal of Taxonomy 212: 1-31 (2016)

Table 3. Armature of legs 1-4 in CIV instar of Cyclops bohater Kozminski, 1933 (coding system as in

Table 2).

Coxopodite

Basipodite

Exopodite

Endopodite

Leg 1

0-1

1-1

1-1; 11-1,2-3

0-1; 1—1,1-5

Leg 2

0-1

1-0

1-1; 111—1,1—4

0-1; 1—1,1-5

Leg 3

0-1

1-0

I—1; Hl-1,1-4

0-1; 1—1,1—4

Leg 4

0-1

1-0

1-0; HIM, 1-4

0-1; 1—II—3

spinules tiny or absent on maxillulary palp, and maxilliped syncoxopodite adorned with single group of

spinules midway between median and distalmost setae.

P1-P4 segmentation and setation as in female (Table 2). Medial spine of PI basipodite (Fig. 5C) with

few long setules proximally and short spinules more distally, and reaching distal margin of enp2 to

proximal ~l/6 of enp3. Tong spinules arranged in arc on frontal surface of PI basipodite between

insertions of exo- and endopodite. Lateral spine of P2 expl more slender than other spines of exopodite,

and bearing long setules (Fig. 5D cf. Fig. 5E). Intercoxal sclerites naked in P1-P3 and sparsely pilose on

caudal surface in P4. Caudal surface ornamentation of P4 coxopodite (Fig. 5F) similar to that in female,

but spinules/hairs missing on lateral margin. Coxopodal seta conspicuously longer (~1.7-1.8x) than

height of medial expansion of basipodite. Medial expansion of basipodite apically pilose in P1-P4. P4

enp3 2.9-3.2 times as long as wide. Of apical spines medial one 1.9-2.1 times as long as lateral spine

and as long or slightly (~l.lx) longer than segment.

P5 (Fig. 4C) distal segment 2.0-2.4 times as long as wide, medial spine inserted near half-length of

segment (0.47-0.58 segment length measured from proximal margin), medial spine 0.56-0.71 times as

long as distal segment. Lateral seta on P5 proximal segment 0.68-0.81 times as long as apical seta and

2.9-3.6 times as long as distal segment. Apical seta 4.1-4.8 times as long as distal segment. Spinules

present at insertion of median spine and apical seta, but absent at insertion of lateral seta.

P6 (Fig. 4C) composed of three elements; median- and lateral seta 0.94-1.05 times and 1.9-2.4 times as

long as median spine, respectively. P6 flap naked.

Copepodid IV instar (Lake Brigene, Latvia)

Copepodid shows some important species diagnostic characters, such as: inner medial seta of antennal

coxobasis with short setules and outer medial seta naked; proximal seta of maxillulary palp and at least

one seta of lateral lobe of palp with long setules; medial spine of PI basipodite with few distinctly

longer setules proximally and short spinules distally; long spinules arranged in arc on frontal surface of

basipodite between insertions of exo- and endopodite; P5: spinules present at insertions of medial spine

and apical seta, but absent at insertion of lateral seta; lateral seta (110 pm) relatively long (0.79x apical

seta length).

Other features: Anal somite with continuous row of spinules along posterior margin. Caudal rami

relatively short (175 pm), bearing medial hairs, longitudinal crest absent on dorsal surface. Short

transverse row of robust spinules present in anterior third at presumptive insertion site of anterolateral

accessory seta (I), and caudal seta II and III. Seta II (50 pm) inserted at distance of 0.31 ramus length

measured from posterior margin. Length of setae VII, VI, V, IV and III, 153 pm, 260 pm, 565 pm,

425 pm, and 137 pm, respectively.

HOLYNSKAM. & DIMANTE-DEIMANTOVICAI., Redescription of Cyclops bohater and C. lacustris

Table 4. Overview of the published records of Cyclops bohater Kozminski, 1933. For comments on the

data see “Discussion”. ? = dubious identification according to Maemets et al. 1996; * = world-catalogue

or regional checklist reports without specific information on the localities and authors; n.d. = no data; E

= East; N =

North; S = South; W = West.

Country

Region

Habitat

Altitude (m)

References

Sweden

Central (Svealand), L. Largen

lake

reported here

Estonia (?)

Peipus

lake

Maemets etal. 1996

Latvia

Latgale, L. Brigene

lake

148

Deimantovica 2010

Lithuania

Vilnius/Trakai

lakes

-100-150

Wierzbicka 1936

Ukraine

Dnieper

river

Monchenko 2003

Poland

NE: Suwalszczyzna

lakes

-130

Kozminski 1933

NW: Pomerania

lakes

50-135

Patalas 1954

Central: Warsaw

clay pit pond

-80-115

Wierzbicka 1960

Germany

N: Schleswig Holstein,

Lower Oder Valley

lakes and alluvial ponds

-5-45 m

Einsle 1968; Prisch 2002

S: Lake Constance

lake

395

Einsle 1964, 1996a

Austria

N: Salzburg, Upper Austria

lakes

-480-505

Gaviria 1998

S: Carinthia

lakes

-440-750

Einsle 1971;

Gaviria 1998

Croatia

NW: Drava River basin

gravel pit lake

119

Stankovic & Temjej 2007

Switzerland

W, NE and Central: Bern,

Zurich, Schwyz

lakes

-405-645

Kiefer & Einsle 1962;

Kiefer 1978; Stebler 1979

* Czech Rep.

n.d.

Dussart & Defaye 2006

Prance

E: PrenchAlps

eutrophic with

decomposed plants

-370-570

Dussart 1958

W: Pays de la Loire

n.d.

-70-170

Dussart 1969

Spain

N: Burgos

lakes

-1900

Velasco et al. 2005

* Italy

n.d.

Dussart & Defaye 2006

Georgia

W: Racha (L. Shaori)

dam

1105

Monchenko 1974

* Turkey

n.d.

Ustaoglu 2004

Antennule (Fig. 6A-C) 10-segmented, homologous with ancestral segments I-V, VI-XI, XII-XIII,

XIV, XV-XVI, XVII-XX, XXI-XXIII, XXIV, XXV, XXVI-XXVIII. Seta formula 5, 6,2,2,2,3,2+ae,

2, 2+ae, 7+ae. Anterodistal seta on segment 4 (XIV), spine-like in the adult, relatively long, reaching

slightly beyond distal margin of succeeding segment (Fig. 6A, arrowed). Length of aesthetascs on

segment 7 (XXI-XXIII) and 9 (XXV), 60 pm and 62 pm, respectively (Fig. 6 B-C). Second endopodal

segment of antenna bearing 7 setae. P1-P4 rami two-segmented (Fig. 6D-E), setation shown in Table 3.

