European Journal of Taxonomy Ressource électronique

European Journal of Taxonomy 23: 1-19
http://dx.doi.org/10.5852/ejt.2012.23

ISSN 2118-9773
WWW. europeanj oumaloftaxonomy. eu
2012 • Sidorov D. & Palatov D.

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

Research article

um:lsid:zoobank.org:pub:FFD87F78-3590-4C2A-A256-BC6379739106

Taxonomy of the spring dwelling amphipod Synurella ambulans
(Crustacea: Crangonyctidae) in West Russia: with notes on its

distribution and ecology

Dmitry SIDOROV ' * & Dmitry PALATOV ^

^Department of Zoology, Institute of Biology and Soil Seienee FEB RAS, 100-let Vladivostoku Av.

159, Vladivostok 690022, Russia. E-mail: sidorov@biosoil.ru
um:lsid:zoobank.org:author:55C1264F-67C0-4C3F-8E6B-ECElE821457B
^Department of Hydrobiology, Moseow State University, Feninskie Gory 1/12, Moseow 119991,

Russia. E-mail: triops@yandex.ru

um:lsid:zoobank.org:author:25BBE8A0-81C4-4B42-859A-3A7BA3EE3C0E

* Corresponding author: E-mail: sidorov@biosoil.ru

Abstract. This study deals with taxonomie problems of the semi-subterranean erangonyetid amphipod
Synurella ambulans (F. Muller, 1846), well-known from various freshwater habitats in Europe. The
taxonomy of the speeies S. ambulans and the generie diagnosis for the genus Synurella are revised. A new
synonymy is proposed: Synurella ambulans (F. Muller, 1846) = Synurella ambulans meschtscherica
Borutzky, 1929, syn. nov. The affinity with the related groups, distribution and eeology of the speeies
are examined.

Key words. Amphipoda, Crangonyetidae, Synurella ambulans, stygophile, distribution, eeology,
springs.

Sidorov D. & Palatov D. 2012. Taxonomy of the spring dwelling amphipod Synurella ambulans (Crustacea:
Crangonyctidae) in West Russia: with notes on its distribution and ecology. European Journal of Taxonomy 23:
1-19. http://dx.doi.org/10.5852/eit.2012.23

Introduction

The amphipod genus Synurella Wrzesniowski, 1877 is stygophile, but laeking typieal stygomorphie
features sueh as anophthalmy, depigmentation and redueed feeundity (the females normally produee no
less than 11-20 small-sized eggs). Synurella oeeurs in semi-subterranean freshwaters and eoastal plain
braekish habitats throughout the Holaretie region (Karaman 1974a, 1990; Barnard & Barnard 1983).
To date, approximately 19 speeies of Synurella have been deseribed, ineluding 6 from Russia. The
geographieal reeords of Synurella in western Russia are eonfined to three deseribed speeies: S. donensis
Martynov, 1919 (springs of Rostov-on-Don vieinity), S. derzhavini Behning, 1928 (wells of Saratov
vieinity) and A meschtscherica (Borutzky, 1929) (springs and spring-runs of Mesehtsehera).

The goal of our study was to examine taxonomie boundaries between S. ambulans and S. meschtscherica.
Some authors previously had doubts about the validity of S. meschtscherica (see Karaman 1974a) and

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European Journal of Taxonomy 23: 1-19 ( 2012 )

assumed it was a junior synonym of S. ambulans, because of the lack of clear differences between
these taxa. However, it was thus far never formally synonymized. It should be noted, that initially
S. meschtscherica was reported by Borutzky (1927) under the name S. ambulans, who then later
described it as a subspecies of that species (Borutzky 1929). Birstein (1948) provisionally evaluated the
status of S. ambulans meschtscherica and suggested that it deserved species status. However, this was
done without sufficient argumentation.

The taxonomy of the genus is confusing. Some forms of the large species-complex S. ambulans are
considered distinct species by some researchers, whereas other workers only rank them as subspecies or
consider them as local forms. Moreover, poorly known nominal species such as S. donensis, S. ambulans
taurica Martynov, 1931 and S. philareti Birstein, 1948 may simply be aberrant forms of other, better-
known, species. Consequently, the number of species to be included in the genus is uncertain. The
species S. ambulans was described as Gammarus ambulans by Friedrich Muller (1846) from ditches at
Greifswald in Mecklenburg-Vorpommern (N Germany) (Muller 1846). It has been suggested previously
that the species is extremely polymorphic and widespread in Europe and Asia Minor (Ruffo 1974;
Karaman 1974a, 2003; Nesemann 1993; Musko 1994; Kontschan 2001; Pezzoli 2010). However, the
morphological variability of the species was interpreted too liberally by the former authors and without
good reasons. As a consequence of this incomplete taxonomic knowledge, erroneous conclusions about
species origin and distribution were formulated. Since the recent discovery of S. ambulans in the Black
Sea area (Ketelaars 2004), Ukraine (Alexandrov et al. 2007) and in the Belgian province of West-
Flanders (Boets et al. 2010), it was assigned to a group of Ponto-Caspian invaders, although there is no
sound evidence for this assumption.

