regeneration patterns of natural populations of herbaceous perennials in spruce forests of the...

ISSN 1995�4255, Contemporary Problems of Ecology, 2013, Vol. 6, No. 4, pp. 402–407. © Pleiades Publishing, Ltd., 2013.Original Russian Text © G.G. Zhilyaev, 2013, published in Sibirskii Ekologicheskii Zhurnal, 2013, No. 4, pp. 535–542.

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Looking back at the history of population studies, itis clear that, until about the last third of the 20th cen�tury, populations were considered fairly simple deter�ministic systems with a regular pendulum dynamics oftheir elements. This meant that, after determining thetrends of population dynamics in relatively small timeintervals, one could predict their development for anyperspective.

Only after a scrupulous generalization of a vastarray of research results that were accumulated in thefield of population biology has it become clear thatsuch ideas about natural populations are either incon�sistent or exceptional [1–10]. The principle of lineardeterministic interdependence between their elementsis valid only for small fragments (population fractionsand remnants) left after the collapse of natural popu�lations [11]. This seems to be one of the reasons for theexisting inconsistency (conflict) between the researchgeneralizations of the basic biology and the results ofeconomic exploitation or environmental practices.

In the new paradigm, natural (natural–historical)populations are represented as supercomplex systemswith a multivariate relationship of their elements and athreshold principle of responding to the impact [12]. Itfollows that an adequate assessment of the state andforecast of development of the population, not interms of chance, requires an infinitely accuratedescription of the totality of their current states. Rec�ognition of the fact that natural populations have nocreator makes the task of finding universal elementswhich can enable population self�assembly clear.

Other than technical difficulties, its solution is com�plicated due to the increasing degradation of the natu�ral environment, when it is difficult to find natural–historical populations whose structure had not under�gone anthropogenic deformations and which can serveas adequate reference objects for such studies. Thisarticle presents the results of perennial registrations ofthe parametric dynamics and analysis of the self�regeneration patterns of the sample population of Sol�danella hungarica Simonk. in reserve conditions.

OBJECTS AND METHODS

The studies were carried out on permanent sampleplots of the Pozhizhevskii biological station, Instituteof Ecology of the Carpathians, National Academy ofSciences of Ukraine, assigned in the general contourof the Carpathian National Nature Park (CNNP).They were launched in 1974 and have not been inter�rupted to this day. The duration of registration on per�manent sample plots and objects is the only one of itskind in the world.

The studies were based on traditional methods ofpopulation analysis, which do not require furtherexplanation [13–20]. A natural–historical populationof Soldanella hungarica Simonk. was chosen as a sam�ple object.

In the Carpathians, in the conservation area at aheight of 1340 m above sea level, we laid a transect of5 × 13 m (65 m2) with a corresponding number of ele�mentary squares, each 1 m2, in a typical section of

Regeneration Patterns of Natural Populations of Herbaceous Perennials in Spruce Forests of the Carpathians

G. G. ZhilyaevInstitute of Ecology of the Carpathians, National Academy of Sciences of Ukraine, ul. Kozel’nitskaya 4, Lviv, 79026 Ukraine

e�mail: [email protected]

Abstract—According to the results of the long�term (1974–2011) monitoring of permanent sample plots atthe Institute of Ecology of the Carpathians, National Academy of Sciences of Ukraine, located at the upperborder of spruce forests of the Carpathians, patterns of characteristic stages of self�regeneration (regenera�tion) of the subpopulation structure of Soldanella hungarica Simonk. after severe local damage are general�ized. It is found that, under experimental conditions, these point exposures do not cause the general destabi�lization of population processes. However, they initiate behavioral responses for the mobilization of the pop�ulation reserve of S. hungarica on nearby areas. The varying role of individual and group effects in theseprocesses is revealed. The conclusion of the decisive importance of the vitality composition for the results oflocal regeneration of the S. hungarica population is made. It is stated that the self�regeneration of the S. hun�garica structure on the experimental plots is a time�consuming process that requires several decades.