P4: intercoxal sclerite naked, medial expansion of basipodite apically pilose; medial apical spine of

endopodite much longer (3.6x) than lateral apical spine. P5 distal segment relatively short (length,

36 pm; width, 28 pm), medial spine (24 pm) inserted at distance of 0.72 segment length, measured from

proximal margin. P6 armed with medial spine (50 pm) and lateral seta (81 pm).

Remarks

Cyclops bohater was originally described from Lake Wigry (max. depth 73 m, surface area 21.15 km 2 )

and some smaller lakes (Okrpgle, Dlugie, Muliczne and Rzepiskowe) within the vicinity of L. Wigry in

the Suwalki-Augustow Lake District, NE Poland (Kozminski 1933). The holotype of C. bohater was not

designated in the original description (Kozminski 1933), and we are not aware of any institution where

it might be deposited. The Hydrobiological Station at Wigry Lakes, where Zygmunt Kozminski worked

and his copepod material might be left, was completely destroyed during World War II (Gieysztor 1963).

We have not found any type(s) at the Museum and Institute of Zoology PAS (Warsaw) either, though

European Journal of Taxonomy 212: 1-31 (2016)

part of Kozminski’s samples (e.g., those from the Tatra Mts. and Warsaw region) are deposited here.

However, some specimens from the terra typica and identified by Kozminski himself as C. bohater

fortunately have remained in the collections of Staatliches Museum fur Naturlcunde in Karlsruhe and

Natural History Museum in London. The collecting data of these specimens suggest that they might

constitute the material which Kozminski’s original description was based on; therefore, we selected one

of those specimens as the lectotype of the species (NHMUK 2016.38). In choice of the lectotype the

NHM series was preferred to the material from Karlsruhe Museum, because more characters (incl. those

of the mouthparts and the morphometric traits) could be verified in the NHM specimens which were

dissected by us, and the collecting data of the NHM material were fully consistent with the information

provided in the original description.

Ecology

Permanent lakes and ponds, eutrophic to oligotrophic. The species is rather rare in the surface plankton,

both the larvae and adult seem to prefer the deep or near-bottom waters (Kozminski 1933, 1936;

Wierzbicka 1936). The same distribution pattern was observed in a population in Lake Largen, Sweden

(Dimante-Deimantovica & Walseng, unpublished data). Adults and copepodids occurred in large number

in the littoral zone of a clay pit pond in Poland (Warsaw) in March, but disappeared from the littoral in

late spring (Wierzbicka 1960). Einsle (1988) reported on the permanent near-shore occurrence of the

species in Lake Constance (Obersee).

Cyclops bohater usually has one generation per year: reproduction from November to February, youngest

copepodid instars appear early spring, copepodid V instar enters summer diapause in May and stays in

the sediment on the bottom of the lake until late autumn when the larvae moult to adults. (Einsle 1988;

Frisch 2002). A second summer generation appeared in the Wigry lakes (Kozminski 1933), and Einsle

(1988) also hypothesized more than one generation in the permanently littoral populations.

Distribution

Verified occurrences from Northern (Sweden, Latvia) and Central Europe (Poland, Austria) (Table 4). For

a critical overview of the literature data on the geographic distribution of C. bohater , see the Discussion.

Cyclops lacustris G O. Sars, 1863

Figs 7-10

Cyclops lacustris G.O. Sars, 1863: 30

Cyclops lacustris - Sars, 1914: 35-36, pi. 18.—Rylov 1948: 187-189, fig. 38, tab. 13.—Lindberg 1957:

85-87, figs 90-94, tab. 13. —Einsle 1993: 141-142, fig. 84; 1996a: 41-43, fig. 26. —Deimantovica

2010: 216-222, figs 2-5, tabs 1, 3M.

Cyclops strenuus f. lacustris -Lilljeborg 1901: 31-32, pi. 2, fig. 23.

Type material

Lectotype (designated here)

SOUTHERN NORWAY: dissected on two slides labelled as “lectotypus b” and “lectotypus c”. Lake

Mjosa, 60°42' N, 11°1' E, 121 m a.s.l., surface area 369.2 km 2 , max. depth 443 m, G.O. Sars coll., prep.

Fr. Kiefer, 27 Aug. 1971 (ZMO).

Paralectotypes

SOUTHERN NORWAY: 1 undissected $ labelled as “lectotypus a”, 1 S labelled as “lectotypus d”,

same locality as in lectotype, G.O. Sars coll., prep. Fr. Kiefer, 27 Aug. 1971 (both ZMO); 2 $5 an d

1 S dissected by the authors from vial “Sars Coll. 12977” (ZMO), and 1 $ and 1 $ dissected by the

HOLYNSKAM. & DIMANTE-DEIMANTOVICA I., Redescription of Cyclops bohater and C. lacustris

Fig. 7. Cyclops lacustris G.O. Sars, 1863, §. A. Habitus, dorsal (SMNK: 9567). B-C. Urosome,

ventral (Lake Aspem, Norway - MIZ 2/2016/20). B. Urosomites 1-4. C. Anal somite and caudal rami.

D. Ante nn ular segments 12—17, ventral (Lake Aspern, Norway - MIZ 2/2016/20). E—F. Ante nn al

coxobasis, caudal. E. $ from Lake Mjosa, Norway (ZMO: Sars Coll. 12977 $-2). F. $ from Lake

Aspem, Norway (MIZ 2/2016/20). G-H. Maxillulary palp. G. $ from Lake Mjosa, Norway (ZMO:

Sars Coll. 12977 $-2). H. § from Lake Aspern, Norway (MIZ 2/2016/20). Scale bars: A-H = 100 pm.

European Journal of Taxonomy 212: 1-31 (2016)

authors from vial “Sars Coll. 12976” (ZMO); two antenna of single $, “ Cyclops lacustris $ 2 A2 Kiefer

26.4.83, Norwegen, Mjosen 4894, Oslo Mus. F 12977” (SMNK: 11366); 1 undissected $ “ Cyclops

lacustris Sars, $ Kiefer 31 Aug. 71, Norwegen Mjosen” (SMNK: 9567); and 2 5? dissected on two

slides “ Cyclops lacustris Sars, §, C. Kiefer 31 Aug. 71, Norwegen Mjosen” (SMNK: 9568 (Abd+P5),

SMNK: 9569 (A1-P4).

Fig. 8. Cyclops lacustris G.O. Sars, 1863. A-B. §. A. Leg 1, frontal (ZMO: lectotype designated

herein). B. Leg 4, caudal - distal margin of the first and second exopodal segments bearing spinules on

the frontal surface only (ZMO: Sars Coll. 12977 $-2). C-F. S- C. Anal somite and caudal rami, ventral

(ZMO: Sars Coll. 12977 c?-l). D. Urosomites 1-4, ventral (ZMO: Sars Coll. 12977 c?-l). E. Antennal

coxobasis, caudal (ZMO: Sars Coll. 12976 F. Maxillulary palp (ZMO: Sars Coll. 12976 c?-l).