Some authors have considered integrating the genus Synurella into the genus Stygobromus, owing to the
lack of clear morphological and geographical boundaries between these groups (Birstein 1948; Karaman
1974a, 1974b), or directly united them (Barnard & Barnard 1983). However, some researchers have
expressed the opposite view (Holsinger 1977; Bousfield 1977). In our opinion, significant morphological
differences between these genera are primarily observed in the general body morphology. Moreover, the
genus Synurella is not monophyletic according to Martynov (1931), who assigned all the Far Eastem-
Siberian and one Alaskan species to the subgenus Eosynurella. The latter group differs markedly
from the European taxa, except for Synurella dershavini Behning, 1928 and from the North American
Synurella, by the pear-shaped gnathopod 2 propodi and the structure of uropod 3 with a strongly reduced
terminal segment. It is possible that the biogeography of the genera Synurella and Stygobromus cannot
be explained by a simplified barrier-insulation approach proposed by several authors (Birstein 1948;
Karaman 1974a, 1974b), but should apparently be explained by isolation through the existence of
ancient seas, straits, and by evolutionary differences.

Synurella meschtscherica from the Meschtschera Eowland is now formally considered a junior synonym
of S. ambulans. In our study we examined samples from the topotype locality of S. meschtscherica. We
further include detailed distributional information based on morphological comparison of material from
scattered localities in West Russia.

Material and Methods

Taxonomic sampling

The specimens of S. meschtscherica were collected and studied from an extensive territory on the East
EuropeanPlain (Fig. 1), ranging from the Baltic Seabasin (Pskov administrative area) to the Meschtschera
Eowland in the east (Moscow, Ryazan, Vladimir areas) and in the south to the Central Russian Upland in
the Oka River basin (Kaluga area) as far as the upper part of the Dniepr River basin (Bryansk area). The

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SIDOROV D. & PALATOV D., Taxonomy of Synurella ambulans

comprehensive eolleetion of speeimens was earried out mostly by the seeond author using a eommon
hand net. The samples were preserved in 80% ethanol and are kept at the Institute of Biology and
Soil Seienee (Vladivostok). Preserved material of S. ambulans at the Museum of Naturkunde (Berlin),
eolleeted and determined by M.L. Zettler from Kassow near Rostoek (Meeklenburg-Vorpommem,
Germany), was also used in this study for eomparison of two speeies.

Morphology

All relevant morphologieal struetures were examined and measured. To measure the body length, more
preeisely the distanee along the dorsal side of the body from the base of the first antenna to the base of

30 to the East o f Greenwich

Fig. 1. Map indieating the geographie distribution of Synurella ambulans (F. Muller, 1846) in western
Russia. Legend: “empty” springs display undisturbed springs with a rieh erenophilous fauna without S.
ambulans. Literature data: Borutzky (1929), Chertoprud (2006a, 2006b). Souree: Blank Mapping Tools,
Moseow (2009).

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European Journal of Taxonomy 23: 1-19 ( 2012 )

the telson, the speeimen was held in a ventro-dorsal position. A Lomo MBS-9 stereomieroseope with
a sealed mierometer eyepieee was used to make this measurement and appendages were drawn using
a Carl Zeiss NU-2 eompound mieroseope equipped with a drawing deviee as modified by Gorodkov
(1961). The permanent preparations were made using polyvinyl laetophenol (PVL) and a methylene
blue staining solution. A lens adapter LSN-23D by Zarf Enterprises for Nikon CoolPix 995 was used for
digital photomierography of the lateral eephalie lobes and appendages.

The term “defining angle” of the gnathopod propodi refers to the angle formed at the end of the palm
and beginning of the posterior margin or the point at whieh the tip of the daetylus eloses on the propodus
(see Holsinger 1974). The nomenelature for setal patterns on segment 3 of the mandibular palp follows
the standard introdueed by Karaman (1970). The deseriptive terminology follows a elassifieation system
based on the homology eoneept proposed by Watling (1989). The following deseription was generated
from a DELTA database (Dallwitz 2005) for the erangonyetid genera and speeies of the world.

Abbreviations

BN = Bryansk area, Navlinsky.

FENU = Zoologieal Museum of the Far East National University, Vladivostok.
IBSS = Institute of Biology and Soil Seienee, Vladivostok.

KF = Kaluga area, Ferzikovsky.

MO = Moseow area, Orekhovo-Zuevo.

MSU = Zoologieal Museum of Moseow State University, Moseow.

PP = Pskov area, Pustoshkinsky.

RK = Ryazan area, Klepiki.

VP = Vladimir area, Petushinsky.

