Genus Vol. 15(1): 13-24 Wroc³aw, 30 III 2004Ecdyonurus austriacus nataliae n. ssp., a new subspecies of theEcdyonurus helveticus-group from Ukraine(Ephemeroptera: Heptageniidae)ROMAN J. GODUNKO1 and MA£GORZATA K£ONOWSKA-OLEJNIK21State Museum of Natural History, National Academy of Sciences of Ukraine, Teatral’na 18,L’viv 79008, Ukraine; e-mail: godunko@museum.lviv.net2Department of Hydrobiology, Institute of Environmental Sciences, Jagiellonian University,Gronostajowa 3, 30-387 Kraków, Poland; e-mail: klon@eko.uj.edu.plABSTRACT. A new subspecies, Ecdyonurus austriacus nataliae ssp. n., belongingto the helveticus-group, is described and illustrated at all stages (imagines, subimagines,
nymphs and eggs) from material collected in the Gorgany Mts. in the Eastern Carpathians,
Ukraine. The new subspecies is closely related to Ecdyonurus austriacus austriacus
Kimmins. Its affinities, distribution and ecology are discussed.Key words: entomology, taxonomy, Ephemeroptera, Heptageniidae, new subspecies,nymph, imago and subimago, egg, distribution, ecology, Eastem Carpathians,
Ukraine.INTRODUCTIONSpecies of the Ecdyonurus helveticus-group are characterized by the lack oflong and dense setae on the distal part of larval hypopharynx, and by the presence
of a laterally elongated apical sclerite of the penis lobes of imago (HEFTI & TOMKA1986, STUDEMANN et al. 1992). Some taxonomical revisions and descriptions ofnew species from the E. helveticus-group were published (KIMMINS 1958, BOGOESCU& TABACARU 1962, JACOB & BRAASCH 1984, HEFTI et al. 1986, 1989, 1991,BELFIORE & BUFFAGNI 1994, BAUERNFEIND 1997). The Central European fauna ofthe E. helveticus-group includes 12 taxa (HEFTI et al. 1989). Five species knownfrom literature are found in the Ukrainian Carpathians. Ecdyonurus subalpinus
(KLAPÁLEK, 1907) from the Gorgany Mts. was described by KLAPÁLEK (1907). In14ROMAN J. GODUNKO, MA£GORZATA K£ONOWSKA-OLEJNIKthe same paper, KLAPÁLEK cited Ecdyonurus helveticus (EATON, 1885) from theChornogora Mts. Ecdyonurus picteti (MEYER-DüR, 1864) from the Chornogoramountain range was described as Heptagenia nigrescens (DZIÊDZIELEWICZ &KLAPÁLEK 1908a, 1908b) and as Ecdyonurus nigrescens (MIKULSKI 1936). Somedata about the distribution of species of the E. helveticus-group in the Ukrainian
Carpathians were also presented by DZIÊDZIELEWICZ (1907, 1919), MIKULSKI (1935),PAW£OWSKI (1959), GODUÑKO (1997, 1999), and GODUNKO (1998). Recently, a newsubspecies belonging to the E. helveticus-group from the Gorgany Mts. in the
Eastern Carpathians, Ukraine has been found and described.Ecdyonurus austriacus nataliae n. ssp.ETYMOLOGYThe new subspecies is named in honor of NATALIA O. GODUNKO, wife of thefirst author.MALEIMAGOSize: body length: 13.4-15.7 mm; fore wings: 13.6-14.5 mm; cerci: 28.2-35.1mm. Head light brown with a lighter margin and yellowish-gray spots. Scapus and
pedicle yellowish-brown. Eyes light gray with a submarginal dark band. Ocelli
white at the apical part with a brownish band in border. Thorax light brown. Fore
legs of the same coloration as the thorax; tibia and tarsi brownish, generally paler
than the femora; coxa and trochanter light yellowish-brown. Middle and hind legs
paler, yellowish-brown, unicolorous; tibia and tarsi paler, yellowish and light
yellowish-brown; femora yellowish-brown, light brownish-gray distally; coxa and
trochanter of the middle and hind legs slightly darker. Fore wings hyaline,
transparent, light milk color, grayish, unicolorous; venation dark, well visible;
C, Sc and R paler, yellowish-brown; other longitudinal and transversal veins
darker, brownish-castaneous; costal and subcostal area milky; pterostigmatic area
brownish and opaque. Hind wings of the same color as the fore wings, with dark
venation. Abdomen yellowish-brown, sometimes with small reddish-brown spots
in the lateral part; lateral part of the terga with the typical pattern of the
E. helveticus-group (as in female imago, Fig. 3); surface of the terga with two
small visible paler spots in the central part; posterior margins of the terga slightly
darker; sterna paler, yellowish-brown with well visible dark violet nerve ganglia
and four small spots (two in the central part and two laterally oriented). Genitalia:
styliger plate brown, posterior margin with two strong protuberances. In dorsal
view the outlines of the penis lobes are rounded and slightly stretched laterally.