Scale bars =100 pm.

HOLYNSKAM. & DIMANTE-DEIMANTOVICA I., Redescription of Cyclops bohater and C. lacustris

Other material examined

NORWAY (southern): 1 §, Oymarksjoen, 59°23' N, 11°39' E, 107 m a.s.l., surface area 14.3 km 2 , max.

depth 37.6 m, 18 Jul. 2012, leg. I. Dimante-Deimantovica & B. Walseng (MIZ 2/2016/16); 1 $, Store

Le, 59°17' N, 11°48' E, 102 m a.s.l., located on the border between Norway and Sweden, surface area

136.1 km 2 , of which 14.88 km 2 in Norway, max. depth 58.2 m, 2011, R2872, leg. I. Dimante-Deiman¬

tovica & B. Walseng (MIZ 2/2016/17); 1 $, Aremark, 59°15' N, 11°40' E, 105 m a.s.l., surface area

7.47km 2 ,max. depth39.5m, 18 Jul. 2012,leg. I. Dimante-Deimantovica&B. Walseng (MIZ 2/2016/18);

2 5 ?, 1 S, Aspern, 59°9' N, 11°43' E, 105 m a.s.l., surface area 6.76 km 2 , max. depth 46.5 m, Jul. 2011,

leg. I. Dimante-Deimantovica & B. Walseng (MIZ 2/2016/19-20 and MIZ 2/2016/21, respectively).

SWEDEN: 4 $ §, K. Lindberg collection Lund, jar nr. 109, S. 28 (ZMLU).

LATVIA: 1 $ dissected (MIZ 2/2016/22) and 1 ? in alcohol (IDD), 1 CV $ parts (A2, Mxl, PI) (MIZ

2/2016/23), 2 CIV $ $ parts (Al, A2, Mxl, PI) (MIZ 2/2016/24-25), 4 CIV parts (A2, Mxl, PI) (MIZ

2/2016/26-29), Lake Svente, 55°51' N, 26°22' E, southeastern part of Latvia, 137 m a.s.l., surface area

7.35 km 2 , max. depth 38 m, 30 Aug. 2007, leg. I. Deimantovica.

Description

Female

Body length 1375-1800 pm (the female here designated as lectotype was originally dissected by

F. Kiefer, and no data are available on the morphometric traits of the prosome and the total body

length). Cephalothorax length/width 1.1—1.3, cephalothorax width/genital double-somite width 2.6-2.8,

Fig. 9. A-B. Cyclops lacustris G.O. Sars, 1863, S (ZMO: Sars Coll. 12976 cJ-1). A. Leg 1, frontal. B. Leg

4, caudal. C. Cyclops vicinus Uljanin, 1875, maxilliped syncoxopodite, frontal (Lake 0stensjovann,

Norway - MIZ 2/2016/30). Scale bars = 50 pm.

European Journal of Taxonomy 212: 1-31 (2016)

prosome length/urosome length 1.4-1.7. Pediger 2 (Fig. 7A) without distinct posterolateral lobes, and

not wider than succeeding somites. Pediger 5 not protruding laterally (Fig. 7B vs. Fig. IB).

Genital double-somite (Fig. 7B) longer than its greatest width. Seminal receptacle (Fig. 7B) as typical in

genus, transverse ridge present next to copulatory pore. Posterior margin of anal somite with continuous

row of spinules, anal sinus without surface ornamentation, anal operculum weakly developed. Caudal

rami (Fig. 1C) length/width 4.3-7.5 (lectotype, 4.9), medial margin bearing hairs. Midline crest running

almost full length of ramus on dorsal surface. Few spinules present on lateral margin in anterior third,

indicating presumptive insertion site of anterolateral accessory seta (I). Spinules also present at insertion

of caudal setae II and III. Caudal seta II inserted at distance of 0.20-0.26 ramus length measured from

posterior end (lectotype, 0.25). Relative length of caudal setae VII, VI, V, IV, and III: 1.2-2.0 (lectotype,

1.5), 1.8-2.6 (lectotype, 2.3), 3.7-6.0 (lectotype, 5.3), 3.2-4.9 (lectotype, 4.0), 1.0. Seta VI (medialmost)

1.0-1.4 (lectotype, 1.35) times as long as caudal rami, seta V (longest) 0.63-1.0 (lectotype, 0.95) times

as long as urosome. Caudal setae homonomously setulose, setae IV and V with breaking plane.

Antennule reaching middle of pediger 3, segmentation and setation pattern as in C. bohater. Aesthetasc

on segment 12 (Fig. 7D) reaching middle to distal margin of segment 14; aesthetasc length 2.9-3.6%

of body length (n = 10; no data on lectotype). Aesthetasc on segment 16 reaching near insertion site of

medial seta of segment 17 (approx, middle of terminal segment); aesthetasc length 2.6-3.6% of body

length (n = 9; no data on lectotype). Spinules only present on ventral surface of segment 1. Last three

segments of ante nn ule with hyaline membrane.

Antenna composed of coxobasis and three-segmented endopodite, and bearing 3, 1, 9 and 7 setae,

respectively. Inner medial seta of coxobasis (Fig. 7E-F) with long setules, outer medial seta naked.

Exopodite seta reaching beyond enp3 and bearing setules conspicuously longer in proximal than in distal

section of seta. On frontal surface of coxobasis, spinules (4, 5) present in -4/10 of segment and group

of longer spinules appear more proximally next to lateral margin, few long spinules usually present in

oblique/longitudinal row near middle line in proximal third of segment (for similar pattern see Fig. 2A).

Caudal surface of coxobasis (Fig. 7E-F) with double row of long spinules near long proximal spinules

on lateral margin, long and thin spinules (5-7) in longitudinal row near lateral margin, and oblique field

of small spinules proximally to insertions of medial setae.

Labrum with distal teeth in arc, lateral lobe with small spinules. Distal hairs long and arranged in two

groups. Epistoma and vertical cleft naked. Segmentation and setation of mandible, maxillule, maxilla

and maxilliped as in C. bohater. No cuticular ornamentation present next and proximally to mandibular

palp. Maxillulary palp usually without spinules (Fig. 7G) (spinules absent in lectotype). Proximal seta

and at least one (sometimes two or all three) of lateral lobe setae of maxillulary palp with distinctly

long setules (Fig. 7G-H). Claw-like attenuation of basipodite of maxilla with spinules on both concave

(inner) and convex (outer) margin. Basipodite seta inserted in front of claw-like attenuation with

long setules on both concave and convex edge in proximal half, and tiny spinules on convex edge in

distal half (for similar pattern see Fig. 3 A). Maxilliped: first endopodal segment bearing long and thin

spinules on frontal surface; basipodite with spinules arranged in two groups on caudal surface and

long and thin spinules on frontal surface; syncoxopodite frontally adorned with small spinules midway

between median and distalmost setae, tiny spinules sometimes present (Sweden, $ -2) near finger-like

membraneous element (for similar pattern see Fig. 3B).