Results

Taxonomy

Order Amphipoda Latreille, 1818
Family Crangonyetidae Bousfield, 1973

Genus Synurella Wrzesniowski, 1877

Synurella Wrzesniowski, 1877: 403.

Gop/a/ra Wrzesniowski, 1879: 299.

Boruta WrzQsniowski, 1888: 44.

Eosynurella MartynoY, 1931: 531.

Diasynurella Bohning, 1940: 43.

Type species

Gammarus ambulans F. Muller, 1846 (= Synurella ambulans (F. Muller, 1846) designated by
Wrzesniowski (1877)).

Revised diagnosis (related to sub-family group 1 sensu Bousfield 1977: 302)

Closely allied with Stygobromus Cope, 1872, but with the following eharaeteristie features: head, lateral
eephalie lobe broadly rounded without inferior sinus (exeept Synurella osellai); antenna 2 of male with
paddle-shaped ealeeoli; gnathopod 1 propodi sub-quadrate; gnathopod 2 propodi with well-developed
posterior margins, propodi always larger than the same of gnathopod 1; eoxal plates 1-3 deep, mueh
longer than broad; eoxal plate 4 deep, with exeavation; urosomites partially or entirely fused; telson
apieal margin distinetly notehed or lobate; oostegites 2-5 large, ovoid.

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SIDOROV D. & PALATOV D., Taxonomy of Synurella ambulans

Remarks

In our opinion, the Crimean form Synurella ambulans taurica Martynov, 1931, with a slightly extended
basipodite of pereopod 7, is related to the southern speeies eomplex of S. intermedia and S. tenebrarum
rather than to the nominative speeies. However, owing to the poor deseription it is diffieult to reaeh a
definite eonelusion.

Synurella ambulans (F. Muller, 1846) {sensu stricto)

Figs 2-9

Gammarus ambulans F. Muller, 1846: 296, Taf 10, figs A-C (original deseription).

Synurella ambulans Stebbing, 1906: 369.

Synurella polonicaWxzQsmimski, 1877: 403.

Synurella ambulans meschtscherica Borutzky, 1929: 30, figs 1-17, syn. nov.

Synurella meschtscherica ^ixstQm, 1948: 70.

Stygobromus ambulans BdLmdLxd, 1983: 438.

Stygobromus meschtschericus^dimdLYd, 1983: 440.

Synurella ambulans - Sehafema 1922: 57, tab. 1 (10), tab. 2 (1-4), text-figs 26-29. — Borutzky 1927:
63. — Sehellenberg 1942: 85, Fig. 66.

Synurellapolonica - Stebbing 1906: 369. — Jaroeki & Krzysik 1924: 555.

Synurella ambulans meschtscherica - Straskraba 1962: 132.

Synurella meschtscherica -^dimmd 1958: 75. — Straskraba 1967: 208. —Karaman 1974a: 124.
Stygobromus meschtschericus - Starobogatov 1995: 192. — Chertoprud 2006a: 19; 2006b: 382.

Diagnosis

Medium-sized speeies with marked seeondary sexual dimorphism. Body pigmented. Gnathopod 2 larger
than gnathopod 1. Pereopod 6 longer than pereopod 7. Pereopod 7 basis without distinet posterior lobe.
Coxal gills on pereopods 2-7, gill 7 very small. Sternal gills arrangement as following: pereonite 2 (-2-),
pereonite 3 (-2-), pereonite 6 (1-1), pereonite 7 (1-1), pleonite 1 (1-1). Brood plates 2-5 (oostegites)
rather broad, with long marginal setae. Body length 3.5 - 6.0 mm (5), 3.0 - 4.5 mm (3).

A distinetive feature of this speeies is a well-marked broad yellowish spot (Fig. 2A) loeated on the dorsal
surfaee of the head between eyes. The spot is diseemible only in living animals.

Fig. 2A-C. Synurella ambulans (F. Muller, 1846). A. Yellow spot on the dorsal surfaee of the head of
live speeimens (front and left side), MO. B. S, 4.2 mm, FENU X34906/Cr-1406, KF. C. $, 5.5 mm,
FENU X34906/Cr-1406, BN, left side (preserved speeimens).

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European Journal of Taxonomy 23: 1-19 (2012)

Material examined

GERMANY. All specimens (3 $ 1 (?) completely dissected and mounted on a single slide per number:

[MSU Mb-1146] $ (oostegites developed, setose) 5.7 mm and f 4.2 mm, [FENU X34906/Cr-1406] $
(oostegites developed, setose) 5.5 mm and $ (oostegites developed, setose) 5.2 mm. Mecklenburg-
Vorpommern, Kassow (53°87’76.3”N 12°07’67.2”E), 21 May 1997, collected by M. Zettler.