The distal borders of the lobes are rounded (Fig. 1). The inner margin of the apical
sclerite has one strong stout tooth directed towards the center of the lobe (some-
times with two strong stout teeth) and with some small submarginal teeth. The
lateral sclerite is large without deep narrowing in the external part and softly15ECDYONURUS AUSTRIACUS NATALIAE N. SSP.1-4. Ecdyonurus austriacus nataliae: 1-2. Male imago: 1-male genitalia, ventral view; 2 - penis
lobe, dorsal view; 3-4. Female imago: 3 - posterior part of abdomen, lateral view; 4 - posterior partof abdomen, ventral view16ROMAN J. GODUNKO, MA£GORZATA K£ONOWSKA-OLEJNIKrounded in the internal part; the basal sclerite usually supports one well visible
tooth and 2-4 irregular small teeth. A small protuberance is often presented
outside the penis stem (Fig. 2). Cerci brown.FEMALEIMAGOSize: body length: 11.9-14.6 mm; fore wings: 19.6-23.7 mm; cerci: 13.3-16.4mm. General color of the body brownish with reddish-brown spots on the lateral
part of the abdominal and thoracic segments, darker as with the imago male. Head
brown with yellowish-gray and reticulated dark spots. Eyes brownish-gray with a
submarginal dark band. Ocelli brownish-gray. Thorax dark, brownish-gray, some-
times brownish-castaneous. Fore legs of the same coloration as the thorax. Middle
and hind legs yellowish-brown with darker tarsi. Fore wings hyaline, transparent;
venation dark, well visible; costal and subcostal zones light yellowish;
pterostigmatic area milky, sometimes brownish. Hind wings hyaline with dark
venation. Terga brown and brownish-castaneous with reddish spots laterally.
Sterna of the same coloration as the terga, with dark nerve ganglia. The drawing
on the surface of the sterna is similar to that of the male imago. Subgenital plate
relatively wide; posterior margin slightly rounded. An apical outline of the
subanal plate can be seen in Figs. 3, 4. Cerci brown.MALESUBIMAGOSize: body length: 13.9 mm; fore wings: 15.7 mm; cerci: 15.9 mm. Generalcolor of the body yellowish-brown with reddish-brown spots around the typical
drawing on the abdominal terga. Thorax and legs with small reddish spots. Head
grayish-brown with paler spots on margin; antennae brownish with a lighter
scapus; eyes and ocelli gray. Dorsal part of the thorax yellowish with brown and
yellowish-brown bands; lateral and ventral part yellowish-brown; segments of the
thorax are contrasted with darker abdominal segments. General color of legs
yellowish brown; fore legs darker with paler tarsi; middle and hind legs light,
grayish on the dorsal part of the femora. Fore wings opaque with yellowish-gray
venation; well visible drawing formed by transversal, intensive grayish bands
(zigzag type) pointed at the distal part. Hind wings with only one paler band.
Terga brownish-gray with a drawing on the surface similar to the male imago.