P1-P4 setation formula same as in C. bohater (Table 2). Medial spine of PI basipodite (Fig. 8A): setules

distinctly longer near base of seta and becoming abruptly shorter towards distal end, or length of setules

almost gradually decreasing (lectotype). Medial spine reaching beyond distal margin of enp2 up to

middle of enp3. Long spinules arranged in arc on frontal surface of PI basipodite between insertions of

HOLYNSKAM. & DIMANTE-DEIMANTOVICA I., Redescription of Cyclops bohater and C. lacustris

exo- and endopodite. Intercoxal sclerites naked on frontal and caudal surfaces in P1-P3, and sparsely

pilose on caudal surface in P4. Obtuse protuberances of P4 intercoxal sclerite low, slightly extending

beyond distal margin of segment (Fig. 8B). Medial expansion of basipodite usually pilose in P1-P3 and

naked inP4; basipodite sometimes naked inP2-P4 (Sars Coll. 12977 $-1) or pilose in all swimming legs

(Sweden, $-2). Caudal surface ornamentation ofP4 coxopodite (Fig. 8B) composed of several groups of

spinules (groups are coded according to Einsle 1996a): A-with 6-9 spinules in medial section and 5-10

spinules in lateral section; B - few spinules (5), group sometimes absent; C - 5-7 spinules, sometimes

accompanied with 2-4 spinules directly above former row; D - usually single spinule, sometimes group

of spinules (7 + 1); E - 5-8 spinules. P4 coxopodal seta conspicuously longer (1.6-2.lx; lectotype,

1.6x) than height of medial expansion of basipodite. On caudal surface, distal margin of first and second

exopodal segment of P4 naked or bearing spinules. P4 enp3 2.8-3.8 (lectotype, 3.0) times as long as

wide; of apical spines medial one 2.2-3.8 (lectotype, 3.4) times as long as lateral and 0.8-1.1 (lectotype,

0.95) times as long as segment.

P5 (Fig. 7B) segmentation and setation as typical in genus. Distal segment 1.9-2.3 times as long as

wide. Medial spine inserted near half-length of segment (0.47-0.64 segment length measured from

proximal margin; 0.60 segment length in lectotype), medial spine 0.33-0.62 (lectotype, 0.37) times as

long as distal segment. Lateral seta on proximal segment relatively long, 0.70-0.90 (lectotype, 0.70)

times as long as apical seta and 2.5-4.0 (lectotype, 2.5) times as long as distal segment. Apical seta

3.3-4.6 (lectotype, 3.6) times as long as distal segment. Spinules present at insertion of median spine

and apical seta, but absent at insertion of lateral seta.

Fig. 10. Original drawings of Cyclops lacustris , made by G.O. Sars (Archives of the National Library

of Norway, archive items - Ms.Fol. 1109: 333; Ms.Fol. 1109: 421). A. Habitus, dorsal. B. Antenna,

caudal - note the long setules on the inner medial seta of the coxobasis. C. Leg 5, third endopodal

segment of leg 4, and caudal rami with setae. All drawings show specimens from Lake Mjosa, Norway.

European Journal of Taxonomy 212: 1-31 (2016)

Male

Body length 1135-1325 jli m , cephalothorax length/width 1.3-1.4, prosome length/urosome length ca

1.6. Pediger 5 laterally not protruding. Caudal rami (Fig. 8C) without dorsal crest, 3.5-5.1 times as

long as wide, pilose on medial margin. Short transverse row of spinules present in anterior third at

presumptive insertion site of ancestral anterolateral accessory seta (I). Spinules also present at insertion

of caudal seta II and III. Caudal seta II inserted at distance of 0.26-0.30 ramus length measured from

posterior end. Relative length of caudal setae VII, VI, V, IV, and III: 1.8-2.5, 2.3-2.8, 4.1-5.4, 3.3-4.4,

1.0. Seta VI 1.5-1.7 times as long as caudal rami, seta V 0.81-0.98 times as long as urosome.

Antennule 17-segmented, but two terminal segments (ancestral segment XXV and compound segment

XXVI-XXVIII) fused on ventral surface. Armature formula as in C. bohater. Antenna segmentation and

setation as in female, except for enp2, which has 8 setae. Spinule ornamentation of antennal coxobasis

(Fig. 8E) similar to that in female, inner medial seta bearing long setules, outer seta naked. Mouthparts

similar to those in female. Lateral lobe setae of maxillulary palp sometimes lacking long setules (Fig.

8F). Maxilliped syncoxopodite bearing small spinules midway between median and distalmost setae

(verified in male from Lake Aspern, Norway).

P1-P4 segmentation and setation as in C. bohater (Table 2.). Medial spine of PI basipodite usually

with homonomous setulation (Fig. 9A) and exceeding distal margin of enp2; distinctly long setules

sometimes present near base of seta (Lake Aspern, Norway). Long spinules arranged in arc on frontal

surface of PI basipodite between insertions of exo- and endopodite. Lateral spine of P2 expl more

slender than other spines of exopodite, bearing long setules. Intercoxal sclerites naked in P1-P3 and

sparsely pilose on caudal surface in P4. Caudal surface ornamentation of P4 coxopodite composed of

groups A, C and E; group B sometimes present (Lake Aspern, Norway) (Fig. 9B cf. Fig. 8B). Coxopodal

seta ca 1.5x longer than height of medial expansion of basipodite. Medial expansion of basipodite

apically pilose in P1-P3, naked (paralectotype - Sars Coll. 12976) or pilose (paralectotype -Sars Coll.

12977; Lake Aspern, Norway) in P4. P4 enp3 2.7-3.3 times as long as wide. Of apical spines medial one

2.5-3.7 times as long as lateral spine, as long or slightly (~l.lx) longer than segment.

P5 (Fig. 8D) distal segment 1.6-1.9 times as long as wide, medial spine inserted near half-length of

segment (0.5-0.6 segment length measured from proximal margin), medial spine 0.41-0.50 times as

long as distal segment. Lateral seta on P5 proximal segment 0.60-0.74 times as long as apical seta and

3.0-4.2 times as long as distal segment. Apical seta 5.0-6.0 times as long as distal segment. Spinules

present at insertion of median spine and apical seta, but absent at insertion of lateral seta.