RUSSIA. All specimens completely dissected and mounted on a single slide [FENU X34906/Cr-1406],
8 2 (?(?: $ (oostegites developed, setose) 6.0 mm, Pskov area, Pustoshkinsky, near Yezerische

Fake, Kholodny brook (56°24’10”N 29°08’33”E), 20 Aug. 2010, collected by D. Palatov; 2 $$
(oostegites developed, setose) 3.5 mm and 5.5 mm, Bryansk area, Navlinsky, Desna River basin, near
Partizanskoye, pond (52°45’77”N 34°22’72”E), 17 Sep. 2009, collected by D. Palatov; $ (oostegites
developed, non-setose) 5.5 mm and $ (oostegites developed, setose) 4.0 mm, Kaluga area, Ferzikovsky,
Oka River basin, spring (54°29’47”N 36°21’41”E), 02 Jul. 2007, collected by D. Palatov; $ (oostegites
developed, non-setose) 5.2 mm and f 4.5 mm, Moscow area, Orekhovo-Zuevo, ~ 3.5 km E of Voinovo,
Chernaya River (55°50’42”N 39°04’82”E), 02 May 2009, collected by D. Palatov; $ (oostegites

Fig. 3A-B. Synurella ambulans (F. Muller, 1846), (?, 4.5 mm, FENU X34906/Cr-1406, MO.
A. Gnathopod 1. B. Gnathopod 2. Scale bars 0.2 mm.

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SIDOROV D. & PALATOV D., Taxonomy of Synurella ambulans

Fig. 4A-L. Synurella ambulans (F. Muller, 1846), S, 4.5 mm, FENU X34906/Cr-1406, MO.
A. Antenna 1. B. Antenna 2. C. Maxilla 1. D. Maxilla 2. E. Lower lip. F. Left mandible. G. Right
mandible. H. Upper lip. I. Maxilliped. J. Maxilliped, inner plate. K. Maxilliped, outer plate.
L. Distal part of maxilliped palp, female, 6.0 mm, FENU X34906/Cr-1406, PR Seale bars 0.2 mm.

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European Journal of Taxonomy 23: 1-19 (2012)

developed, setose) 5.0 mm. Nerskaya River basin, near Podosinky, brook (55°34’18”N 38°49’12”E),
27 Aug. 2005, eolleeted by D. Palatov; 2 $ $ (oostegites developed, non-setose) 4.0 mm and 3.8 mm,
~ 2.5 km NE of Aneiferovo, “Aneiferovsky Spring” (55°33’85”N 38°48’17”E), 8 Jan. 2010, eolleeted
by D. Palatov; f 4.0 mm. Vladimir area, Petushinsky, Markovo, brook (55°52’11”N 39°17’15”E), 22
Apr. 2007, eolleeted by D. Palatov.

Additional material examined

All speeimens measured, partially disseeted and stored in different vials [IBSS 17/2SD], ea. 82
33 (?(?: 3 $ $, 4 (f(f, Vladimir area, Petushinsky, ~ 3 km SE of Usad, small floodplain lake (55°51 ’27”N
39°08’76”E), 02 May 2009, eolleeted by D. Palatov; 4 $ Gus-Khrustalny, near Shestimirovo, Buzha
River basin, brook (55°27’09”N40°13’68”E), 14May 1994, eolleeted by M. Chertoprud and D. Palatov;

Fig. 5A-E. Synurella ambulans (F. Muller, 1846), 4.5 mm, FENU X34906/Cr-1406, MO.

A. Pereopod 3. B. Pereopod 4. C. Pereopod 5. D. Pereopod 6. E. Pereopod 7. Seale bars 0.2 mm.

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SIDOROV D. & PALATOV D., Taxonomy of Synurella ambulans

4 Ryazan area, Klepiki,- l.SkmNW Shmeli,Yalma River basin, spring (55°12’93”N39°55’63”E),

02 Oet. 2006, eolleeted by M. Chertopmd and D. Palatov; 38 near Velikodvorye, Yalma River
basin, springs (55°12’46”N 39°59’12”E), 20 Oet. 2006, eolleeted by D. Palatov; ea. 50 Kaluga
area, Ferzikovsky, ~ 2 km E of Majakovsky, Oka River basin, spring (54°29’47”N 36°2E4r’E), 30
Apr. 2011, eolleeted by D. Palatov; 5 $ $, 4 S'S', Bryansk area, Navlinsky, near Dumeha, Dumeha River
basin, springs (52°49’35”N 34°10’48”E), 19 Sep. 2009, eolleeted by D. Palatov; 2 $$, Pskov area,
Pustoshkinsky, Velikaya River basin, ~ 2 km W of Vysotskoe, brook (56°26’68”N 29°22’06”E), 16
Aug. 2010, eolleeted by D. Palatov.

Type locality

Germany, Meeklenburg-Vorpommem, Greifswald (approx. 54°5’N, 13°23’E), ditehes (F. Muller, 1846).
Type material stored in the zoologieal eolleetion of the Greifswald University (Zettler 1998: 57).

