Phylogenetic Relationships of Coelacanthia and Archaschenia, Two Spinose Rissoids (Mollusca, Gastropoda) from the Miocene of the Eastern Paratethys
Миоценовые риссоиды родов Archaschenia Zhgenti, 1981 и Coelacanthia Andrusov, 1890 не связаны друг с другом непосредственным родством, однако имеют общих предков в эволюционной ветви рода Mohrensternia Stoliczka, 1868. Они произошли от разных видов Mohrensternia, которые проникали в Понто-Каспийскую...
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irk-123456789-32202009-09-01T17:23:10Z Phylogenetic Relationships of Coelacanthia and Archaschenia, Two Spinose Rissoids (Mollusca, Gastropoda) from the Miocene of the Eastern Paratethys Anistratenko, V.V. Миоценовые риссоиды родов Archaschenia Zhgenti, 1981 и Coelacanthia Andrusov, 1890 не связаны друг с другом непосредственным родством, однако имеют общих предков в эволюционной ветви рода Mohrensternia Stoliczka, 1868. Они произошли от разных видов Mohrensternia, которые проникали в Понто-Каспийскую область Восточного Паратетиса из Средиземноморья. Archaschenia merklini Zhgenti, 1981 и A. iljinae Zhgenti, 1981 произошли от Mohrensternia barboti Andrusov, 1890 или близкого вида в карагане, тогда как вид Coelacanthia quadrispinosa Andrusov, 1890 обособился от Mohrensternia subinflata Andrusov, 1890 в мэотисе. Имеющиеся морфологические данные убеждают, что Rissoa Desmarest, 1814, Mohrensternia, Archaschenia и Coelacanthia должны рассматриваться как особые роды в пределах семейства Rissoidae. Недавнее установление для этих групп (Анистратенко, 2003) семейства Mohrensterniidae Korobkov, 1955 (с подсемействами Mohrensterniinae Korobkov, 1955, Coelacanthiinae Anistratenko, 2003 и Archascheniinae Zhgenti, 1991) признается теперь излишним. 2004 Article Phylogenetic Relationships of Coelacanthia and Archaschenia, Two Spinose Rissoids (Mollusca, Gastropoda) from the Miocene of the Eastern Paratethys / V. V. Anistratenko // Вестн. зоологии. — 2004. — Т. 38, № 2. — С. 3-12. — англ. 0084-5604 http://dspace.nbuv.gov.ua/handle/123456789/3220 564.329”6235”(262) en Інститут зоології ім. І. І. Шмальгаузена НАН України |
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Миоценовые риссоиды родов Archaschenia Zhgenti, 1981 и Coelacanthia Andrusov, 1890 не связаны друг с другом непосредственным родством, однако имеют общих предков в эволюционной ветви рода Mohrensternia Stoliczka, 1868. Они произошли от разных видов Mohrensternia, которые проникали в Понто-Каспийскую область Восточного Паратетиса из Средиземноморья. Archaschenia merklini Zhgenti, 1981 и A. iljinae Zhgenti, 1981 произошли от Mohrensternia barboti Andrusov, 1890 или близкого вида в карагане, тогда как вид Coelacanthia quadrispinosa Andrusov, 1890 обособился от Mohrensternia subinflata Andrusov, 1890 в мэотисе. Имеющиеся морфологические данные убеждают, что Rissoa Desmarest, 1814, Mohrensternia, Archaschenia и Coelacanthia должны рассматриваться как особые роды в пределах семейства Rissoidae. Недавнее установление для этих групп (Анистратенко, 2003) семейства Mohrensterniidae Korobkov, 1955 (с подсемействами Mohrensterniinae Korobkov, 1955, Coelacanthiinae Anistratenko, 2003 и Archascheniinae Zhgenti, 1991) признается теперь излишним. |
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Anistratenko, V.V. |
| spellingShingle |
Anistratenko, V.V. Phylogenetic Relationships of Coelacanthia and Archaschenia, Two Spinose Rissoids (Mollusca, Gastropoda) from the Miocene of the Eastern Paratethys |
| author_facet |
Anistratenko, V.V. |
| author_sort |
Anistratenko, V.V. |
| title |
Phylogenetic Relationships of Coelacanthia and Archaschenia, Two Spinose Rissoids (Mollusca, Gastropoda) from the Miocene of the Eastern Paratethys |
| title_short |
Phylogenetic Relationships of Coelacanthia and Archaschenia, Two Spinose Rissoids (Mollusca, Gastropoda) from the Miocene of the Eastern Paratethys |
| title_full |
Phylogenetic Relationships of Coelacanthia and Archaschenia, Two Spinose Rissoids (Mollusca, Gastropoda) from the Miocene of the Eastern Paratethys |