P6 (Fig. 8D) composed of three elements; median and lateral seta 1.1-1.4 times and 3.0-4.3 times as

long as median spine, respectively. P6 flap naked.

Morphology of the species diagnostic characters in the late copepodid instars (Lake Svente, Latvia)

Antennule segmentation and setation in CIV instar as in C. bohater. Inner medial seta of antennal

coxobasis with conspicuously long setules, outer medial seta naked in CIV and CV; second endopodal

segment with 7 and 8 setae in CIV and CV (male), respectively. Long setules present on proximal seta

of maxillulary palp and at least one seta of lateral lobe of maxillulary palp both in CIV female and C V

male. Setulation of medial spine of PI basipodite homonomous or heteronomous (few longer setules

present proximally) in CIV, and heteronomous in CV male (verified in single specimen).

Remarks

The species was originally described from two large lakes (Mjosa (or Mjosen) and Tyrifjord) in Norway;

a holotype was not designated by the author (Sars 1863). The Zoological Museum in Oslo holds the

syntypes of C. lacustris collected by Sars from Lake Mjosa, comprising alcohol material and few

HOLYNSKAM. & DIMANTE-DEIMANTOVICA I,, Redescription of Cyclops bohater and C. lacustris

microscope slide preparata made by F. Kiefer; the fate of the original material from Tyrifjord is unknown

to us. Other parts of the Sars legacy, such as notes, manuscript drafts and unpublished drawings, are

kept in the archives of the National Library of Norway. The first description of the species was part of a

two-volumed hand-written manuscript (archive item Ms. Fol. 1109: 613) entitled “Om de i Christiania’s

Omegn foreko mm ende Ferskvandskrebsdyr. I og II” (On the freshwater crustaceans occurring in the

vicinity of Christiania) and devoted to the Cladocera, Copepoda and Ostracoda; this work as a whole has

never been published. In the archives we did not manage to identify the drawings of C. lacustris which

Sars might have added to the above mentioned manuscript; in any case, the published description of the

species (Sars 1863) did not contain illustrations. Nevertheless, the published diagnosis, the illustrations

that later appeared in the monograph of the Crustacea of Norway (Sars 1914) and some fine pencil

drawings of C. lacustris from Lake Mjosa (Fig. 10A-C; date of origin of these drawings is unknown),

deposited in the archives, allow us to identify the species with great certainty.

Kiefer (1978) provided drawings of the female habitus and the third endopodal segment of leg 4 of the

“lectotypus” of C. lacustris , which most likely show one of the two females that he selected from among

the syntypes as “lectotype” in the early seventies. These two females (one dissected on two slides, and

one undissected specimen) plus one male, all of which were labelled by Kiefer as “lectotypus”, are

deposited in the Zoological Museum in Oslo. As the lectotype designation by Kiefer is invalid (only one

specimen can be choosen as the lectotype, ICZN art. 74.1), we designated the lectotype of C. lacustris

herein, which is the dissected female labelled by Kiefer as lectotypus “B” (containing Abd+P5) and “C”

(containing A1-P4).

Ecology

Cyclops lacustris is a true limnetic species, mostly known from comparatively large and deep

oligotrophic to slightly eutrophic lakes. Adult individuals are generally found in the epilimnion, close

to the surface (Sars 1914); however, the species has also been recorded down to 20 m depth and deeper

(Rylov 1948). In Lake Mjosa a significant part of the population has been found in deep waters. The

vertical distribution depends on the season and developmental stage (Huitfeldt-Kaas 1946; Jarl Eivind

Lovik, unpublished data).

In Norway C. lacustris often co-occurs with typical glacial relict species (Spikkeland et al. 2012).

In 2006, when C. lacustris was evaluated for the Norwegian Red List database, it was only known

from Lake Mjosa and got included in the Red List as a critically endangered species, threatened by

eutrofication (incl. oxygen decrease) and water temperature increase due to the global warming. Later,

by applying new micro-characters in species identification, C. lacustris was discovered in several other

lakes in SE Norway. The new finds indicated that the species had a wider distribution in the country than

was thought before. At the moment the Red List status of C. lacustris in Norway has been changed from

endangered to least concern (Bjorn Walseng, personal communication). The example of C. lacustris

illustrates how alpha taxonomy may contribute to a better understanding of the actual geographic

distribution of and environmental threats to a species.

Distribution

Verified occurrences from Northern Europe (Norway, Sweden, Latvia) (Table 5). For an overview of the

literature data on the geographic distribution of C. lacustris , see the Discussion.

European Journal of Taxonomy 212: 1-31 (2016)

Table 5. Overview of the published records of Cyclops lacustris G.O. Sars, 1863, including the forms

C. 1. suecicus Lindberg, 1957 and C. 1. finlandicus Lindberg, 1957. For comments on the data see

“Discussion”. * = there are no recent records of the species; ** = records that need further verification;

n.d. = no data; E = East; N = North; S = South; W = West.

Country

Region

Habitat

Altitude (m)

References

Norway

SE: L. Mjosa, L. Tyrifjord *,

L. Store Le,

L. Gymarksjoen, L.

Rodenessjoen, E. Aspern,

L. Femsjo, L. Skulerudsjoen,

L. Romsjoen,

E. Aremark (1)

lakes

-70-140

Sars 1863, 1914; Spikkeland

et al. 2012; (1) (this paper).

Sweden

South (Gotaland) and Central

(Svealand),

- inch the largest lakes

Vanern, Vattern, Malaren

lakes

-15-270

Lilljeborg 1901; Lindberg

1957

Finland

S: L. Lohjanjarvi

lake

Lindberg 1957

lakes from both the southern

and

northern (Finnish Lapland)

regions

lakes

Silfverberg 1999

Latvia

E: L. Svente L. Ilzas-

Geranimova,

L. Raznas,

E. Baltas, L. Kustaru

lakes

130-170

Line 1966; Deimantovica 2010

Estonia

n.d.

lakes.

n.d.

Monchenko 1974

Russia

NW: L. Onega, L. Ladoga

Several lakes from former

Karelo-Finnish SSR

lakes

-4-130

Rylov 1948,

SW: Volgograd reservoir,

Veselovsky reservoir

n.d.

Monchenko 1974

S Siberia: Lower Multinskoe

Lake in Altai

lake

1710

Rylov 1948

W Siberia: Lower Ob

river

n.d.

Yukhneva 1970

NE Siberia: Kolyma

river

n.d.

Monchenko 1974

Belarus**

S: Pripyat river basin

n.d.

Monchenko 1974

Poland**

NE: L. Kolje, L. Perty

(Suwalszczyzna)

lakes

-150

Karabin & Ejsmont-Karabin

1993

Germany* *

N: E. Plon (Schleswig-

Holstein)

lake

Herbst 1951

France**

n.d.