Sterna yellowish with dark nerve ganglia. General coloration of genitalia yellow-
ish-gray with darker forceps. In dorsal view, the outlines of the penis lobes are
rounded and slightly stretched laterally. Base of forceps with two strong lateral
protuberances. Cerci light brown.FEMALESUBIMAGOSize: body length: 12.1-14.2 mm; fore wings 15.5-16.8 mm; cerci 11.9-15.9mm. General color similar to the male subimago. Reddish-brown spots are pre-
sented on the lateral part of the thoracic and abdominal segments, as well as on the
femora.17ECDYONURUS AUSTRIACUS NATALIAE N. SSP.MATURENYMPHSize: body length: 10.3-15.5 mm; caudal filaments: 9.5-11.0 mm. Generalcolor of body yellowish-brown to brown, with a light grayish tinge. Head rela-
tively wide, yellowish-brown to brown, with paler spots or a light zone on the
anterior part. Antennae brown. Labrum relatively wide with well developed5-16. Ecdyonurus austriacus nataliae, nymph: 5 - outline of labrum; 6 - bristles on the ventral side
of labrum; 7 - hypopharynx; 8 - fragment of labium with glossa and paraglossa; 9 - lateral pronotum
expansion; 10 - hind leg; 11 - spines of central part of dorsal surface of the femora; 12 - tarsal claw18ROMAN J. GODUNKO, MA£GORZATA K£ONOWSKA-OLEJNIK13-16. Ecdyonurus austriacus nataliae, nymph: 13 - abdomen, dorsal view; 14 - central part ofposterior margin of fifth abdominal tergum; 15 - first gill; 16 - fourth gill19ECDYONURUS AUSTRIACUS NATALIAE N. SSP.lateral projections and a slight depression in the middle of anterior margin (Fig.
5). The bristles in the rows on the ventral side of the labrum are rather slender.
The first 1-4 bristles are shorter than the others (Fig. 6). Basal part of superlinguae
of hypopharynx relatively wide and massive (Fig. 7); the pilosity of the hypophar-
ynx is typical of the E. helvetius-group. Maxilla: 5-11 fine bristles on the fore
margin of the first segment of maxillary palpus; more than 40 bristles along the
outer margin; more than 30 bristles on dorsal side of the galea-lacinia. Glossa of
labium as in Fig. 8; paraglossa massive with a rounded outer lateral projection.
External margins of glossae and paraglossae with numerous fine setae. Thorax:
general coloration yellowish-brown to brown with small reddish spots on the
surface. Pronotum with paler spots and bands. External outline of the lateral
pronotum expansion not regularly curved and rounded distally. Apex of the
pronotum expansion strong, directed towards the body (Fig. 9). Legs yellowish-
brown to brown with darker tarsi. Femora sometimes with small brownish-red
spots. Femora slender and long. Length/width ratio of hind femora on average
2.84 (s = 0.203; n = 20). Surface of femora with a more or less visible drawing
(four dark spots on a paler background, as in Fig. 10). Femoral spines rounded at
the apex, generally in the form of a spatula (Fig. 11). Ventral side of posterior
margin of hind femora with 23-34 (mean 28) spines and a rather short setae. Tarsal
claws generally with 1-2 teeth (seldom 0-3) (Fig. 12). Abdomen: general color
yellowish-brown to brown with small reddish spots. Lateral expansion of the
abdominal segments short and pointed. Surface of the terga with a more or less
distinct drawing: I-II segments relatively paler and centrally smudged; III-IV
segments uniformly colored, with two paler spots on central part of the segment
and two paler spots laterally; V segment sometimes lighter, with one central paler
spot and two paler spots laterally; VI-VII segments uniformly colored, with three
small paler spots placed centrally and two paler spots laterally; VIII-IX segments
uniformly colored, with a big central paler spot and a darker lateral part of the
segments; X segment generally dark, with only two small paler spots (Fig. 13).