| title_fullStr |
Phylogenetic Relationships of Coelacanthia and Archaschenia, Two Spinose Rissoids (Mollusca, Gastropoda) from the Miocene of the Eastern Paratethys |
| title_full_unstemmed |
Phylogenetic Relationships of Coelacanthia and Archaschenia, Two Spinose Rissoids (Mollusca, Gastropoda) from the Miocene of the Eastern Paratethys |
| title_sort |
phylogenetic relationships of coelacanthia and archaschenia, two spinose rissoids (mollusca, gastropoda) from the miocene of the eastern paratethys |
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Інститут зоології ім. І. І. Шмальгаузена НАН України |
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2004 |
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http://dspace.nbuv.gov.ua/handle/123456789/3220 |
| citation_txt |
Phylogenetic Relationships of Coelacanthia and Archaschenia, Two Spinose Rissoids (Mollusca, Gastropoda) from the Miocene of the Eastern Paratethys / V. V. Anistratenko // Вестн. зоологии. — 2004. — Т. 38, № 2. — С. 3-12. — англ. |
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2025-07-02T06:29:12Z |
| last_indexed |
2025-07-02T06:29:12Z |
| _version_ |
1836515592942649344 |
| fulltext |
UDC 564.329”6235”(262)
PHYLOGENETIC RELATIONSHIPS OF COELACANTHIA
AND ARCHASCHENIA, TWO SPINOSE RISSOIDS
(MOLLUSCA, GASTROPODA) FROM THE MIOCENE
OF THE EASTERN PARATETHYS
V. V. Anistratenko
Schmalhausen Institute of Zoology of NAS of Ukraine,
vul. B. Khmelnits’kogo, 15, Kyiv-30, MSP, 01601 Ukraine
Accepted 11 February 2004
Phylogenetic Relationships of Coelacanthia and Archaschenia, Two Spinose Rissoids (Mollusca,
Gastropoda) from the Miocene of the Eastern Paratethys. Anistratenko V. V. — The Miocene rissoids
Archaschenia Zhgenti, 1981 and Coelacanthia Andrusov, 1890 are not directly related, but have
common ancestors in the Mohrensternia Stoliczka, 1868 lineage. They have evolved from the different
species of Mohrensternia, which immigrated into the Ponto-Caspian area of the Eastern Paratethys from
the Mediterranean. Archaschenia merklini Zhgenti, 1981 and A. iljinae Zhgenti, 1981 have evolved from
Mohrensternia barboti Andrusov, 1890 or a related species during the Karaganian, whereas Coelacanthia
quadrispinosa Andrusov, 1890 evolved from Mohrensternia subinflata Andrusov, 1890 during the Maeo-
tian. The morphological data available suggest that Rissoa Desmarest, 1814, Mohrensternia, Archasche-
nia and Coelacanthia should be considered as separate genera within the family Rissoidae. The recently
introduced for these taxa (Anistratenko, 2003) family Mohrensterniidae Korobkov, 1955 (with the
subfamilies Mohrensterniinae Korobkov, 1955, Coelacanthiinae Anistratenko, 2003 and Archascheni-
inae Zhgenti, 1991) is considered as insufficiently substantiated.
Ke y wo r d s: Gastropoda, Rissooidea, phylogeny, taxonomy, Miocene, Paratethys.
Ôèëîãåíåòè÷åñêèå îòíîøåíèÿ Coelacanthia è Archaschenia, äâóõ ðîäîâ «øèïàñòûõ» ðèññîèä
(Mollusca, Gastropoda) èç ìèîöåíà Âîñòî÷íîãî Ïàðàòåòèñà. Àíèñòðàòåíêî Â. Â. — Ìèîöåíîâûå
ðèññîèäû ðîäîâ Archaschenia Zhgenti, 1981 è Coelacanthia Andrusov, 1890 íå ñâÿçàíû äðóã ñ
äðóãîì íåïîñðåäñòâåííûì ðîäñòâîì, îäíàêî èìåþò îáùèõ ïðåäêîâ â ýâîëþöèîííîé âåòâè ðîäà
Mohrensternia Stoliczka, 1868. Îíè ïðîèçîøëè îò ðàçíûõ âèäîâ Mohrensternia, êîòîðûå ïðîíèêàëè
â Ïîíòî-Êàñïèéñêóþ îáëàñòü Âîñòî÷íîãî Ïàðàòåòèñà èç Ñðåäèçåìíîìîðüÿ. Archaschenia merklini
Zhgenti, 1981 è A. iljinae Zhgenti, 1981 ïðîèçîøëè îò Mohrensternia barboti Andrusov, 1890 èëè
áëèçêîãî âèäà â êàðàãàíå, òîãäà êàê âèä Coelacanthia quadrispinosa Andrusov, 1890 îáîñîáèëñÿ îò
Mohrensternia subinflata Andrusov, 1890 â ìýîòèñå. Èìåþùèåñÿ ìîðôîëîãè÷åñêèå äàííûå óáåæ-
äàþò, ÷òî Rissoa Desmarest, 1814, Mohrensternia, Archaschenia è Coelacanthia äîëæíû ðàññìàòðè-
âàòüñÿ êàê îñîáûå ðîäû â ïðåäåëàõ ñåìåéñòâà Rissoidae. Íåäàâíåå óñòàíîâëåíèå äëÿ ýòèõ ãðóïï
(Àíèñòðàòåíêî, 2003) ñåìåéñòâà Mohrensterniidae Korobkov, 1955 (ñ ïîäñåìåéñòâàìè Mohren-
sterniinae Korobkov, 1955, Coelacanthiinae Anistratenko, 2003 è Archascheniinae Zhgenti, 1991)
ïðèçíàåòñÿ òåïåðü èçëèøíèì.