Boxshall & Defaye 2013

Spain**

n.d.

Boxshall & Defaye 2013

Ukraine**

SW: Dniester,

river

Monchenko 1974

S: Lower Dnieper valley

river, liman

—

Monchenko 1974

Georgia* *

Abkhazia

lake

Derevenskaya & Mingazova

2015

Turkey* *

n.d.

Ustaoglu 2004

Uzbekistan **

E: Kattakurgan (Zeravshan

River valley)

reservoir

503-543

Monchenko 1974

Kazakhstan * *

N: Kokshetau district

lakes

n.d.

Monchenko 1974

Kirgyzstan * *

N: Chu River valley

n.d.

Monchenko 1974

Orto Tokoy

reservoir

n.d.

Monchenko 1974

“Caspian Sea basin” **

n.d.

Monchenko 2003

HOLYNSKAM. & DIMANTE-DEIMANTOVICA I., Redescription of Cyclops bohater and C. lacustris

Identification key to the Cyclops species of Fenno-Scandinavia:

Character states are diagnostic both in the adult female and male (§, (?), or in the adult female (?) only.

1. PI: spinules absent or tiny on frontal surface of basipodite near distal margin between insertions

of exo- and endopodite (§,(?) .2

- PI: arc (or group) of large spinules present on frontal surface of basipodite near distal margin

between insertions of exo- and endopodite ($, (?) (Fig. 3C, arrowed).5

2. Pediger 4 with wing-like posterolateral protrusions, that is pediger 4 wider than pediger 3 (width

measured as distance between posterolateral angles of the somite) (§). Intercoxal sclerites of PI¬

PS (and usually also P4) naked on caudal surface (§). Mandibular coxopodite with group of large

spinules near palp on frontal surface ($, (?) (site is arrowed in Fig. 2D).3

- Pediger 4 without wing-like posterolateral protrusions, that is pediger 4 narrower than pediger 3

(width measured as distance between posterolateral angles of the somite) (§). Intercoxal sclerite of

P3 (sometimes also P2) with hairs on caudal surface, P4 intercoxal sclerite caudally naked or pilose

(5). Mandibular coxopodite without spinules near palp (Fig. 2D) (§, (?).4

3. Antennal coxobasis: spinules small or sometimes absent on frontal surface at mediodistal angle

of segment (site is shown by arrow in Fig. 2A) (§, (?); inner medial seta with long setules (for a

similar setulation pattern see Fig.7E-F) (§, (?). Frontal surface of maxilliped syncoxopodite with

a distal (midway between insertions of median and distal seta) and proximal (midway between

insertions of median and proximal seta) group of long spinules, and smaller spinules arranged in

nearly ellipse line laterally to proximal group of long spinules; scattered tiny spinules sometimes

also present near lateral margin (§, (?) (Fig. 9C) . C. vicinus Uljanin, 1875

- Antennal coxobasis: spinules large on frontal surface at mediodistal angle of segment ($, (?);

setules absent or short on inner medial seta ($, (?). Frontal surface of maxilliped syncoxopodite

with two groups of long spinules only: distal group (midway between insertions of median and

distal seta) and proximal group (midway between insertions of median and proximal seta) ($.,(?)

. C kikuchii Smirnov, 1932

4. Antennule usually 17-segmented (18,17,16, and 14-segment states were found in C.furcifer caspicus

Findberg, 1942 from a saline marsh in Golestan Prov., Iran) (5). Anterolateral (II) caudal seta

inserted at distance of 0.18-0.23 ramus length measured from posterior end (§). Maxillulary palp:

long setules present almost at full length of proximal seta and one (proximalmost) seta of lateral lobe

(5, c?). P4 coxal seta slender: seta thickness and setulation similar to those in corresponding setae

of P1-P3 ($) . C. furcifer Claus, 1857

- Antennule 14-segmented (§). Anterolateral (II) caudal seta inserted at distance of 0.24-0.29 ramus

length measured from posterior end (§). Maxillulary palp: proximal seta and lobe setae without

long setules (§, (?). P4 coxal seta usually swollen at its base (seta thicker than corresponding setae

in P1-P3), and setules of seta short and robust ($) . C. insignis Claus, 1857

5. Maxillulary palp: proximal seta bearing long setules ($, (?) (shown by arrow at bottom inFig. 2E)... 6

- Maxillulary palp: proximal seta without long setules (§,(?).10

6. Maxillulary palp: at least one of the lateral lobe setae also bearing long setules (§) (Fig. 2E).7

- Maxillulary palp: lateral lobe setae lacking long setules ($).9

7. Antennal coxobasis: inner medial seta with long setules (§, (?) (Fig. 7E-F). Dorsal (VII) caudal

seta 1.2-2.1 times as long as posterolateral (III) caudal seta (§). C. lacustris G.O. Sars, 1863

European Journal of Taxonomy 212: 1-31 (2016)

- Antennal coxobasis: inner medial seta naked or bearing short setules ($-(?). Dorsal (VII) caudal seta

shorter or just slightly longer (max. 1.2x) than posterolateral (III) caudal seta (§).8

8. PI basipodite: medial spine proximally bearing long setules/spinules ($, S) (Fig. 3C, arrowed).

Terminal accessory (VI) caudal seta longer than caudal ramus (§). Dorsal caudal seta as long

or slightly longer than posterolateral (III) caudal seta ($). Caudal surface ornamentation

of P4 coxopodite: group “E” present (Fig. 3D) (§) .C. bohater Kozminski, 1933

- PI basipodite: medial spine with tiny spinules only (§, <$). Terminal accessory (VI) caudal seta

shorter than caudal ramus (§). Dorsal caudal seta usually shorter than posterolateral (III) caudal seta

(5). Caudal surface ornamentation of P4 coxopodite: group “E” absent or present (cf. Fig. 3D) (5)

. C. heberti Einsle, 1996

(Occurrence of the species is expected in Fenno-Scandinavia.)

9. Maxillulary palp with tranverse row or large spinules near base of palp and field of tiny spinules more

apically (§, S) .C. divergens Lindberg, 1936

(Synonyms: C. abyssorum divulsus Lindberg, 1956 and C. singularis Einsle, 1996)

- Maxillulary palp naked or if spinules present they are tiny and scattered ($, S) .

. C. abyssorum G.O. Sars, 1863

10. Pediger 4 with wing-like posterolateral protrusions, that is pediger 4 wider than pediger 3 (width

measured as distance between posterolateral angles of the somite) ($). Anterolateral (II) caudal

seta inserted at distance of > l A ramus length, measured from posterior end ($). P5, proximal

(first) segment: spinules absent at insertion of lateral seta (§, $). Male antennule: first segment

with single (distal) aesthetasc .C. scutifer G.O. Sars, 1863

- Pediger 4 with or without wing-like posterolateral protrusions, that is pediger 4 wider or narrower

than pediger 3 (width measured as distance between posterolateral angles of the somite) (§).

