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A new record of Crenobia alpina (Dana, 1776) (Platyhelminthes, Tricladida) in Romania
Author: Anda-Felicia Babalean
University of Craiova, Faculty of Horticulture, Department of Biology and Environmental Engineering, 13 A.I. Cuza Street, 200583, Craiova, Romania. email@example.com
In this paper, it is recorded for the first time the presence of Crenobia alpina in Parâng Mountains – the Southern Carpathians. Crenobia alpina populations have been sampled in the summer of 2016 and 2017 from a small area, of about 9 km2, between N 45° (17ʹ - 19ʹ) – E 23° (40ʹ - 41ʹ). In the studied area, Crenobia alpina is represented by both the monopharyngeal Crenobia alpina alpina (Dana) and the polypharyngeal Crenobia alpina montenegrina (Mrazek). The two forms appear to be spatially isolated by the biotope: the polypharyngeal C. a. montenegrina was found in springs (the place of emergence of the water from the ground) and the spring-fed streams, while the monopharyngeal C. a. alpina was found in streams with a higher flow.
Key words: Crenobia alpina alpina, Crenobia alpina montenegrina, Parâng Mts., springs, spring-fed streams, ecological isolation?
Introduction and historical background
Parâng Mountains are composed of certain old mountains consisting of crystalline and granitic schists on which different types of acidic soils with pH between 3.8 and 4.5 develop (Buia et al., 1961/1962). The maximum altitude is reached by Parângu Mare Peak - 2518 m (The Geographic Atlas of the World, 1992).
The vegetation of Parâng Mts., as well as the one of the entire Carpathian chain, presents several levels imposed by altitude: the nemoral level (Fagus), the boreal level (Picea, Abies), the subalpine level (juniper trees, rare spruce trees) and the alpine level (alpine meadows and dwarf shrubs) (Drugescu 1994). The soils of Parâng Mts., develop small marshy areas with a vegetation characteristic to peat acid soil: Eriophorum latifolium, E. vaginatum, Dactylorhiza maculata, Carex spp., Sphagnum spp., Pohlia elongata, Philonotis fontana. The vegetation in the proximity of certain springs is characterized by the presence of the insectivorous plant Pinguicula vulgaris. The species Oxalis acetosella is frequently met within Parâng Mts., being considered a good indicator of soil acidity.
In Romania, Crenobia alpina is represented by two subspecies the copulatory apparatus of which is identical, but they are different in terms of the number of pharynxes: Crenobia alpina alpina – the monopharyngeal form and Crenobia alpina montenegrina – the polypharyngeal form. Data regarding the geographical distribution, age and origin of the two subspecies within the Romanian Carpathians are brought by Codreanu (1956), Năstăsescu (1973, 1974), Botoșăneanu (1977).
Geographical distribution in Romania
Crenobia alpina (Crenobia alpina alpina): Mehedinți Mts., Retezat Mts., Bucegi Mts., the vicinity of Piatra Craiului Mts., Ciucaș Mts., Vrancea Mts., Godeanu Mts., Călimani Mts., Muntele Mic Mts., the Lotru River basin, the Jiu River basin, the Olt River basin, the Olteț River basin, the Siret River basin, the Bistrița Moldovenească River basin, the Bistrița Valley.
Crenobia alpina montenegrina: Mehedinți Mts., Retezat Mts., the vicinity of Piatra Craiului Mts., Bucegi Mts., Gârbova Mts., the Olt River basin, the Olteț River basin, the Jiu River basin.
With regard to the origin and age of the two subspecies, different authors formulated and brought arguments for different hypotheses:
Crenobia alpina (monopharyngeal) is supposed to be:
1. glacial relict - argument: cold-water stenothermic species (Codreanu 1956; Brandle et al. 2007 and included reference)
2. Tertiary, pre-glacial element – arguments:
a. the palearctic distribution (Codreanu 1956 citing Chappuis 1927 and Thienemann 1950)
b. its presence in Anatolia (Codreanu 1956 citing Kosswig and Recai 1950)
The Tertiary age is also supported by recent molecular studies (Brandle et al. 2007), which bring strong evidence that “Crenobia alpina is a superspecies, a complex of distinct lineages or cryptic species that date back to the late Miocene”.