Êëþ÷åâûå ñ ëîâ à: Gastropoda, Rissooidea, ôèëîãåíèÿ, ñèñòåìàòèêà, ìèîöåí, Ïàðàòåòèñ.
Introduction
The Rissooidea is one of the largest and most diversified gastropod groups worldwide. It occupies full
marine to fresh-water habitats (Wenz, 1938–1944; Ponder, 1985; Anistratenko, Starobogatov, 1994). The
Miocene fossil record of Paratethys reveals a wide range of rissoid morphologies, which are usually related to
different ecological conditions. During this time many endemic lineages have evolved, of which the best
known one is the Mohrensternia Stoliczka, 1868 lineage. The gastropods of this genus became one of most
common components of the semi-marine to brackish-water gastropod associations in the Paratethys
(Stoliczka, 1867–1868; Andrusov, 1890, 1906; Friedberg, 1911–1928; Kolesnikov, 1935; Zhizhchenko, 1936;
Svagrovsky, 1971; Il’ina, 1972 a, b, 1979; Il’ina et al., 1976).
Although the genus Mohrensternia is well known to West European readers (see review by Kowalke,
Harzhauser, 2004), other related gastropods have been studied only by paleontologists of the former USSR.
Vestnik zoologii, 38(2): 3–12, 2004
© V. V. Anistratenko, 2004
The subfamily Mohrensterniinae was introduced by Korobkov (1955), but its taxonomy and generic/specific
content have not been adequately evaluated (Badzoshvili, 1991; Zhgenti, 1981, 1991; Il’ina, 1993).
The aim of this paper is to discuss possible taxonomic significance of shell morphological characters of
some Mohrensternia-related gastropods and to trace their possible phylogenetic relationships. A review of the
poorly known homeomorphic genera Coelacanthia Andrusov, 1890 and Archaschenia Zhgenti, 1981 is
provided and the systematic position of these taxa is evaluated.
Material and methods
The present investigation is mainly based on large collections of Miocene gastropods, which are housed
at the Institute of Geological Sciences of the National Academy of Sciences of the Ukraine and at the
Schmalhausen Institute of Zoology NAS of Ukraine (Kyiv). The material comes from natural outcrops and
borehole of Sarmatian deposits in western Ukraine. More than 3 000 specimens of the Mohrensternia and
Rissoa Desmarest, 1814 identified in studied samples. Also I examined three specimens of Coelacanthia
quadrispinosa Andrusov, 1890 from the type collection of Andrusov (1890). Andrusov collected this material
from Maeotian deposits near Pavlovsky Cape on Kerch Peninsula (fig. 1). These samples quite possibly are
paralectotypes for this species as Il’ina (2001) designated the lectotype from the same locality.
An optical stereoscopic microscope (MBS-9) and a simple “drawing tube” were used for studying and
drawing the shell characters. The SEM micrographs were made at the Institute of Paleobiology, Polish Academy
of Sciences in Warsaw (Poland), abbreviated ZPAL Ga. 11/1; the illustrated specimen is housed there.