Redescription

Male

Eength. 4.5 mm, FENU X34906/Cr-1406.

Habitus. (Fig. 2B) Not stygomorphie.

A

Fig. 6A-H. Synurella ambulans (F. Muller, 1846), S, 4.5 mm, FENU X34906/Cr-1406, MO.
A. Pleopod 1. B. Pleopod 2. C. Pleopod 3. D. Epimera 1-3. E. Uropod 1. F. Uropod 2. G. Uropod 3.
H. Telson. Seale bars 0.2 mm.

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European Journal of Taxonomy 23: 1-19 (2012)

Body. Slender with elongate appendages, eolor yellowish.

Head. Eyes (Figs 2B; 9) vestigial, blaek; yellow spot (Fig. 2A) loeated on the dorsal surfaee of the
head between eyes eharaeteristie for living speeimens. Antenna 1 (Fig. 4A) 55% length of body, 30%
longer than antenna 2; peduneular segments 1-3 in length ratio 1:0.8:0.6; primary flagellum with 13
segments; aesthetases present. Antenna 2 (Fig. 4B), peduneular segments 4 and 5 in lengths ratio 1:1;
flagellum with 5 segments; ealeeoli present. Feft mandible (Fig. 4F) ineisor 5-dentate; laeinia mobilis
5-dentate; setal row with 3 serrate setae. Right mandible (Fig. 4G) ineisor 5-dentate; laeinia mobilis
trifureate. Molar proeess (Fig. 4F, G) triturative, with aeeessory seta. Palp mandible (Fig. 4G) segment
2 slightly longer than segment 3; segment 3 with 1 A-seta, 2 C-setae, 6 D-setae and 4 E-setae. Fower lip
(Fig. 4E), inner lobes present; mandibular proeess indistinet (broad). Maxilla 1 (Fig. 4C), inner plate
with 7 plumose setae; outer plate with 7 serrate setae; palp segment 2 about 2x longer than segment 1.
Maxilla 2 (Fig. 4D), inner plate with 6 plumose setae. Maxilliped (Fig. 41-K) inner plate with 3 strong
apieal setae; outer plate broad. Foregut lateralia with 8 strong peetinate setae.

Fig. 7A-B. Synurella ambulans (F. Miiller, 1846), $, 4.0 mm, FENU X34906/Cr-1406, KF.
A. Gnathopod 1. B. Gnathopod 2. Seale bars 0.2 mm.

10

SIDOROV D. & PALATOV D., Taxonomy of Synurella ambulans

Pereon. Gnathopod 1 (Fig. 3A), propodus palm beveled, defining angle distinet, palmar modified
setae at defining angle present, palm with eutting margin smooth, palm with 19 simple strong setae
in two rows; daetylus, inner margin smooth. Gnathopod 2 (Fig. 3B), propodus larger than gnathopod
1 propodus; palm distinetly beveled, defining angle distinet, palmar modified setae at defining angle
present, palm with eutting margin smooth, palm with 24 simple strong setae in two rows; daetylus,
inner margin smooth. Pereopod 6 longer than pereopod 7. Pereopods 5-7 (Fig. 5C-E) bases expanded,
posterior margins with serration. Pereopods 3-7 (Fig. 5A-E) daetyli elongated, about 40-50% length of
eorresponding propodi. Coxal gill 7 present. Paired median sternal gills on pereonite 2 and pereonite 3.
Single lateral sternal gills on pereonite 6, pereonite 7 and pleonite 1.

Peeon. Epimeron 1 (Fig. 6D), posteroventral eomer aeute or sub-aeute, ventral margin unarmed. Epimera
2-3 (Fig. 6D), posteroventral eomer aeute or sub-aeute, ventral margins armed. Pleopods 1-3 (Fig. 6A-
C), peduneular segments with 2 eoupling setae (retinaeulae). Uropod 1 (Fig. 6E), inner ramus 80% as
long as pedunele, distal peduneular proeess absent. Uropod 2 (Fig. 6F) about 65% as long as uropod 1,

Fig. 8A-J. Synurella ambulans (F. Muller, 1846), $, 4.0 mm, FENU X34906/Cr-1406, KF. A. Eateralia.
B. Antenna 2. C. Pereopod 5. D. Pereopod 6. E. Pereopod 7. F. Epimera 1-3. G. Uropod 1. H. Uropod
2.1. Uropod 3. J. Telson. Seale bars 0.2 mm.

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European Journal of Taxonomy 23: 1-19 (2012)

peduncle shorter than inner ramus; inner ramus longer than outer ramus. Uropod 3 (Fig. 6G) uniramous,
peduncle or/and lateral margin of ramus armed. Telson (Fig. 6H) not tapered distally, rather elongate,
1.8x longer than broad, about 10% longer than uropod 3, apical margin cleft on 1/3 of total length, with
6 strong notched setae on each lobe.