Keywords: population, subpopulation locus, vitality, vital state, viability, ontogeny

DOI: 10.1134/S199542551304015X

CONTEMPORARY PROBLEMS OF ECOLOGY Vol. 6 No. 4 2013

REGENERATION PATTERNS OF NATURAL POPULATIONS 403

blueberry (Piceetummyrtillosum) spruce forest (Fig. 1).It had been studied previously in terms of standarddemographic variables, age, and vitality composition;all the specimens of S. hungarica were marked andtheir spatial position was mapped. Then, in 1975, infive central squares of the transect (type T�1), all theplants of S. hungarica were excavated. As a result, eachof them (C�III, C�V, C�VII, C�IX, and C�XI) was sur�rounded by undamaged plots of four types: T�2 (C�V,C�VI, C�VIII, and CX) contacting damaged plots onboth sides, T�3 (B�III, B�V, B�VII, B�IX, B�XI, C�II,C�XII, D�III, D�V, D�VII, D�IX, D�XI) contactingdamaged plots on one side, T�4 (B�II, B�IV, B�VI,B�VIII, B�X, B�XII, D�II, D�IV, D� VI, D�VIII, D�X,D�XII) contacting only at one or two corner points,and T�5 (from A�I to A�XIII, from E�I to E�XIII, B�I,C�I, D�I, B�XIII, C�XIII, D�XIII) not contacting theplots of T�1. These notations of types and the alpha�betic–numerical scheme of coordinates of specificplots are used in the text and figures of this paper.

In the percentage of the total transect area, the pro�portion of each type (T�1, T�2, T�3, T�4, and T�5) ofthe sample plots was 8, 6, 18, 18, and 50%, respec�tively. In addition to the external, territorially contin�uous type (T�5), the squares of the other types withinthe transect can be considered discrete, regular frag�ments. In general, the experiment imitates a typicalsituation of natural local damages of herbaceous tiersof the Carpathian spruce forests. This scheme allowedresearchers to organize the detailed perennial registra�tions on a relatively small area of the sample transectand to observe the character stages of local self�regen�eration of the S. hungarica population after experi�mental impacts.

The text and figures of this paper use a generallyaccepted age periodization [21] and ontogenic (age)notations: young growth (p), juvenile (j), immature(im), virginile (v), young generative (g1), mature gen�erative (g2), old generative (g3), subsenile (ss), andsenile (s); and vitality differentiation [22–23] of the

specimens: high (V�1), medium (V�2), and low (V�3)vitality.

RESULTS AND DISCUSSION

At the beginning of the experiment, in terms of thedynamics and the system stability, the population ofS. hungarica was qualified as definitive with a fluctua�tion period of 4–5 years. In terms of the ontogenic(age) structure, it was qualified as a normal middle�aged population with a maximum of mature generativespecimens (Fig. 2), which fits well within the range ofthe basic ontogenetic range of S. hungarica [24]. Interms of the ratio of vitality groups (V�1 : V�2 : V�3 =4.4 : 2.3 : 1), it is a thriving population [16, 25].

Due to the high content of reproductive (v�g3) indi�viduals, regular vegetative and seed reproductionoccurs in this population and the average number ofindividuals ranges from 101 to 125 pcs/m2, in accor�dance with periodicity of fluctuation waves and sea�sonal dynamics of germs. In addition, the populationhas an additional reproductive reserve in the form ofnumerous renewing buds, temporally nonflowering,dormant, and quasi�senile generative individuals. It isby their activation that the fluctuation processesoccurring in it are determined.

In terms of all the above, this population wasfounded as a reference one [24], where at each level ofvitality some basic variants of sinontogeny are imple�mented with a steady and regular change of genera�tions [26].

The experiments imitated the real situation of thepopulation field of S. hungarica in the blueberry

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Fig. 1. Scheme of experimental transect plots for the studyof the local regeneration of Soldanella hungarica Simonk.population in blueberry spruce forest.Types of plots: 1, T�1; 2, T�2; 3, T�3; 4, T�4; 5, T�5.Coordinate grid of plots is denoted by horizontal numeri�cal row I–XIII and vertical letter row A–E.