4 V. V. Anistratenko
Fig. 1. Location of the outcrops and borehole in the western Ukraine and on Kerch peninsula: PIL — outcrop
between Pilyava and Ivankovcy villages, Staraya Sinyava district, Khmelnitsky region; KOS — outcrop
between Kos’kov and Grycev villages, Shepetovka district, Khmelnitsky region; LIT — borehole 1529 near
Litinka village, Litin district, Vinnitsky region; ANT — outcrop near Antonovka village, Letichev district,
Khmelnitsky region; LET — outcrop near Letichev, Khmelnitsky region; SAT — outcrop near Satanov town,
Khmelnitsky region; IZA — precipice on the bank of river Goryn, vicinity of Izyaslav town, Khmelnitsky
region; A — Andrusov’s (1890) record of Coelacanthia quadrispinosa in Kertsch peninsula — type locality.
Ðèñ. 1. Ìåñòîíàõîæäåíèå îáíàæåíèé è ñêâàæèí â çàïàäíîé ÷àñòè Óêðàèíû è íà Êåð÷åíñêîì ï-âå:
PIL — îáíàæåíèå ìåæäó ñåëàìè Ïèëÿâà è Èâàíêîâöû, Ñòàðîñèíÿâñêèé ð-í, Õìåëüíèöêàÿ îáë.;
KOS — îáíàæåíèå ìåæäó ñåëàìè Êîñüêîâ è Ãðèöåâ, Øåïåòîâñêèé ð-í, Õìåëüíèöêàÿ îáë.; LIT —
ñêâàæèíà 1529 âîçëå ñ. Ëèòèíêà, Ëèòèíñêèé ð-í, Âèííèöêàÿ îáë.; ANT — îáíàæåíèå âîçëå ñ. Àíòî-
íîâêà Ëåòè÷åâñêîãî ð-íà., Õìåëüíèöêàÿ îáë.; LET — îáíàæåíèå âîçëå Ëåòè÷åâà, Õìåëüíèöêàÿ îáë.;
SAT — îáíàæåíèå âîçëå ã. Ñàòàíîâ, Õìåëüíèöêàÿ îáë.; IZA — áåðåãîâîé îáðûâ ð. Ãîðûíü â îêð. ã. Èçÿ-
ñëàâ, Õìåëüíèöêàÿ îáë.; A — ìåñòî îáíàðóæåíèÿ Coelacanthia quadrispinosa Àíäðóñîâûì (1890) íà
Êåð÷åíñêîì ï-îâå — òèïîâîå ìåñòîíàõîæäåíèå.
Results and discussion
The genus Mohrensternia was erected by Stoliczka (1868) for a few Miocene ris-
soid species (Rissoa angulata Eichwald, 1830, R. inflata Ho�rnes, 1856 and others),
which occur in the brackish or fresh-water deposits of central and Eastern Europe.
Since that time the number of species classified in Mohrensternia has increased signifi-
cantly (Andrusov, 1890, 1906; Zhizhchenko, 1936; Zhgenti, 1981) although some of
them were transferred later to morphologically similar genus Rissoa (Il’ina, 1979; Ba-
dzoshvili, 1991). The shell of Mohrensternia differs from Rissoa one by having relative-
ly thin walls and not thickened outer lip of aperture; the ornament consists of delicate
spiral threads and strong usually curved axial ribs.
The fossil record of Mohrensternia strongly suggests that the genus has evolved from
Rissoa as early as latest Oligocene/earliest Miocene (Andrusov, 1890; Zhizhchenko,
1936; Korobkov, 1955). In the middle part of Miocene Mohrensternia is already well
diversified. For instance the Tarkhanian and Chokrakian seas were inhabited by Mo-
hrensternia protogena (Andrusov, 1890), M. subprotogena Zhizhchenko, 1936, M. nitida
Zhizhchenko, 1936, M. pseudosarmatica Friedberg, 1923. In Karaganian Sea lived the
Mohrensternia grandis (Andrusov, 1890), M. barboti (Andrusov, 1890), M. subglobosa
Zhgenti, 1981, M. karaganica Zhgenti, 1981, M. gratiosa Zhizhchenko in Il’ina, 1993.
From Sarmatian deposits are known the Mohrensternia angulata (Eichwald, 1830),
M. inflata (Ho�rnes, 1856), M. hydrobioides Hilber, 1897, M. pseudoinflata Hilber, 1897.
At last the Maeotian Sea was inhabited by the Mohrensternia carinata (Andrusov, 1890),
M. subangulata (Andrusov, 1890) and M. subinflata (Andrusov, 1890). Actually these
are only our preliminary survey, because the exact faunal composition is still incom-
plete, and is a subject for discussions and contradictions.
The decrease of the salinity in the Ponto-Caspian basins forced Mohrensternia to
adapt these conditions. As a result the homeomorphic genera Archaschenia and Coela-
canthia have developed in two different time horizons (Karaganian and Maeotian
respectively) independently.