Dimorphism

Female

Length. 5.5 mm, FENU X34906/Cr-1406), sexually dimorphic characters.

Body. (Fig. 2C) Stout, appendages shortened. Antenna 1 45% longer than antenna 2. Antenna 2 (Fig. 8B)
flagellum with 6 segments; calceoli absent. Gnathopod 1 (Fig. 7A), propodus palm transverse or scarcely
sub-transverse with cutting margin acanthaceous, palm with 8 simple strong setae in two rows; dactylus,
inner margin with setae. Gnathopod 2 (Fig. 7B), propodus palm with cutting margin acanthaceous,
palm with 7 simple strong setae in two rows; dactylus, inner margin with setae. Pereopods 3-7
(Fig. 8C-E), dactyli about 45-50% length of corresponding propodi. Uropod 1 (Fig. 8G), inner ramus as
long as peduncle. Uropod 2 (Fig. 8H) about 60 % as long as uropod 1. Telson (Fig. 8J) somewhat tapered
distally, slightly elongate. Fix longer than broad, as long as uropod 3. Oostegites 2-5 large, ovoid with
long marginal setae.

Variability

Karaman (1974a) pointed out a significant variability in several morphological features for S. ambulans.
However, in our analysis of individuals from the different parts of its range in Russia and Germany, we
could not discover any significant variance in the shape of the lateral cephalic lobes, epimera, uropods,
telson or bases of pereopod 7 (Fig. 9). In adults we observed elongation of the pereopod 7 bases and the
presence of many robust setae on the lower edge of the epimera 2-3. Ommatidia were larger in young
animals but their number was smaller than the one in adults. We also noted a slight variation in the
length of the antenna 1 50-55% length of body and 40^5% longer than antenna 2, and a considerable
variation in the length of pereopods 3-7 dactyli (35-50% length to corresponding propodi). The number
of segments in the flagellum of antenna 1 equals 12-16. The specimens from Kholodny brook, near
Yezerische Take (Russia, Pskov area, Pustokshinsky) have a slightly different setation pattern of
maxilliped palp segments 3 and 4 (Fig. 4E), but are otherwise indistinguishable.

Remarks

Borutzky (1929: 32) adduced several distinctive characters which, in his opinion, were sufficient to
distinguish S. a. meschtscherica from S. a. ambulans: relative length of the both antennae, the stronger
armament of mouthparts (viz., presence of scopiform bundles of setules on palpi of mandibles and
maxilliped, presence of molar setae), armament of uropod 3 peduncle, the shape and armament of telson.
After analyzing Borutzky’s description, we are convinced that he has mixed details (p. 33) of males
and females without explanation: the cited characters of the antenna 2 and the gnathopods belong to
the female, while the telson characters are typical of the male. Moreover, Borutzky (loc. cit.) compared
his own “mixed” description to the incomplete description by Schafema (1922), who also depicted
the female’s telson without indication of gender. Borutzky (loc. cit.) evidently did not have Muller’s
(1846) original description at hand, in which the latter explains why he attributed subspecies status to
his specimens.

The comparison of the material of S. ambulans from Russia, previously identified as Synurella
meschtscherica, with that from Germany revealed no morphological differences between them.
Comparison of the variability of the original samples with species descriptions by Mtiller (1846),
Schafema (1922) and Bomtzky (1929) showed that both species are identical. We therefore consider

12

SIDOROV D. & PALATOV D., Taxonomy of Synurella ambulans

Synurella ambulans meschtscherica Bomtzky, 1929 a junior synonym of nominative S. ambulans (F.
Muller, 1846).

The taxonomie status and geographie distribution of previously deseribed forms of S. ambulans are in
need of a substantial revision. In our opinion the eomplex elassifieation of S. ambulans is eaused by: 1)
a poor first deseription of the speeies by F. Muller and 2) a relatively wide distribution of the genus in
Europe. It is possible that S. ambulans, ranging widely in Europe and Asia with signifieant variability
reported by some authors (see above), is aetually a series of several eryptie speeies.

A few diserepaneies were found in the eomparison with the original deseription. Bomtzky (loe. eit.)
reported the body length of individuals within the range of 6-12 mm for mature speeimens without
an indieation of the method of measurement. Our largest individual has a body length of 6.0 mm. We
have also studied the samples from Velikodvorskye springs of Ryazan area, previously also explored
by Bomtzky, where females up to 5.0 mm body length were found. Bomtzky (1929: 32) also found
a somewhat larger number of segments of the fiagellum of the antenna 1, 18-24 (males) and 16-22
(females), and reported on eyeless individuals whieh are absent in our eolleetions. However, the observed

Fig. 9. Variability of (A) lateral eephalie lobe of Synurella ambulans (F. Muller, 1846) from different
populations; (B) posterior margin of pereopod 7 basis; (C) epimera 2 and 3. (Digital Photomierography).