In case of Crenobia alpina montenegrina, the Balkan Peninsula is considered the “incontestable genetic centre”, the Southern Carpathians representing the northern limit of the subspecies areal (Codreanu 1956). The age of Crenobia alpina montenegrina is supposed to be Tertiary (Neogene) (Codreanu 1956 citing Beklemișev 1923 and Stanković 1935). In 1927, Chappuis gave an even more exact temporal and spatial dating for the polypharyngeal form, namely the older Tertiary - Pliocene – the Southern Carpathians from Romania, arguing that “before this age, Cerna Mountains were separated from the Aegean land by a branch of the Sarmatian Sea” (Codreanu 1956, p. 393).
Codreanu (1956) considered that the polypharyngeal form is not the result of “a repeated and recent process of polytopic genesis” of the monopharyngeal form.
Material and methods
The material was collected from typical reocrene springs, reocrene springs with accumulation reservoirs with sandy bottom, spring-fed streams and slope streams with higher flow from the mountains belonging to the secondary range of Parâng Mts., between the Gilort and the Galbenul Valleys: Dâlbanul, Rânca and Tidvele Mountains. The springs and streams are located in the upper part of the boreal zone, at the contact with the subalpine zone (1443 – 1841 m altitude), on routes starting from the Botanical Chalet “Marin Păun” in Rânca Tourist Resort:
1. Botanical Chalet to the Romanul Stream – the 1st of July 2016; the 27th of June 2017
- typical reocrene springs and spring water collecting pipe – Fig. 1a, b (collecting, un-separated samples) – N 45º 17′ 781″, E 23º 40′ 666″, alt. 1443 m.; the 27th of June, 2017 (only observations).
2. Botanical Chalet to the Dâlbanul Stream (Paltinul Mt.) – the 2nd of July 2016
a) reocrene spring with sandy bottom accumulation reservoir, located within a marshy area (Pinguicula vulgaris present) – Fig. 1c – N 45° 19′ 136″, E 23° 40′ 136″, alt.1706 m.
b) stream at the foot of Paltinul (Mohorul) Mt. – N 45° 19′ 457″, E 23° 40′ 545″, alt. 1774 m.
3. The Tidvele Stream – the 10th of July 2016
a) Tidvele reocrene spring – (Pinguicula vulgaris present) – Fig. 1d, e – N 45°19′ 444″, E 23° 41′ 751″, alt. 1841 m.
b) spring-fed stream resulting from Tidvele spring – N 45° 19′ 249″, E 23° 41′ 774″, alt. 1737 m.
c) spring-fed stream, the same as the one mentioned above, but at a lower altitude – alt. 1700 m.
4. Tidvele Stream – marshy area – the 28th of June 2017
a) artificial reservoir with a drainage pipe (cross) – N 45° 18′ 595″, E 23° 41′58″, alt. 1670 m.,
b) two reocrene springs with sandy bottom accumulation reservoirs – Fig. 1f and
c) the resulting streams at about 20 m from the spring.
5. stream with a higher flow in the skilift area – the 28th of June 2017; N 45° 18′ 834″, E 23° 41′ 109″, alt.1660 m.
The water temperature at the sampling sites varied between 6º and 12º C in different moments of the day – 9 a.m. – 1 p.m.
The collected planarians were fixed with the Beauchamp fixator for 12 + 24 hours or 24 hours and preserved in 75 ° ethyl alcohol.
Fig. 1a – reocrene spring on the route Botanical Chalet – Romanul Stream
Fig. 1b - water collecting pipe on the route Botanical Chalet – Romanul Stream
Fig. 1c – spring-fed stream within a marshy area on the route Botanical Chalet – Dâlbanul Stream
Fig. 1d –Tidvele reocrene spring
Fig. 1e – Pinguicula vulgaris
Fig. 1f – spring with sandy bottom reservoir within Tidvele marshy area
146 monopharyngeal and polypharyngeal specimens of Crenobia alpina were collected. Their distribution on sampling sites, morphological features and biocoenosis are further rendered:
1. Botanical Chalet to the Romanul Stream – the 1st of July 2016; the 27th of June 2017
- reocrene springs and spring water collecting pipe, alt. 1443 m.: 31 specimens of which 28 polypharyngeal (7 – 12 pharynxes) – Fig. 2b and 3 monopharyngeal. Most specimens have visible copulatory area and 2 eyes; very few specimens with 1, 3 and 4 eyes.