The question of affinity and distinction between Mohrensternia and Rissoa, Mo-
hrensternia and Archaschenia, and also Mohrensternia and Coelacanthia were widely dis-
cussed in the literature (Stoliczka, 1868; Il’ina, 1972 a, b; Zhgenti, 1981, 1991; Badzo-
shvili, 1991). Zhgenti (1991) provided the hypothesis on the origin of Archaschenia
while phylogenetic relationships between Mohrensternia and Coelacanthia have been
postulated by Il’ina et al. (1976).
The monotypic genus Coelacanthia was established by Andrusov (1890) for C. qua-
drispinosa from Maeotian deposits of Kerch Peninsula. Later Andrusov (1906) repeat-
ed the original description of the species, which he supplemented by three drawings and
one photograph of the shells. Although these images are of rather poor quality they
allow nevertheless to note the main diagnostic characters. The shell of C. quadrispinosa
is characterized by very thin walls, conical shape and lacking both spiral and axial orna-
mentation. Apart from fine growth lines it bears also spine-like projections with a nar-
row slit along their posterior side (fig. 2, 3). Andrusov (1890, 1906) classified Coelacan-
thia with some doubts in the family Rissoidae and its systematic position is still not fully
understood (Wenz, 1938–1944; Starobogatov, 1970). In the original description of the
species as well as in the formal diagnosis of the genus, Andrusov (1890, 1906) has
emphasized the distinctiveness of the Coelacanthia sculpture elements and supposed
that the spines are the outer lip projections sealed by the following whorls.
C. quadrispinosa has spines that are highly modified axial ribs (such ribs are always
present at the Mohrensternia species), which were inflected into strongly stretched pro-
jections and not outer lip through projections (as e. g., in Typhis Montfort, 1810 or
Murex Linnaeus, 1758). They form a semiclosed cone narrowing towards its end (fig. 2,
3). Only the outer layers of the shell (i. g., periostracum and outer layer of ostracum)
5Phylogenetic relationships of Coelacanthia and Archaschenia...
were engaged in forming the spines (as in the forming of the axial ribs of Rissoa and
Mohrensternia). The aperture of adult C. quadrispinosa is simple, holostomous without
any channel extended into the spine as it was mentioned in the original description of
Andrusov (1890).
All these facts strongly suggest that Coelacanthia belongs to Rissoidae, not to
Hydrobiidae or Pyrgulidae (= Micromelaniidae) as it was suggested by some authors
(Wenz, 1938–1944; Starobogatov, 1970; Badzoshvili, 1991).
The fossil record suggests also that C. quadrispinosa has evolved from a species of
Maeotian Mohrensternia, most probably M. subinflata (Andrusov, 1890). The transfor-
mation of the Mohrensternia axial ribs into spines of Coelacanthia is quite simple from
the morphological point of view (see the reconstruction of those projections on fig. 3,
A1–A4). The collection of Andrusov, deposited at the Paleontological Institute of
Russian Academy of Sciences (Moscow) consists also of some intermediate forms
(specimens were collected from the same strata) showing the continuous transforma-
tion M. subinflata into C. quadrispinosa (fig. 3, B–D). It is clearly visible, that the inter-
mediate forms differ from typical smooth Coelacanthia by its wider and thickened shell
ornamented with fine spiral threads and remnants of axial ribs (fig. 2, 3). This orna-
mentation is similar to the one present among M. subinflata specimens but the presence
of the spines and more slender shell shape are typical of C. quadrispinosa. Although the
intermediate forms between M. subinflata and C. quadrispinosa are known nevertheless
the transformation was so fast, that it might be considered as “instantaneous”. M. subin-
flata, when evolved towards C. quadrispinosa, lost the axial and spiral sculpture but
acquired spines and thinner-walled shell instead.
These transformations seem to be connected with a rapid change of habitat, which
probably had an impact on their mode of life. Mohrensternia most probably lived in sea
grass meadows similarly to many modern species of the Rissoa (Fretter, Graham, 1963;
Anistratenko, Stadnichenko, 1995). The long and fragile spines cannot be regarded as
adaptive to such conditions. The thin-walled shells and spines of Coelacanthia suggest
that the mollusks inhabited most likely a rather soft substratum (Il’ina, 1979).
A similar scenario as with Coelacanthia led probably to the development of
Archaschenia merklini Zhgenti, 1981 and Archaschenia iljinae Zhgenti, 1981, the
Karaganian rissoids of the Crimean-Caucasian region of the Paratethys. These gas-
tropods are known only from the Varnensky substage of the Karaganian and, accord-
ing to Zhgenti (1991), are derived from Mohrensternia barboti (Andrusov, 1890). Also in
this case a sequence of intermediate forms (from the same strata) is known (fig. 3, E–G).