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Fig. 2. Changes in ontogenic ranges after local damage tothe Soldanella hungarica Simonk. population.

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spruce forest damaged by natural fall of trees or actionsof burrowing animals.

In 1975, before the start of the growing season, allthe specimens of S. hungarica were dug up andremoved from each of the squares (T�1) of thetransect. The response consisted of the fast (before theend of the current growing season) activation of gener�ative individuals (g1, g2, g3) on the adjacent (T�2, T�3,T�4) nondamaged plots of the transect. Depending onthe vitality (V�1, V�2, V�3) and ontogenetic status,before going under the snow, such specimens formed12–20% more buds than their statistical norm. Whilethe number of buds of vegetative regeneration did notchange as the main prerequisite for the activation ofvegetative propagation, most of the other individualtraits did not change at this point either.

However, starting in 1976, there is a quite clearactivation of other elements of the reproductivereserve. The fact that this process involves not onlyspecimens from the adjacent plot, but also from moreremote ones (T�5) which generally do not have com�mon borders with damaged ones (T�1) was interestingand quite unexpected. Although information on localdamages comes here with a delay of 1.5 or 2 years, itcauses the same activation of elements of seedrenewal. To some extent, these findings suggest a fairly

high speed and reliability of this information distribu�tion in the population field. However, because we donot yet have actual material for a discussion of thephysical principles and agents that transfer such infor�mation, we can only state the adequacy of response tosuch impacts on both the close and peripheral plots ofthe transect.

Though the indicators of the potential and actualseed productivity almost have not changed, the yieldof seeds of S. hungarica within the transect increasednearly sevenfold. Such an explosive dynamics was notso much a consequence of an increase in the level ofindividual indicators (percentage of fruit flowering,number of generative shoots per specimen, etc.) as itwas of the number of such effects of the group [27–28]like the massive activation of temporally nonflower�ing, dormant, and quasisenile specimens. Although atdifferent stages of recovery the level of group effectsmay change, their priority over the individual reac�tions is maintained (Fig. 3).

If in the first 4–5 years the reliability of seed repro�duction is provided by the maximum participation ofthe entire pool of the population reserve specimens,the situation changes after this period. Gradually theactivity of these animals decays, and they again returnto the standard periodicity in shifts of their active andpassive states. Accordingly, their contribution to theformation of the yield of seeds of the invasion flow tothe damaged area slows down. However, by this pointof time, the first genets, capable of supporting furtherregeneration in the damaged fragments of the S. hun�garica populations by themselves, had alreadyentrenched on these specimens and reached thereproductive state. The accelerated rates of develop�ment of such genets is a consequence of temporalmodifications of sinontogeny [29] and violations byimmature and virginile specimens of the stereotypedsequence in the change of their age states. As a result,already by the fifth or sixth year (rather than 15–20thyear, as is normal [26, 30–31]) there are young andmature generative specimens on the damaged plots,although in terms of many criteria the ontogenic rangestill retains features of the invasive one (see Fig. 2). Atthe same time, and in many ways as a result of theabovementioned changes, the total number of speci�mens grows rapidly. Fourteen or 15 years later (1990–1991) it reaches its maximum (225 psc/m2), whichnoticeably (by 1.8–2.2 times) exceeds the averagepopulation one. However, unlike the rest of the popu�lation, 90–97% of this local population consists of anextremely unstable pool of undergrowth: p, j, and im.