Biocoenosis – Gammaridae, Plecoptera larvae and adults, Trichoptera larvae – Fig. 3a, b, c.
- the 27th of June 2017: uncollected specimens, observed in the same springs but with a smaller flow compared to the previous year.
Biocoenosis – Gammaridae.
2. Botanical Chalet to the Dâlbanul Stream – the 2nd of July 2016
a) spring (with Pinguicula vulgaris), alt. 1706 m.: 23 polypharyngeal specimens (7 – 12 pharynxes); the copulatory area and the female gonads (ovaries and vitelogenae) visible in most specimens. Three specimens show ocular asymmetry – 3, 4, respectively 5 eyes.
b) stream at the foot of Paltinul Mt. (Mohorul), alt. 1774 m.: 12 monopharyngeal specimens – Fig. 2c with visible copulatory area and female gonads. One specimen presents ocular asymmetry – 2+1.
Biocoenosis: Ephemeridae larvae, Plecoptera larvae, Trichoptera larvae – Fig. 3d.
3. The Tidvele Stream – the 10th of July 2016
a) Tidvele spring, with Pinguicula vulgaris, alt. 1841 m.: 16 polypharyngeal specimens (7 – 11 pharynxes) with visible copulatory area; the female gonads visible in some specimens.
b) spring-fed stream resulted from Tidvele spring, alt. 1737 m.: 6 polypharyngeal specimens (9 – 11 pharynxes) with visible copulatory area; the female gonads visible in some specimens.
c) spring-fed stream – alt. 1700 m.: 9 polypharyngeal specimens.
4. Tidvele Stream – marshy area – the 28th of June 2017
a) artificial reservoir with drainage pipe (cross), alt. 1670 m: no specimen.
b) two springs: 26 polypharyngeal specimens (9 – 12 pharynxes), most of them with visible copulatory area; the female gonads visible in some specimens.
c) 20 m downstream the springs: 20 polypharyngeal specimens (7 – 11 pharynxes), most of them with visible copulatory area; the female gonads visible in some specimens. Three specimens with 3, 5 and 9 exerted pharynxes – Fig. 2a, e.
Biocoenosis – Gammaridae, Trichoptera, Gordiidae (Gordiidae only in the area of the artificial reservoir with drainage pipe) - Fig. 3e, f, g, h.
5. stream – skilift area – the 28th of June 2017, alt. 1660 m.
- 3 monopharyngeal specimens – Fig. 2d.
The body length of the specimens varied between 6 mm and 15 mm in fixed animals. Body colour ranges from light grey with pink shades to blackish grey in living animals – Fig. 2b – and from yellow-grey to dark grey in fixed animals.
Fig. 2a – Crenobia alpina montenegrina (polypharyngeal) from a spring within a marshy area
Fig. 2b – Crenobia alpina montenegrina (polypharyngeal) –specimens collected from a reocrene spring on the route Botanical Chalet – Romanul; living animals in a glass crystallizer
Fig. 2c – Crenobia alpina alpina (monopharyngeal) – specimen collected from a brook located at the foot of Paltinul Mt.; living animal in a glass crystallizer
Fig. 2d - Crenobia alpina alpina (monopharyngeal) –specimen collected from a brook – skilift area; fixed animal
Fig. 2e – Crenobia alpina montenegrina – specimen collected from a reocrene spring with sandy bottom reservoir within Tidvele marshy area; fixed animal with 9 exerted pharynges
Fig. 3a – Trichoptera larva – reocrene spring biocoenosis on the route Botanical Chalet – Romanul
Fig. 3b – Plecoptera – reocrene spring biocoenosis on the route Botanical Chalet – Romanul
Fig. 3c – Plecoptera larva – reocrene spring biocoenosis on the route Botanical Chalet – Romanul
Fig. 3d – Trichoptera larvae – stream biocoenosis on the route Botanical Chalet – Dâlbanul
Fig. 3e – gammarid – spring biocoenosis within the marshy area of Tidvele spring
Fig. 3f – Trichoptera larva – spring biocoenosis within the marshy area of Tidvele spring
Fig. 3g – Trichoptera larva – spring biocoenosis within the marshy area of Tidvele spring
Fig. 3h – Gordiid – spring biocoenosis artificial reservoir, Tidvele stream - marshy area
The present study points the presence of monopharyngeal and polypharyngeal Crenobia within a mountain area composed of crystalline schists and with acid soil at altitudes between 1443 m and 1841 m.