6 V. V. Anistratenko
Fig. 2. Shell of paralectotype of Coelacanthia quadrispinosa in different position showed the characteristics of
sculpture (A1–A3) and base part of a broken spine (A4).
Ðèñ. 2. Ðàêîâèíà ïàðàëåêòîòèïà Coelacanthia quadrispinosa â ðàçëè÷íûõ ïîëîæåíèÿõ; ïîêàçàíû îñîáåí-
íîñòè ñêóëüïòóðû (A1–A3) è áàçàëüíîãî ó÷àñòêà îòëîìèâøåãîñÿ øèïà (A4).
Shell of Archaschenia differs from Coelacanthia one practically only by lower spire and
having an extremely fragile periostracal plate that covers the slit which is along the
spine-like projections.
An much harder task is to solve the problem of demarcation between Mohrensternia
and Mohrensternia-derived Archaschenia and Coelacanthia. The fossil record of both
7Phylogenetic relationships of Coelacanthia and Archaschenia...
Fig. 3. The sculptural elements of Coelacanthia quadrispinosa (A1–A4) and reconstructions of the morphological
lineage Mohrensternia subinflata (B) — Coelacanthia quadrispinosa (C, D1, D2) and Mohrensternia barboti (E) —
Archaschenia merklini (F, G). B–D after Il’ina et al., 1976, modified; E–G after Zhgenti, 1991, modified.
Ðèñ. 3. Ýëåìåíòû ñêóëüïòóðû Coelacanthia quadrispinosa (A1–A4) è ðåêîíñòðóêöèè ìîðôîëîãî-
ýâîëþöèîííûõ ðÿäîâ Mohrensternia subinflata (B) — Coelacanthia quadrispinosa (C, D1, D2)) è Mohren-
sternia barboti (E) — Archaschenia merklini (F, G). B–D ïî Èëüèíîé è äð., 1976, ñ èçìåíåíèÿìè; E–G
ïî Æãåíòè, 1991, ñ èçìåíåíèÿìè.
cases is complete enough to trace continuous lineages. The evolution of these gas-
tropods may be divided into four stages as follows.
1. Typical Mohrensternia ornamented with strong axial ribs.
2. Shells with sinusoidal and relatively weaker but wider ribs.
3. Shells with clamp-shaped ribs, which express a tendency to form high half-
moon-shaped scales. Both features (clamp-shaped ribs and sinusoidal ribs) can be noted
sometimes even on the same shell.
4. Shells with both half-moon-shaped scales and spines typical of Archaschenia and
Coelacanthia.
The shells, which have axial ribs with half-moon-shaped scales, even low and not
inflected into incipient spines, have to be excluded from Mohrensternia. Spines and
scales as modifications of the sculpture are typical of Archaschenia and Coelacanthia and
gastropods with such features should be classified in those genera.
8 V. V. Anistratenko
Fig. 4. The hypothetical scheme of the phylogenetic relationships of some Miocene rissoids.
Ðèñ. 4. Ïðåäïîëàãàåìàÿ ñõåìà ôèëîãåíåòè÷åñêèõ îòíîøåíèé íåêîòîðûõ ìèîöåíîâûõ ðèññîèä.
In result Archaschenia and Coelacanthia although strikingly similar are only hete-
rochronous homeomorphic taxa developed in two different time horizons from two dif-
ferent species of Mohrensternia, which immigrated repeatedly from the Mediterranean
into the Ponto-Caspian area of the Eastern Paratethys (Anistratenko, 2001). Thus the
position of these genera among Rissoidae is doubtless. The hypothetical scheme of the
phylogenetic relationships of mentioned taxa is presented in figure 4.
The Rissoidae in broad sense is a conglomerate of families, which are united into
the superfamily Rissooidea (Ponder, 1985; Anistratenko, Starobogatov, 1994). Karaga-
nian Archaschenia, Maeotian Coelacanthia and also their ancestor group Mohrensternia
should be considered as separate genera. Taking into consideration mainly the unusual
characteristics of the adult shell (namely lacking of spiral and axial ornamentation and
bearing the spine-like projections instead of), the level of their distinctiveness was
recently evaluated to merit subfamily rank (Anistratenko, 2003). This decision has been
made in a similar way as the taxonomic treatment of the recent Rissoiformes, based on
a complex evaluation of conchological, anatomical and other characters (Golikov,
Starobogatov, 1975, 1989; Ponder, 1985; Anistratenko, Starobogatov, 1994; Anistraten-
ko, Stadnichenko, 1995).