Posterior margins of the terga with two kinds of spines: the first is rather small and
pointed at the apex; the second is large, extended and rounded at the apex in the
form of a spatula, sometimes with a laterally pointed one from a common basis
(Fig. 14). Surface of the terga with small pointed and rounded spines and long fine
setae. Gills milky to yellowish in color, sometimes slightly grayish or reddish-
brown. Gill 1 relatively long (Fig. 15); gill 4 wide and asymmetrical (Fig. 16).
Length/width ratio of the 4th gill on average 1.93 (s = 0.153; n = 20). Caudal
filaments brown to dark brown.EGGEgg oval. Size: length:173-181µm; width 107-115 µm (Fig. 17). Chorionicsurface with a clear pattern (tubercles and attachment structures). Small rounded
tubercles (0.8-1.6 µm in diameter) are quite densely and evenly distributed on the
chorionic surface. There is a delicate granular ground matrix covering both the20ROMAN J. GODUNKO, MA£GORZATA K£ONOWSKA-OLEJNIKchorionic surface and the tubercles (Fig. 18). Attachment structures are character-
ized by knob-terminated coiled threads (KCTs) (KOSS & EDMUNDS 1974). Theyare small, unanimous in size (2.2-2.7 µm in diameter) and cover the whole
chorionic surface of the egg quite densely (4.4-8.0 µm distance between them)
(Fig. 20). There are no concentrations of KCT attachment structures on the poles
of the egg. Three to five micropyles can be found in the subequatorial area. Sperm
guide ovoidal, 8.5-10.4 µm in length and 6.5-7.7 µm in width; a micropylar
opening is situated to the side. Micropylar rim thin, with a few sparsely distributed
tubercles (Fig. 19).17-20. Ecdyonurus austriacus nataliae, egg: 17 - general outline of the egg, scale bar = 50 µm; 18-
chorionic surface, scale bar = 10 µm; 19 - micropyle, scale bar = 5 µm; 20 - detail of chorionic
surface with tubercles and attachment structures (KCTs) (ground matrix visible), scale bar = 1 µm(JEOL JSM 5410 scanning microscope)21ECDYONURUS AUSTRIACUS NATALIAE N. SSP.MATERIALEXAMINEDHolotype: 1 male imago, Ukraine, Ivano-Frankivs’k Region, Gorgany Mts.,Natural Reserve “Gorgany”, the Dzhurdzhi brook, 900-950 m. a.s.l., 48°29'14" E,
24°19'21" N, 23-26.VII.2000, leg. R.J. GODUNKO, M. K£ONOWSKA-OLEJNIK,K. PRZYBY£A & P. ZASÊPA. Paratypes: 4 male imagines (reared from larvae, partlyon microscopic slides), 9 female imagines, 2 female subimagines (reared from
larvae, partly on microscopic slides), 6 subimaginal skins, 2 nymphal exuviae, 28
nymphs, from same locality and date. Other material: 4 larvae, Ukraine, Ivano-
Frankivs’k Region, Gorgany Mts., Natural Reserve “Gorgany”, the Dzhurdzhi
brook, 900-950 m. a.s.l., 24.VII.1997, leg. R.J. GODUNKO & M. K£ONOWSKA-OLEJNIK; 9 nymphs, same locality, 20.VI.1999, leg. R.J. Godunko; 7 femaleimagines, 1 male subimago, 4 female subimagines, 3 subimaginal skins, 96
nymphs (partly on slides), same locality, 28.VI.2000, leg. R.J. GODUNKO, M. PUTZ& M. ŠTROJSOVá; 5 nymphs (partly on slides), Ukraine, Ivano-Frankivs’k Region,Gorgany Mts., Natural Reserve “Gorgany”, the Dovzhynets’stream, Ozirnyi dis-
trict, 800-900 m. a.s.l., 48°22'48" E, 24°17'57" N, 22.VII.1997, leg. R.J. GODUNKO& M. K£ONOWSKA-OLEJNIK; 10 nymphs (partly on slides), same locality, 29-30.VII.1998, leg. R.J. GODUNKO; 10 nymphs, same locality, 25.VII.2000, leg. R.J.GODUNKO & M. K£ONOWSKA-OLEJNIK; 8 nymphs, Ukraine, Ivano-Frankivs’k Re-gion, Gorgany Mts., Bohdan stream, 750 m a.s.l., 48°21'40" E, 24°28'42" N,
18.VII.1999, leg. R.J. GODUNKO, M. K£ONOWSKA-OLEJNIK, K. PRZYBY£A & P.ZASÊPA. All specimens were preserved in 75% alcohol. The holotype and some ofthe paratypes are housed in the collection of the State Museum of Natural History,
National Academy of Sciences of Ukraine, L’viv, Ukraine. Other material can be
found in the second author’s collection at Jagiellonian University, Cracow, Poland.AFFINITIESE. austriacus nataliae belongs to the E. helveticus-group due to the followingcharacteristics: 1) distal part of larval hypopharynx without long and dense setae;
2) lateral expansion of larval abdominal segments short and directed in parallel to
the axis of the body; 3) apical sclerite of the penis lobes of imago laterally
elongated; 4) lateral part of the adult terga with a characteristic pattern.