Within the area and during the period the study took place, the monopharyngeal and polypharyngeal populations of Crenobia alpina seemed to be spatially isolated by biotope, excepting the sampling site nr. 1 (Botanical chalet to the Romanul Stream). The co-existence (mixture) of populations at this point should be questioned because the samples were not separated when collected from the springs and the collecting pipe.
The polypharyngeal populations occupy the area of springs – at the site of water emergence for typical reocrene springs and the reservoirs for reocrene springs with sandy bottom reservoirs. The spring-fed streams at a distance of approx. 20 m from the spring are populated by polypharyngeal Crenobia.
The monopharyngeal populations occupy the area of the streams with higher flow.
The separation of the two types of populations is correlated with the biotope (spring, greater flow stream) and does not appear to be influenced by altitude. From the altitudinal point of view, the monopharyngeal population from the point 2b) – stream, altitude 1774 m, is located between the altitudes occupied by polypharyngeal populations, for example between 1706 m altitude (spring from point 2.a)) and 1841 m (Tidvele spring from point 3a).
These data contradict the data provided by Codreanu (1956). The author points out two aspects for the collecting sites from the Southern Carpathians: 1) the "constant mixture of polypharyngeal and monopharyngeal Crenobia populations" and 2) "the mutual distribution of the two forms in the Prahova Valley: the polypharyngeal Crenobia do not exceed the low springs, up to 900 m, while the monopharyngeal Crenobia populate the springs of the whole alpine domain up to the plateau "(Codreanu 1956, p. 392). Codreanu considers as a possible means of reproductive isolation of the two subspecies (!) the different reproduction period. This seems to be all year long at the monopharyngeal Crenobia and limited to the interval September-October till January and perhaps at the beginning of summer for the polypharyngeal Crenobia (Codreanu 1956).
Wright determined in N-Wales that 2 factors are involved in the distribution of Crenobia alpina – the gradient of the stream and inter-specific competition with Polycelis felina, while other parameters do not appear to be important – temperature, water chemistry, predation and parasitism (Lock and Reynoldson 1976).
Codreanu states that, in the process of biogeographical evolution, "Polypharyngeal Crenobia has succeeded in almost completely replacing the monopharyngeal Crenobia" (Codreanu 1956, p. 392). Since the mixture of the two types of populations in the Southern Carpathians was found by Codreanu about 70 years ago it is possible that in Parâng Mountains we witness a going-on process of separation of the two types of populations due to the competition between them, competition in which polypharingy might represent an advantage. The possible adaptive role of polypharingy is discussed in some marine flatworms (Curini-Galletti & Cannon 1997).
Taking into account the conclusions of the study by Brandle et al. (2007) for Central Europe, namely "Crenobia alpina is a complex of distinct lineages or cryptic species that date back to the late Miocene", two considerations derive for the Crenobia fauna of the Romanian Carpathians (and not only):
1. Monopharyngeal and polypharyngeal populations should not necessarily be regarded as subspecies. They should be rather regarded as distinct species, namely twin species, morphologically identical at the level of the copulatory apparatus, and reproductively isolated by other means – maybe an ecological isolation by different biotopes.
2. As during the Miocene the Romanian Carpathians were barely rising (Drugescu 1994), it remains to investigate the biogeographical history of the lineages of the Carpathian Mountains. Such an investigation should include a long-term and on-whole areal monitoring of the presence, reproduction biology and food spectrum for the two forms.
A. F. Babalean is deeply grateful to Prof. Dr. Maria Năstăsescu for kindly provided the doctoral thesis for consultation.
A. F. Babalean also acknowledges the help of Dr. Daniel Răduțoiu and the students of the Faculty of Horticulture Craiova –Department of Biology and Environmental Engineering in the field work, Aurel Babalean for preparing the images (figures) and Dr. Alina Vlăduț for the English translation of a former draft of the manuscript.
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Babalean, A. (2017). A new record of Crenobia alpina (Dana, 1776) (Platyhelminthes, Tricladida) in Romania. PHILICA.COM Article number 1107.