Homeomorphy, so common in different groups of gastropods, may be present
among species of the same or closely related genera but commonly also different gen-
era (e. g. Archaschenia and Coelacanthia) or even families. The latter has been exem-
plified by Anistratenko (1998). It was shown that species of the recent genera Haurakia
Iredale, 1915 and Mutiturboella Nordsieck, 1972 (Haurakiidae) are very similar con-
chologically to Rissoa and Benzia Nordsieck, 1972 (Rissoidae); Thalassobia Bourguignat
in Mabille, 1877 (Cochliopidae Tryon, 1866 [syn. Littoridinidae Thiele, 1928]) is anal-
ogous to Hydrobia Hartmann, 1821 (Hydrobiidae). Anatomically all these pairs of gen-
era differ enough to place them in easy distinguishable families — more detail see
Ponder (1985) and Anistratenko, Stadnichenko (1995).
When the higher level taxonomic units of any group are defined it is important to
follow certain principles. One of the most important, the principle of a uniform level
of taxonomical distinctiveness, requires that all taxa of the same rank have to be based
on the same level of distinctiveness (Mayr, 1971; Golikov, Starobogatov, 1989).
Morphological characters of three reviewed genera were considered as sufficiently dis-
tinct to place them in three different monotypic subfamilies (Mohrensterniinae
Korobkov, 1955, Coelacanthiinae Anistratenko, 2003 and Archascheniinae Zhgenti,
1991) and to unite them in the separate family Mohrensterniidae Korobkov, 1955 with-
in Rissooidea (Anistratenko, 2003) (see Appendix).
Some data on protoconch characters of Sarmatian Rissoa and Mohrensternia from
West Ukraine (author’s collection) were recently obtained using the SEM. These are
unpublished data obtained in summer 2003 jointly with Dr. Andrzej Kaim (Institute of
Paleobiology, Polish Academy of Sciences, Warsaw). Unfortunately, the studied shells
condition was not good, and the SEM micrographs are not proper enough for publica-
tion. Nevertheless, the main characters of protoconch and teleoconch both of Rissoa
and Mohrensternia could be evaluated from those SEM micrographs. They suggest to
change the taxonomy significantly. The detailed documentation of Sarmatian Mohren-
sternia and Rissoa from West Ukraine shows the following: 1) the structure and char-
acteristics of Rissoa and Mohrensternia protoconchs are quite similar i. e., both genera
have inflated, rounded protoconchs sculptured only by a few (2–4) fine spiral threads;
2) among the Sarmatian Mohrensternia at least a few distinct species (M. angulata,
M. inflata, etc.), which lived sympatrically, could be recognized. They are similar on
protoconch characters, but distinguished enough on shell geometry, i. e., Raup’s para-
meters (Raup, 1966).
The taxonomically significant differences between Mohrensternia and Rissoa are: in
Mohrensternia the shell is relatively thinner; the outer lip is not thickened, and is slight-
9Phylogenetic relationships of Coelacanthia and Archaschenia...
ly curved and lengthened on the base. Also the axial ribs of Mohrensternia express the
tendency to form high half-moon-shaped scales. Archaschenia as well Coelacanthia usu-
ally lack the thickness of the outer lip which Rissoa has.
Thus the significant morphological similarities suggest a close relationship between
Mohrensternia and Rissoa. The peculiarities of Archaschenia and Coelacanthia suggest
their distinctiveness, although they are definitely connected with Mohrensternia.
Unfortunately we have no specimen of Coelacanthia or Archaschenia preserved well
enough to study the protoconch structure. Therefore our proposed taxonomy is still
quite provisional (Anistratenko, 2003). It follows from the preceding taxonomic inter-
pretation that the separation of Mohrensternia, Archaschenia and Coelacanthia into three
separate monogeneric subfamilies as well as establishing the family Mohrensterniidae
was a premature decision (Anistratenko, 2003). Data available suggest that all men-
tioned taxa (Rissoa, Mohrensternia, Archaschenia and Coelacanthia) should be consid-
ered as independent genera within the family Rissoidae.
Conclusion
1. The species of Archaschenia lived in the Eastern Paratethys during the Karaganian,
whereas the species of Coelacanthia lived only in the Early Maeotian. These taxa have a
very similar morphology, with the resemblance of their sculpture elements being most
conspicuous. The sculptural features are undoubtedly homologous, and they were gener-
ated independently at different times. The morphological similarity is so strong, that some
authors (Badzoshvili, 1986) regard them as species of one genus or even as one species.
2. Although Archaschenia and Coelacanthia are not directly related, they have com-
mon ancestors in the Mohrensternia lineage. These taxa have evolved from the different
species of Mohrensternia, which is known to have immigrated into the Ponto-Caspian
area of the Eastern Paratethys from the Mediterranean.