Anterolateral (II) caudal seta inserted at distance of < l A ramus length, measured from posterior end

(5). P5, proximal (first) segment: spinules usually present at insertion of lateral seta ($, <$)■ Male

antennule: first segment with three aesthetascs .11

11. Spine formula on terminal exopodal segments of P1-P4, 2-3-3-3 (§)... C. kolensis Lilljeborg, 1901

- Spine formula on terminal exopodal segments of P1-P4, 3-4-3-3 ($).12

12. P4 coxopodite seta reaching distinctly beyond distalmost point of medial expansion of P4

basipodite ($, c?). Intercoxal sclerites of P1-P3 naked, P4 intercoxal sclerite caudally pilose

($). P5, distal (second) segment: apical seta long, 2.7-4.2 times as long as segment (§). Caudal

surface ornamentation of P4 coxopodite: groups “FTand “E” usually absent (for coding of

the spinule groups see Fig. 3D) (§, S) .C. strenuus Fischer, 1851

- P4 coxopodite seta not reaching beyond distalmost point of medial expansion of P4 basipodite

(5, c?). Intercoxal sclerites of P3-P4 (sometimes also P2) with hairs on caudal surface (§). P5,

distal (second) segment: apical seta short, 1.6-2.3 times as long as segment ($). Caudal surface

ornamentation of P4 coxopodite: groups “FT and “E” present (for coding of the spinule groups see

Fig. 3D) ($,(?) . C. sibiricus Lindberg, 1949

(This species has often been confounded with C. strenuus. Cyclops sibiricus is distributed in Siberia

and arctic North America; its wester nm ost occurrence is so far known from the Yamal Peninsula

(Lindberg 1957); occurrence in arctic Fenno-Scandinavia is possible.)

Discussion

In the genus only three species, C. bohater , C. lacustris and C. heberti Einsle, 1996, share the characteristic

setulation of the maxillulary palp setae, namely, long setules are present in the proximal half of the

HOLYNSKAM. & DIMANTE-DEIMANTOVICA I., Redescription of Cyclops bohater and C. lacustris

proximal seta and at least one seta of the lateral lobe of the palp (Figs 2E, 5A, 7G—H). This feature

is present not only in the adult female, but it also appears in the adult male (though with intraspecific

variation in C. lacustris ) and copepodid instars (data on CIV and CV) in these species. Both the adult

(female and male) and copepodid instars (data on CIV and C V) of Cyclops lacustris can be distinguished

from C. bohater and C. heberti by the conspicuously long setules on the inner medial seta of the antennal

coxobasis (Figs 7E-F, 8E vs. Figs 2A-B, 4D-E). Cyclops heberti differs from C. bohater in the lack of

long setules on the medial spine of the PI basipodite (copepodid and adult instars, both sexes), and the

relatively short caudal seta VI (seta Vl/caudal ramus length <1) in the adult female.

The terra typica of C. bohater is located in the coldest part of Poland (Kupryjanowicz 2007), mirrored in

the presence of boreal elements in the region; northerly distributed Cyclops species, such as C. scutifer ,

C. abyssorum (boreo-alpine/montane) and C. kolensis occur in Fake Wigry and its neighbourhood.

The geographic occurrence data (Table 4) seem to indicate a lowland in the north and montane in the

south pattern, in a region with humid continental climate with warm su mm er (warmest month average

below 22 °C but above 10 °C) and cold winter (coldest month average below -3 °C) (Peel et al. 2007).

Geographic range limits of the species are poorly understood. Einsle (1996a) wrote that C. bohater

occurred in Asia, yet except for the record (without information on the locality) from Turkey (Ustaoglu

2004), we failed to find any publication describing Asian occurrence. The presence of C. bohater in

Italy is questioned by Pesce (“Checklist of the Italian Copepods” http://www.luciopesce.net/copepods/ ).

Western and Southern Europe, and Western Asia, with hot summer and/or mild winter, are home to

another species, C. divergens , which by its large body size and the appearance of posterolateral lobes

on pediger 2 might be confounded with C. bohater (e.g., in the Ponor Mts, Bulgaria; Pandourski 1997).

Also the “ tatricus ” ecotype of C. abyssorum (as to the taxonomical status of the “ tatricus ” form, we

followed Einsle 1969, 1996a), characterized by the distinct posterolateral lobes on pediger 2, may

approach the lower bound of the body length in C. bohater , as it was observed in a few high mountain

lakes of the Pyrenees (Dussart 1979). Though the large body and the posterolateral lobes on pediger 2

still remain part of the character set that is diagnostic to C. bohater , other characters are also needed

to unambiguously identify the species. Cyclops bohater , C. abyssorum and C. divergens have identical

setulation on the proximal seta of the maxillulary palp (i.e. long setules are present in the proximal half

of the seta); however, in C. bohater long setules are also present on one or more setae of the lateral lobe

of the maxillulary palp (Figs 2E, 5 A), while the lateral lobe setae lack long setules in C. abyssorum and

C. divergens. In C. abyssorum setules are sometimes (rarefy) present on the proximalmost seta of the

lateral lobe (e.g., in 2 $ §, SMNK: 6306 and 8467, collected in Lake Constance and labelled by F. Kiefer

as “C. abyssorum bodanus ”), but they are much shorter (-A of the length of the setules on the proximal

seta of the maxillulary palp) than those in C. bohater (~ 3 A of the length of the setules on the proximal

seta of the maxillulary palp).

Relative length of the terminal accessory (VI) caudal seta in comparison to the caudal ramus length can

also be a helpful taxonomic character. There is a large variation in this trait in C. abyssorum (0.82-1.4)

and C. divergens (0.73-1.1) (Holynska 2008), yet in all the specimens of C. bohater that we have seen

so far, caudal seta VI was longer than the caudal ramus. Therefore, we speculate that those Cyclops

forms in which this index is less than one (e.g., in Thonon in the French Alps - see table II in Dussart

1958) might be not conspecific with C. bohater.

In the male, a useful diagnostic character is the relative length of the median seta of P6, which is as long

as the medial spine of P6 in C. bohater , while it is distinctly longer than the medial spine in C. divergens

(1.4-2.3 times) and C. abyssorum (1.1-1.9 times).