Such transformations are based on the properties ofspecimens with high vitality, with their inherent highrates of development and ability to quickly replenishgenerative groups, bypassing part of the age states [26].This is why, despite the significant reduction in theinvasive inflow of the seeds from neighboring areas, theregeneration of the population field continues, althoughto a greater extent due to the rapid increase of its own

Fig. 3. Changes in individual and group traits of Soldanellahungarica Simonk. at initial stages (1975–1980) of regen�eration of population loci. (A) level of normalized unit.Traits: (1) percentage of fruit�flowering (per individual),(2) number of generative shoots (per individual),(3) potential seed productivity (per individual), (4) realseed productivity (per individual), (5) number of regener�ation buds (per individual), (6) intensity of vegetativepropagation, (7) rates of vegetative overgrowth, (8) num�ber of genets, (9) number of actually flowering individuals,(10) number of temporary nonflowering individuals,(11) number of quasisenile individuals, (12) number ofdormant individuals, (13) seed yield, (14) number ofplantlets (on damaged plots), (15) intensity of seed germi�nation (on damaged plots), (16) death rate in undergrowthgroup (on damaged plots), and (17) speed of germs devel�opment till virginile state (on damaged plots).

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reproductive potential in the loci of T�1. In this period,the content of individuals with high vitality increases hereup to 85%. Though in the future this figure somewhatreduces, but even now (in 2011) specimens of high vital�ity dominate (79%) (Fig. 4). Similar vitality ranges aregenerally characteristic for the S. hungarica popula�tions existing under conditions of nonlimited specialresources [26].

Returning to the ontogenic vitality ranges, itshould be stated that having undergone a number oftransformations, even at the time of the last registra�tions (in 2011), they still have not fully returned totheir original state (Figs. 2, 4). At other comparableproportions, the absolute maximum in the range ofontogenic states currently falls on old rather thanmature generative specimens. In contrast to the initialstages of the experiment, one cannot explain it by thechanges in the basic ontogenic scheme [26]. It is pos�sible that the current situation in the loci of T�1 is justa result of the fact that their regeneration history afterdamages is not long enough and later all the differ�ences are leveled. However, now one can only statethat, after nearly 40 years, the local structure in thedamaged areas still noticeably differs from the struc�ture of the rest of the population.

The generalized picture of the local regeneration ofthe S. hungarica population can be represented as asequence of several stages:

The first response reactions at the adjacent plots(T�2,3,4) appear in the first year of the experiment.Since their speed, synchronicity, and amplitude do notshow dependence on the extension of common bor�ders with the territories (T�1), it is conceivable that theadequacy of the transfer of information on local dam�ages does not depend on it. At greater distances (plotsT�5), the response is delayed for another year or two.However, the fact that it is virtually identical to theresponse in the area of direct contact (T�2, 3, 4) indi�cates the reliability of its transmission channels.

In terms of its duration, the local enhancement ofthe reproductive activity of the undamaged plots of thetransect is proportional to the standard period of pop�ulation fluctuations of S. hungarica in the blueberryspruce forest. Subsequently, its level graduallydecreases to the norm. Since the vegetative mobility ofS. hungarica is weak from the very beginning and thelong�term seed bank is not formed in its population[26], the success of regeneration of the undamagedplots in these years is completely determined by theinvasive influx of the seeds. Under such circum�stances, the seeds of high vitality are able to use theiradvantage of the intensity of germination, survivallevel, and rate of seedling development. In the end,this leads to the absolute dominance of the V�1 frac�tion in the vitality range.

In this period there was no self�maintenance of thelocal clusters of T�1 as a process that can offer a con�tinuity of generational changes [11, 32]. The invasiveprinciple of regeneration still prevails there, which

promotes the preservation of the incomplete ontoge�nic composition of S. hungarica and the reduction ofthe role of the vitality groups V�2 and V�3.

After about 5–7 years (1980–1982), young andmature generative specimens that are replenishedfrom the virginile group start to appear on the dam�aged areas (see Fig. 2). Therefore, the reduction of theseed yield and intensity of their invasive influx fromthe neighboring territories (T�2�5) is partially com�pensated for by the yield of their own seeds on the plotsof T�1.

Subsequently, after 7–10 years, the principle ofinvasive regeneration is more often replaced with self�maintenance, although the number of specimens is40% higher than the average for the population. Sucha high number is explained by the fact that, unlike therest of the population, the plots of T�1 are dominatedin this period by genets, which initially exceed rametsin terms of their reproductive potential [26]. Thisredundancy of the demographic pressure probablymakes the greater part of the generative specimensleave it by entering an inactive state of temporarilynonflowering and dormant ones.