E. austriacus nataliae is closely related to Ecdyonurus austriacus austriacus
KIMMINS, 1958 especially in adult stages. Genitalia of male imagines of E. austriacusnataliae partly resemble E. austriacus austriacus but can be distinguished by: 1)
inner margin of apical sclerite with one (sometimes with two) strong stout tooth
directed towards the center of the lobe and with some small submarginal teeth; 2)
lateral sclerite softly rounded in the internal part; 3) penis lobes are less stretched
laterally; 4) the color and size of body and wings. The combination of these
characteristics clearly distinguishes E. austriacus nataliae from all the East
European species of the E. helveticus-group. E. carpathicus SOWA, 1973 andE. subalpinus are both separable from E. austriacus nataliae because of the
following characteristics: 1) penis lobes are more stretched laterally with more
spherical outline; 2) the lateral sclerite is relatively slender with a deep narrowing22ROMAN J. GODUNKO, MA£GORZATA K£ONOWSKA-OLEJNIKin the external part (SOWA 1973, HEFTI et al. 1989). The shape of basal sclerite ofthe penis lobes distinguishes E. austriacus nataliae from E. picteti as the latter
species bears strong large teeth, projected perpendicularly to the axis of symmetry
of the penis lobes (HEFTI & TOMKA 1986, HEFTI et al. 1989). The lateral sclerite ofthe penis lobes of E. helveticus, in contrast to E. austriacus nataliae, is slender
with a deep narrowing in the external part (HEFTI et al. 1987).The male and female subimagines of E. austriacus nataliae resembleE. austriacus austriacus in terms of general body and wing color (KIMMINS 1958,HEFTI & TOMKA 1986), but is distinguished by: the size of the body and wings; andthe character of the sclerites of penis lobes in male subimago (similar to male
imago). The fore wings of E. austriacus nataliae present a distinct zigzag pattern.