The species range limits of C. lacustris (Table 5) remain poorly understood as well, but the recent

taxonomic literature suggests it being a predominantly northerly distributed species: Fenno-Scandinavia,

European Journal of Taxonomy 212: 1-31 (2016)

Latvia, Estonia, northwestern Russia (Lindberg 1957; Rylov 1948; Monchenlco 1974; Einsle 1996a;

Silfverberg 1999; Dussart & Defaye 2006; Deimantovica et al. 2011). Cyclops lacustris was also reported

from two smaller lakes (Kojle, 17 ha; and Perty, 20 ha) in a limnological study in NE Poland (Karabin

& Ejsmont-Karabin 1993), but the records have not been confirmed by later finds of the species. The

occurrence in northern Germany (Lake Plon) is uncertain as well. In a paper about the crustaceans of

the small bodies of water in Schleswig-Holstein, Herbst (1951: 491) wrote that C. lacustris (along with

C. abyssorum , C. kolensis and C. vicinus ) lived in the plankton of Lake Plon in the spring of 1949;

however, Herbst did not mention C. lacustris in his later studies on the zooplankton of Lake Plon

(Herbst 1953, 1955). Monchenlco (1974) overviewed the taxonomical and limnological literature from

the territory of the former Soviet Union (Table 5), mentioning the alleged occurrence of C. lacustris in

the western/southwestern regions of the USSR, Central Asia and Siberia, and supposed that the species

could have been transported to southern latitudes by rivers. Nonetheless, we think that all southern

records, also those from France, Spain (see Boxshall & Defaye 2013) and the Caspian Sea basin

(literature data cited in Monchenko 2003), need to be confirmed.

Based on morphometric differences Lindberg (1957) described two subspecies from Fenno-Scandinavia:

C. lacustris suecicus Lindberg, 1957 from the Lakes Malaren and Ylen in Sweden, and C. 1. finlandicus

Lindberg, 1957 from the Lake Lohjanjarvi in Finland. Lindberg also reported on the occurrence of

C. lacustris s. str. (among others from Lake Stora Le, from where we also have material) and transitional

forms (in Lake Vanern) between the “typical” form and C. 1. suecicus in Sweden. Table 6 shows the

variation of those morphometric characters which Lindberg used to distinguish the subspecies. We can

conclude that morphometric traits of the specimens here examined from Sweden (Lindberg Coll., there

is no accurate information on the locality) and Latvia are wit hin the range found in the material from

Norway. The single noticeable difference observed in the Latvian specimens is a slight decrease in the

length proportion of caudal setae VI and III. However, the relative lengths of caudal setae VI and III

compared to the body length in the Latvian females are at the lower bound of the range of caudal seta

VI and at the upper bound of the range of caudal seta III measured in the females from Norway. Both

the Malaren and Ylen populations of “C. lacustris suecicus ” differ from C. lacustris s. str. (Norway,

Sweden, Latvia - see Table 6) in the relatively long caudal seta III (see proportions “III/body 1” and

“VI/III”). The Ylen population also has other differences in the P5 morphology (longer medial spine on

the distal segment and shorter lateral seta on the proximal segment), while the Malaren specimens are

distinguished by their large body size. Cyclops lacustris finlandicus differs from the nominate subspecies

in the large body length, wider pediger 5 and the longer caudal seta III. The types of these two subspecies

were not designated in the original description (Lindberg 1957); therefore, examination of the topotype

material might elucidate the taxonomic position of C. lacustris suecicus and C. lacustris finlandicus. In

some older taxonomic works (e.g., Dussart 1969) the taxa originally described as C. strenuus var. laevis

Losito, 1902 (Central Italy) and C. rubens f. corsicana Lindberg, 1955 (Corsica) were also treated as

subspecies of C. lacustris. In the recent literature (Einsle 1975, 1996a; Dussart & Defaye 2006) both

taxa are referred to as subspecies of C. abyssorum. Both the “ laevis ” and “ corsicanus” subspecies need

revision to clear up their accurate taxonomic relationships to C. abyssorum and its close relatives (e.g.,

C. diverge ns).

Acknowledgements

The authors gratefully acknowledge the help of the curators of the Natural History Museum (London),

Staatlisches Museum fur Naturkunde (Karlsruhe), Lund Museum of Zoology, Zoological Museum

Oslo, and the Museum and Institute of Zoology (Warsaw). We are also grateful to the employees of the

National Library of Norway, who made the drawings from the Sars archive accessible to us, and to all

people of the Oslo University and Norwegian Institute for Nature Research Oslo (NINA), who collected

material for this study. Our study was partly based on the research done in the frame of a Synthesys

HOLYNSKAM. & DIMANTE-DEIMANTOVICA I., Redescription of Cyclops bohater and C. lacustris

go ^

p p

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t/3

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C/3

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< < <

o cr

p o

P CP

a- ^

5 s - O

-O

■o

to ov

t!>

-o

t/1

1—l

IP)

-O

' — -

s — /

*PD

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h—*

’

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-o

s — /

s —"

1— k

-P^

vo VO

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V. Ov

2 d 2 £ 2 2 'd d d <

-o

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-o

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CD* w

—1

CD »

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1—*

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1—*

-o

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Table 6. Variability of the morphometric traits in the female of C. lacustris G.O. Sars, 1863. The morphometric characters selected here are those

which showed no or very slight overlapping between C. lacustris lacustris (Sweden), C. lacustris suecicus Lindberg, 1957 and C. lacustrisfinlandicus

Lindberg, 1957 (see Lindberg 1957: 190-191). Number of the specimens examined is given in parentheses after the mean value. * = Data taken from

Lindberg 1957. C. lacustris suecicus was described from the lakes Malaren (Mai) and Ylen (Yle) in Sweden. C. lacustris finlandicus was described

from Lake Lohjanjarvi in Linland. n.d. = no data. Underlined values are those which are distinctly out of the range of the Norwegian populations.

European Journal of Taxonomy 212 : 1-31 ( 2016 )

grant (GB-TAF-308) provided to MH in 2005. We are much obliged to Prof. Grace A. Wyngaard (James

Madison University, Harrisonburg, Virginia USA), who generously sponsored MH’s visit to the Lund

Museum of Zoology in 2008. MH gratefully thanks IDD for sponsoring her research visit in NINA in

2013. Last but not least, we also thank the reviewers for their useful comments.

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Manuscript received: 8 December 2015

Manuscript accepted: 15 March 2016

Published on: 14 July 2016

Topic editor: Rudy Jocque

Desk editor: Kristiaan Hoedemakers

Printed versions of all papers are also deposited in the libraries of the institutes that are members of the

EJT consortium: Museum national d’Histoire naturelle, Paris, France; Botanic Garden Meise, Belgium;

Royal Museum for Central Africa, Tervuren, Belgium; Natural History Museum, London, United

Kingdom; Royal Belgian Institute of Natural Sciences, Brussels, Belgium; Natural History Museum of

Denmark, Copenhagen, Denmark; Naturalis Biodiversity Center, Leiden, the Netherlands.