13

European Journal of Taxonomy 23: 1-19 (2012)

variability was not documented by this author, eyeless individuals were not deseribed and had not been
given a speeial status. The inaeeuraey of Borutzky’s deseription eonfirmed our doubts about the validity
of S. meschtscherica and eonvineed us that only one form of Synurella is present in the Mesehtsehera
Lowland.

Distribution

RUSSIA. Pskov area: Pustoshkinsky region. Vladimir area: Petushinsky and Gus-Khrustalny regions.
Moseow area: Orekhovo-Zuevo, Egoryevsk and Shatura regions (Chertoprud 2006a, 2006b). Ryazan
area, Klepiki (Borutzky 1927, 1929). Kaluga area: Ferzikovsky region. Bryansk area: Navlinsky region.

Although S. ambulans was found in extensive territories in West Russia, it was absent in a number of
different springs (see map) with a rieh erenophilous fauna. This mosaie distribution is apparently eaused
by environmental faetors.

Synurella ambulans has been reported from many eountries situated on the Great European Plain
ineluding Belgium (Boets et al. 2010), Germany (Heekes et al. 1996; Zettler 1998; Eggers & Martens
2001), Poland (Konopaeka & Soboeinska 1992), Eithuania (Arbaciauskas 2008) and Belarus (Giginyak
& Moroz 2000).

Ecology

Stygophile, predominantly oeeupying semi-subterranean habitats. Biotopes mostly ineluding wetlands,
bogs, wetland areas of streams with swampy shores nearly everywhere overgrown with Alnus (see
Borutzky 1929).

Synurella ambulans dwells in various springs, stagnant parts of the rivers and brooks eonneeted with the
ground outlets of subterranean waters, frequently assoeiated with the asellid isopod Asellus aquaticus
(Einnaeus, 1758). A eharaeteristie features of all mierohabitats are their stagnant or very slowly flowing
waters, not exeeeding 0.1 m/see; a water temperature generally ranging between 2.0 and 16.0 °C, a
low oxygen eoneentration of 3.0-9.0 mg/1, a pH between 5.0-8.0 and low mineralization not higher

than 197.5-353.1 mg/1 (onee 510.0 mg/1) (Nesemann et al. 1995; Giginyak & Moroz 2000; Chertoprud
2006a). Springs are often eovered with Lemna and Hydrocharis, or densely grown with Elodea and
Fontinalis; bottoms are eomposed of detritus, sand, mud, snags and leaf litter. Dendrocometes paradoxus
Stein, 1852 (Protozoa, Infusoria, Suetoria) is a eommon eetoparasite on the eoxal gills of S. ambulans
(see Taylor & Sanders 2001).

The rare flndings of S. ambulans in a number of a small floodplain lakes in the spring eould be explained
by the flood drift. However, most interesting is the aeeidental diseovery (by DP) of a mass eongestion
of S. ambulans on the shallows of a large lake in the Velikaya River basin (Pskov area) in winter. It is
possible that these erustaeeans ean survive adverse winter eonditions by “warming up” near oozing from
the bottom fontanels.

Discussion

Comments on biogeography of Synurella species complex

The biogeography of any group is elosely linked with its phylogenetie relationships (Holsinger 1986),
whieh in turn is related to a system of trustworthy diagnostie features. It is well-known that the taxonomy
of the family Crangonyetidae is based largely on the strueture of uropod 3, although the plastieity of this
eharaeter had already been diseussed (Bousfleld 1983) and a reeent moleeular phylogeny eonflrms this
(Hiwatari et al. 2011). The latest results of 18S phylogeny also revealed an aneient radiation of Synurella
with a paraphyletie relationship to the North Ameriean and European groups (Komobis et al. 2011).

14

SIDOROV D. & PALATOV D., Taxonomy of Synurella ambulans

Morphologically, the family Crangonyctidae is not homogeneous and ean be tentatively divided into two
groups of uneven size, but equivalent in terms of their “biogeographieal weighfThe first group ineludes
taxa with the “free eoxal plates \-A'' or eommonly diagnosed as shallow eoxae, when their width is greater
than, or equal to, the height (this group ineludes: Bactrurus, Siberian Stygobromus and the majority of
the North Ameriean Stygobromus). Two poorly deseribed Eurasian Stygobromus apscheronicus