Other species of the E. helveticus-group have no zigzag pattern on the fore wings
or, if it is visible (as in E. austriacus austriacus, E. helveticus, Ecdyonurus
parahelveticus HEFTI, TOMKA & ZURWERRA, 1986 and E. picteti) its color is notintensive. The wings coloration of a new subspecies is similar to that of the
specimens in populations of E. subalpinus KLAPÁLEK, 1907 from the CzechRepublic (SOLDÁN, personal communication). On the contrary, no such relation-ships have been observed in the E. subalpinus populations from the East
Carpathians.In the nymphal stages E. austriacus nataliae is easily distinguishable fromother species belonging to the E. helveticus-group. The tergo-abdominal spines of
larvae of E. austriacus nataliae are of the same type as E. austriacus austriacus,
E. parahelveticus, and Ecdyonurus alpinus HEFTI, TOMKA & ZURWERRA, 1987(HEFTI & TOMKA 1986, HEFTI et. al. 1986, 1987), i. e. large and apically rounded,but their shape and arrangement are very characteristic. Additionally, E. austriacus
nataliae differs from E. austriacus austriacus, E. parahelveticus and E. alpinus
by the number and shape of the teeth on the tarsal claws, the shape of the labrum,
hypopharynx and tergo-abdominal spines and by the drawing of the abdomen and
legs (HEFTI & TOMKA 1986, HEFTI et al. 1986, 1987, 1989). Eggs of E. austriacusnataliae are different from the other species of the E. helveticus-group. A study in
light microscopy showed that the most common chorionic organization is densely
granulated, and the granules are concentrated on both poles of the eggs (HEFTI &TOMKA 1986, HEFTI et al. 1986, 1987, 1989). It seems probable that this concentra-tion concerns KCTs attachment structures. In E. alpinus, E. austriacus austriacus,
E. helveticus and E. parahelveticus KCTs attachment structures are distinctly
concentrated at two poles of the eggs (HEFTI & TOMKA 1986, HEFTI et al. 1986,1987). KCTs in E. zelleri (EATON, 1885) egg poles are less frequent (HEFTI et al.1987). Only E. picteti has KCTs concentration at one pole of the egg (HEFTI &TOMKA 1986). Thus, E. austriacus nataliae with no concentration of KCTsattachment structure at the egg poles is easily distinguishable from previously
mentioned species.23ECDYONURUS AUSTRIACUS NATALIAE N. SSP.DISTRIBUTIONANDECOLOGYE. austriacus nataliae has been found in small mountain brooks and streamsbetween 700-1200 m a.s.l., 1-4 m width and 0.2-1.5 m depth, with cold water (in
summer ~10°C) and current velocity higher than 1 m/s. The bottom is covered
with boulders and cobbles, and the banks are high and shaded by subalpine forest.
The species occurs up to the timberline, the most frequently - between 800-900 m
a.s.l.. In these localities E. austriacus nataliae lives together with Ameletus
inopinatus EATON, Baetis alpinus (PICTET), B. melanonyx (PICTET), Rhithrogenairidina (KOLENATI), R. gorganica KLAPÁLEK, and E. subalpinus. E. austriacusnataliae is a univoltine winter species (Uw) with one generation per year (CLIFFORD1982). Population overwinters in the nymphal stage. The larvae occur gradually
and grow during the winter. The flying period extends from June to August-
September. E. austriacus nataliae has been found only in the Eastern Carpathians,
Gorgany Mts., Bystrytsya Nadvirnyans’ka river-basin (tributaries of the Dnister
river; two localities) and in the Prutets’ Yablons’kyi river-basin (tributaries of the
Prut river; one locality). MIKULSKI (1935), in his paper about the Ephemeropteraspecies of the Chornohora Mts., described and illustrated the larval abdomen of
“Ecdyonurus sp. larvae (subalpinus?)” and “Ecdyonurus sp. larva”. Some details
are similar to the drawing of the abdomen of E. austriacus nataliae (Figs. 1, 2, 4)
but other characteristics needed to determine this taxon are absent in MIKULSKI’sdescription. Moreover, at that time, E. austriacus nataliae was not found in the
Chornohora mountain range. Thus, E. austriacus nataliae seems to be an endemic
of the Eastern Carpathians.ACKNOWLEDGMENTSWe would like to express our sincere thanks to Prof. T. SOLDÁN for his helpand many valuable comments on the manuscript. M. PUTZ, M. ŠTROJSOVÁ, K.PRZYBY£A and P. ZASÊPA helped us with collecting type material. We thank Mr. V.M. KYSLYAK, director of the Natural Reserve “Gorgany” for help in the field work.The SEM photographs of the eggs were taken on a scanning electron microscope
(JEOL JSM 5410) in the Department of Cytology and Histology at Jagiellonian
University, Cracow. We are indebted to Mrs. J. FABER for technical assistance.The research was partially financially supported by Jadwiga Queen Fund of
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