Interestingly, a similar deactivation of specimensalso spreads to the adjacent plots of T�2 and T�3.However, it fades at small distance (0.5–1 m) from theborders with regeneration plots of T�1. At this stage,the range of developmental states gains all the traits ofa normal one (left�sided with the absolute maximumon the virginile group). Because at this time on theT�1 plots there are still no specimens that managed tocompletely undergo their life cycle, there is also noreproductive (ss and s) group. However, the high num�ber of genets, which due to vegetative reproductionhave reached the state of numerous specimens (undi�vided units) [7, 29], creates an increasing deficit of free

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Fig. 4. Local changes in vitality composition after experi�mental damages in population of Soldanella hungaricaSimonk.

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ZHILYAEV

space. As a result, the undergrowth (p + j + im) ofgroup V�1 loses its advantages in survival and is par�tially replaced by undergrowth from groups V�2 andV�3. This leads to the redistribution of the vitalitycomposition in favor of more compact specimens ofV�2 and V�3 (see Fig. 3).

Because of the variability of the starting composi�tion in the particular local clusters of T�1, some differ�ences in the structural and functional characteristicsappear and persist. This somewhat blurs the beginningof the next stage of regeneration, which is difficult todate as definitively as the former. It starts approxi�mately after 18–20 years from the beginning of theexperiment. However, by the time of the last registra�tions (October 2011), the statistical result of all theparticular local transformations is leveled and seemsquite definite. However, currently, in terms of demo�graphic and vitality indicators, it is not quite identicalto the initial state of 1974 (see Figs. 2, 3).

The ontogenic range is right�sided, monocentric,temporarily not complete, and of a normal type. Incomparison to the rest of the population, it still hasalmost two times fewer V�2 (15–17%) and V�3 (3–7%) specimens, and the range of ontogenic states isdominated by the age group g3 rather than g2. There�fore, the current situation in the loci on the plots ofT�1 does not have a complete correspondence with theinitial statistical state which could be interpreted astheir final recovery.

We would like to note that this paper discusses onlyone of the typical situations of fragmented damages ofvegetative populations. There is no doubt a fundamen�tal difference between the patterns of local interactionin the experiment and the global behavior of popula�tions, and algorithms of their regeneration are unlikelyto be completely identical. Of course, they will be dif�ferent in species other than S. hungarica and in otherbiomes.

It can be stated that, although the effects of localdamages in the population of S. hungarica are in prin�ciple reversible, they level out only after a fairly longperiod of rehabilitation.

CONCLUSIONS

Our experiments make it possible to conclud that,after focal damages of the population field of S. hun�garica, the regeneration process is started due to theactivation of the population reserve of neighboringareas. It forms an invasive seed flow, the parameters ofwhich are regulated by integrated effects of the groupand only to a small extent by changes in differentialproperties of separate individuals.

Despite the fact that these local damages do notaffect the general flow of population processes and donot change their regular dynamics, they serve as a“switch” of a number of behavioral reactions thatallow significantly and operationally increasing boththe density of the invasive inflow of seeds and the

speed and efficiency of seed reproduction on the dam�aged areas. Regardless of the extension of commonborders, most reproductive individuals of all the levelsof vitality get involved in this process from neighboringregions. Until the appearance of their own reproduc�tive individuals, the plots of T�1 remain dependentpoints of tension with multiple ways of self�regenera�tion.

The results obtained indicate that even underfavorable circumstances, the effects of such impactspersist after a few decades. And even after them, self�regeneration of the populations of herbaceous peren�nials to the initial entry�level does not seem to be anobvious result. This should be taken into account inthe planning of economic impacts and nature restora�tion activities.

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Translated by K. Lazarev

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regeneration patterns of natural populations of herbaceous perennials in spruce forests of the...

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