(Derzhavin, 1945) and Stygobromus kazakhstanicus Kulkina, 1992 - oeeupy an intermediate position
but they might belong to this group, beeause of their shallow eoxal plates 3 and 4. The seeond large
group, however, eonsists of speeies with deep eoxal plates and is distinguished by a eoxal plate 4 with
exeavation. This group ineludes the North Ameriean and European Crangonyx, Synurella, Lyurella
hyrcana Derzhavin, 1939, Stygonyx courtneyi Bousfield & Holsinger, 1989, Amurocrangonyx arsenjevi
(Derzhavin, 1927) and Palaeogammarus. The same eharaeter was used previously by Bousfield (1977)
and Holsinger (1986) in a hypothesized phylogenetie relationship among Crangonyetidae. They aseribed
a plesiomorphie state for the deep eoxae and an apomorphie state for shallow or redueed eoxae in overall
size (see Holsinger 1986: 90). The front edge of the Crangonyetidae head is rather diverse in morphology
(Holsinger 1977) and ean be roundish (Fig. 9) or eharaeterized by the presenee of the inter-antennal lobe
and inferior sinus. However, this feature apparently may not be appropriate for phylogenetie analyses, as
we have observed non-uniform gradation from the roundish form (S. ambulans) with intermediate form
{S. derzhavini) to the expressed “sinusoidal form” (S. osellai) (see Sidorov et al. 2012). Meanwhile,
we have eoneluded that both features of general body morphology are most important for separation of
heterogeneous phyletie groups in future testing of biogeographie hypotheses.

Taxonomic boundaries and eco-geographic distribution of Synurella ambulans

Synurella ambulans has a large distribution in the lowlands of northern Europe, ranging from NW to NE
Europe and possibly reaehing the Blaek Sea area (Dedju 1967). Previously, from pieees of Baltie amber
dated as far baek as the Eoeene, several forms of Synurella have been deseribed as being elosely related
to the southern speeies S. intermedia, beeause of the somewhat extended pereopod 7 bases and the
non-aeute posteroventral epimeral eomers (see Coleman 2004: 3). Perhaps the northern distribution of
S. ambulans may be a relatively reeent phenomenon, after the retreat of the glaeiers in the early Holoeene.
However, we do not know the southern and western boundary of the distribution of this speeies. The
findings of S. ambulans in Southern Europe and Asia Minor indieate that its range is diseontinuous.
This assumption is questionable and requires additional verifieation. In our view, all referenees to
S. ambulans in Asia Minor (Ruffo 1974; Karaman 2003; Ustaoglu et al. 2004) as well as in Southern
Europe (Bonaeina et al. 1992; Stoeh & Dolee 1994; Pezzoli 2010) are ambiguous and should be re-
eompared earefully with S. ambulans from other areas.

Synurella ambulans is absent from a number of intaet springs, unaffeeted by anthropogenie stress
(Fig. 1). These springs were almost always rieh in erenophilous fauna. As noted by Giginyak & Moroz
(2000), Synurella ambulans inhabits springs with water elose to melt water in physieal and ehemieal
properties. The natural toleranee of the studied amphipod speeimens of S. ambulans was limited to low
eoneentration of oxygen and mineralization (see Giginyak & Moroz 2000: 82). Previously, a similar
relationship between life in springs and low water hardness of 1.2-1.6° dGh (= 214.1-285.6 mg/1)
was noted for the Far Eastern Amurocrangonyx (Birstein & Eevanidov 1952). It is interesting that
S. ambulans apparently has a feature that ean distinguish it from the other speeies of the genus, namely
a yellowish spot (url: www.biospeleo.ru/S_ambulans.htm ) whieh is the hypertrophied digestive
gland. Previously, the observation of a yellowish (orange) spot for the speeies was mentioned by
Muller (1846), Stebbing (1906), Borutzky (1929) and Boets et al. (2010). The hypertrophied digestive
gland (HDG) is eharaeteristie also of several deep-sea marine amphipods, e.g. Eurythenes gryllus
Eiehtenstein, 1822 where it performs the leading role in the metabolie response of the defense
antioxidant system (Camus & Gulliksen 2004). In subterranean amphipods the HDG has also been
observed in Amurocrangonyx.

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European Journal of Taxonomy 23: 1-19 (2012)

Acknowledgements

We thank Mikhail Chertoprud (Moscow State University) for collection and forwarding material to us
and Oliver Coleman (Humboldt University, Berlin) for loan of specimens of Synurella ambulans from
the Museum far Naturkunde in Berlin. Both loans made this study possible. We would also like to thank
Rudy Jocque (Royal Museum for Central Africa, Tervuren), Kristiaan Hoedemakers (Royal Belgian
Institute of Natural Sciences, Brussels), Oliver Coleman and an anonymous reviewer for critically
reading and commenting on the manuscript, largely improving content and style of the paper.

The study was funded with partial financial support of the Russian Foundation for Basic Research grant
09-04-98544 and of the Presidium of FEBRAS grants ll-lll-B-06-098 and 12-l-n30-01.

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Manuscript received: 28 April 2012
Manuscript accepted: 30 July 2012
Published on: 27 September 2012
Topic editor: Rudy Jocque

In compliance with the ICZN, printed versions of all papers are deposited in the libraries of the institutes
that are members of the EJT consortium: Museum National d’Histoire Naturelle, Paris, France; National
Botanic Garden of Belgium, 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.

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