3. Archaschenia has evolved from Mohrensternia barboti or a related species during
the Karaganian, whereas Coelacanthia quadrispinosa evolved from Mohrensternia subin-
flata during the Maeotian. The homeomorphic similarity of those taxa was due to con-
vergence in development of homologous structures because of similar ecological pres-
sures related to episodes of paleogeographic isolation and lowering of the salinity of the
Eastern Paratethys sea.
4. All taxa mentioned (Rissoa, Mohrensternia, Archaschenia and Coelacanthia)
should be considered as separate genera within the family Rissoidae.
I wish to thank to Dr. L. B. Il’ina (Paleontological Institute RAS, Moscow) for the donation of three
specimens of Coelacanthia quadrispinosa. I am greatly indebted to Dr. Andrzej Kaim (Institute of Paleobio-
logy PAS, Warsaw) for making the SEM micrographs and for his effort to improve this manuscript. Dietrich
Kadolsky (Great Britain) kindly helped in the proof-reading of the manuscript and provided helpful
suggestions, for which I am very grateful.
The investigation was partly supported by the Paleontological Society International Research
Program — Sepkoski Grants 2002 (Grant RGO — 1337-XX-02). My visit to Warsaw in 2003 was possible
due to financial support of both the Schmalhausen Institute of Zoology NAS Ukraine (Kyiv) and Institute
of Paleobiology, Polish Academy of Sciences (Warsaw).
Appendix
The systematics of Mohrensternia, Coelacanthia and Archaschenia recently reviewed (Anistratenko, 2003). As
this paper was published in Russian, the taxonomic proposals are summarized here in English.
Superfamily Rissooidea Gray, 1847
Family Mohrensterniidae Korobkov, 1955
Typ e g enu s: Mohrensternia Stoliczka, 1868
D i a gno s i s. Shell small, oval to ovate or conical. The ornament consists of fine spiral threads and commonly
strong axial ribs. In some cases spine-like sculptural projections with a narrow slit along their posterior side
are present. The aperture is rounded, holostomous. Umbilicus is absent.
10 V. V. Anistratenko
Di s t r i b u t i o n. Miocene (and Pliocene?) of Europe.
Con t en t. The family includes three subfamilies.
Subfamily Mohrensterniinae Korobkov, 1955
D i a gno s i s. Shell with relatively thin walls, the outer lip not thickened. The ornament consists of delicate
spiral threads and strong usually curved axial ribs.
D i s t r i b u t i o n. Miocene (and Pliocene?) of Europe.
Con t en t. Monogeneric.
Genus Mohrensternia Stoliczka, 1868
Typ e s p e c i e s: Rissoa angulata Eichwald, 1830, by subsequent designation by Nevill, 1885.
D i a gno s i s. As for the subfamily.
D i s t r i b u t i o n. Miocene (and Pliocene?) of Europe.
Con t en t. Approximately 15–20 species.
Subfamily Coelacanthiinae Anistratenko, 2003
Typ e g enu s: Coelacanthia Andrussov, 1890, by original designation.
D i a gno s i s. Shell very thin-walled. The shell usually lacks spiral and axial ornamentation apart from fine
growth lines and spine-like projections with narrow slit along their posterior side.
D i s t r i b u t i o n. Miocene (Maeotian) of Eastern Europe.
Con t en t. Monogeneric.
Genus Coelacanthia Andrusov, 1890
Typ e s p e c i e s: Coelacanthia quadrispinosa Andrussov, 1890, by monotypy.
D i a gno s i s. As for the subfamily.
D i s t r i b u t i o n. Miocene (Maeotian) of Eastern Europe.
Con t en t. Only the type species.
Subfamily Archascheniinae Zhgenti, 1991
(proposed as tribe Archascheniini Zhgenti, 1991)
Typ e g enu s: Archaschenia Zhgenti, 1981.
D i a gno s i s. Shell thin-walled, conical in shape. Spiral and axial ornamentation are absent. The surface
usually bears spine-like projections with narrow slit along their posterior side. The slit is covered by an
extremely fragile periostracal plate.
D i s t r i b u t i o n. (Middle Miocene) Karaganian of East Europe.
Con t en t. Monogeneric.
Genus Archaschenia Zhgenti, 1981
Typ e s p e c i e s: Archaschenia merklini Zhgenti, 1981, by original designation.
D i a gno s i s. As for the subfamily.
D i s t r i b u t i o n. Middle Miocene (Karaganian, Varnensky horizon) of Kerch Peninsula and Transcaucasia.
Con t en t. Two species, type and A. iljinae Zhgenti, 1981.
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