Michał Gradziński, Maria Jolanta Chmiel, Anna Lewandowska & Beata Michalska-Kasperkiewicz

SILICICLASTIC MICROSTROMATOLITES IN A SANDSTONE

CAVE: ROLE OF TRAPPING AND BINDING OF DETRITAL

PARTICLES IN FORMATION OF CAVE DEPOSITS

& Beata MICHALSKA- KASPERKIEWICZ3

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In sti tute of Geo logi cal Sci ences, Jagiel lo nian Uni ver sity, Ole an dry 2a, 30- 063 Kraków, Po land; e- mail: mi chal.gradz in ski@uj.edu.pl, anna.le wan dowska@uj.edu.pl

2

De part ment of Mi cro bi ol ogy, Uni ver sity of Ag ri cul ture in Kraków, Aleja Mick iewicza 24/28, 30- 059 Kraków, Po land; e- mail: mjchmiel@poc zta.onet.pl

3

KTJ Speleok lub Biel sko Bia³a, 1-go Maja 45, 43- 300 Bielsko- Bia³a, Po land; e- mail: beata_mi chals@poc zta.onet.pl Gradzi ñski, M., Chmiel, M. J., Le wan dowska, A. & Michalska- Kasperkiewicz, B., 2010. Sili ci clas tic mi cro-stro mato lites in a sand stone cave: role of trap ping and bind ing of de tri tal par ti cles in for ma tion of cave de pos its. An nales So cie ta tis Ge olo go rum Po lo niae, 80: 303–314.

Ab stract: The ar ti cle deals with finely lami nated mi crostro mato lites com posed of de tri tal sili ci clas tic par ti cles, mainly quartz, feld spars and clay min er als, lin ing the walls of W So potni Wielkiej Cave (Pol ish Outer Car pa -thi ans). Newly pre cipi tated min eral phases do not con trib ute to their growth. The mi crostro mato lites cover ver ti cal and over hang ing walls of the cave. They form sub ho ri zon tal rip ples and tongue- shaped stepped mi croter racet tes. The stro mato lites are con structed by bac te ria and Actino my ce tes. Seven mor pho types of micro- organisms have been dis tin guished. Trap ping and bind ing of de tri tal par ti cles re sult in the mi crostro mato lite growth. The growth is in flu enced by the rela tively close dis tance to the soil cover which pro vides de tri tal min eral par ti cles and by the pres ence of gravi ta tion ally wid ened fis sures which guide the wa ter trans port ing min eral par ti cles down to the cave. The par ti cles are trans ported only dur ing wet pe ri ods. The epi sodic sup ply of the par ti cles re sults in visi ble lami na tion of mi crostro mato lites. The mi croter racet tes form in zones of in creased water flow. The lack of auto -chthonous com po nents most proba bly re flects too low satu ra tion of wa ter to pre cipi tate any min er als.

Key words: bac te ria, Actino my ce tes, biofilm, speleo thems, Outer Car pa thi ans. Manu script re ceived 18 September 2010, ac cepted 18 November 2010

IN TRO DUC TION

Con struc tive role of mi cro-or gan isms in speleothem for ma tion is a fo cus of grow ing in ter est (see re view by Northup & Lavoie, 2001; Jones, 2001, 2010; Taboroši, 2006 and ref er ences quoted therein). A great body of lit er a ture ex ists on the im por tance of bac te ria and fungi for pre -cip i ta tion of car bon ate speleothems, es pe cially for the growth of moonmilk, cave pisoids and some sub aque ous for ma tion (e.g., Gradziñski et al., 1997b; Gradziñski, 2000; Melim et al., 2001; Jones, 2009a; Curry et al., 2009). The plau si ble di rect or in di rect in flu ence of these mi cro-organi-sms on the crys tal li za tion of nee dle-fi bre cal cite in cav ern en vi ron ment is com monly dis cussed (Blyth & Frisia, 2008; Rich ter et al., 2008). The pre cip i ta tion of speleothem-for-ming man ga nese and iron ox ides, opal and phos phates is also me di ated by bi o log i cal ac tiv ity (e.g., Gradziñski et al., 1995; Manolache & Onac, 2000; Aubrecht et al., 2008; Jones, 2009b).

Con versely, lit tle at ten tion has been drawn to trap ping and bind ing of de tri tal grains of var i ous min eral com po si

-tion as a con struc tive mech a nism in speleothem growth. Such mech a nism has been men tioned on the oc ca sion of study ing the microbially driven pre cip i ta tion of di verse speleothems in cav ern en vi ron ment. How ever, it has only been re garded as a sub or di nate con tri bu tion to speleothem growth. Jones and Motyka (1987) showed that al gae trap and bind car bon ate micrite, which even tu ally leads to for ma tion of the well vis i ble lam i na tion. Cyanobacteria, rep re -sent ing Gleocapsa sp., are ca pa ble of trap ping air-born quartz grains and in sect frag ments, which col lec tively con -trib ute to the for ma tion of cray fish-like pro file sta lag mites lo cated in en trance parts of Aus tra lian caves (Cox et al., 1989a, b; James et al., 1994). Jones (1995) noted some ex am ples of trap ping and bind ing of de tri tal grains to the sub -strate by or ganic fil a ments in the biofilm cov er ing walls in twi light zones of caves. Cunningham et al. (1995, fig. 7) illustrated bind ing of cor ro sion re sid uum by fil a men tous miro-or gan isms in Lechugilla Cave (New Mex ico). CaÔa-veras et al. (2001) men tioned trap ping and bind ing of

micritic grains re sult ing from sub strate break down in the fa -mous Altamira cave. Gradziñski and Holúbek (2005) found de tri tal sil i cate and do lo mite grains in cor po rated into sub -aque ous cottonball speleothem in Zlomiská Cave (Slovakia); the cottonballs pre sum ably orig i nated un der bi o log i -cal me di a tion. Baskar et al. (2006) sug gested that de tri tal grains or gan i cally trapped and bound are in cor po rated into car bon ate speleothems in Borra Caves (In dia). Also, biolo-gicaly me di ated opal speleothems from Ven e zue lan caves con tain some grains trapped and bound by mi cro-or gan isms (Aubrecht et al., 2008). De tri tal grains con trib ute greatly to for ma tion of some speleothems in sand stone caves in the Pol ish Outer Carpathians (Ur ban et al., 2007). In some cases they com pose de pos its whose re lief im i tates the mor phol ogy taken by speleothems. Some of these forms are ce -mented with opal.

Clastic for ma tions built of sand- and silt-sized grains are pres ent in many caves (Hill & Forti, 1997, p. 219–221 and ref er ences therein). Some of them are akin to car bon ate speleothems in their shape and di men sion al though they are only ce mented by car bon ate min er als (e.g., Baker, 1942). In fact, many of them rep re sent ero sional rem nants af ter ac cu -mu la tion of cave clastics (e.g., Gradziñski & Radomski, 1960) and they do not share their or i gin with typ i cal speleothems. Ver micu la tions rep re sent an other type of cave for ma tion com posed of siliciclastic ma te rial (Hill & Forti, 1997, p. 221–223). They are found on cave walls or ceil ings as ir reg u lar spots or bi fur cat ing stripes. Their ar ray gives the over all im pres sion of a group of warms or a ti ger or leop ard skin. In the ma jor ity of pa pers no sug ges tions have been made on the pos si ble role of mi cro-or gan isms in the or i gin of the above clastic cave for ma tions, ex clud ing spe cial type

of vermiculation in sulfidic caves. Al though Bini et al. (1978) con sid ered in flu ence of bac te ria on for ma tion of vermiculation, ul ti mately they ruled out this in flu ence.

This pa per docu ments the pres ence of mi crostro mato -lites com posed of de tri tal sili ci clas tic par ti cles lin ing the walls of a sand stone cave. The sig nifi cance of mi croor gan -isms to trap ping and bind ing de tri tal par ti cles, and there fore to form ing the mi crostro mato lites in ques tion, is dis cussed. A hy pothe sis is put for ward on a simi lar con struc tive role of micro- organisms in for ma tion of clastic- rich lami nae within car bon ate speleo thems.

GEO LOG I CAL AND SPELEOLOGICAL

SET TING

Siliciclastic microstromatolites were found in W Sopotni Wielkiej Cave (Jaskinia w SoSopotni Wielkiej) and re -ported dur ing a lo cal sym po sium (Gradziñski et al., 2001). The cave is lo cated in the Beskid ¯ywiecki Mts. From the geo log i cal point of view this re gion be longs to the Outer Carpathians, which are built pre dom i nantly of Cre ta ceous– Palaeogene flysch (Fig. 1). The cave is formed in the Eocene thick-bed ded sand stones of the Magura beds be long ing to the Raèa Unit of the Magura Nappe (Golonka & Wójcik, 1978). The sand stone is fine grained. An X-ray diffractogram of the sand stone re veals the pres ence of quartz, al kali feld spars and two groups of phyllosilicates, with a ma jor or der layer sep a ra tion of 14.5  and 10.0  (see Figs 4A, 5A). The 14.5  min er als are ei ther chlorite or smectite, al though fur -ther anal y ses are nec es sary for ex act iden ti fi ca tion. The 10.0  peak may in di cate the pres ence of illite or micas. Ob ser va tions in op ti cal mi cro scope re vealed the pres ence of mus co vite and bi o tite micas (see Fig. 5A); a con fir ma tion of the illite pres ence re quires < 2 µm frac tion sep a ra tion.

The cave main en trance is lo cated at the altitude of 840 m (Klas sek, 1997). The cave is com posed of a se ries of pas sages up to 2 m wide and 5 m high, with to tal 101 m in length (Fig. 2). The pas sages dis play rec tan gu lar or tri an gu -lar cross- sections. Their floors are lit tered with sand stone de bris. The cave origi nated due to wid en ing of frac tures un der the ac tion of downslope creep of sand stone blocks to wards the neigh bour ing val ley. Hence, it rep re sents a crev ice type cave (sensu Pal mer, 2007, p. 7). The cave is pro -tected as a ‘na ture monu ment’.

The mi crostro mato lites cover the walls of the Ko mora Trójk¹tna (Tri an gu lar Cham ber) lo cated ca. 12 m from the cave en trance. They were noted by Mikuszewski (1973) and Klas sek (1997) but rec og nized as car bon ate flow stones. The cham ber is com pletely dark; its walls are cov ered by drops of con den sa tion wa ter. The in ter nal tem pera ture is 5.5 °C (Klas sek, 1997). The thick ness of the roof rock over the Tri -an gu lar Cham ber is es ti mated at 6–8 m. The area over the cave is for ested.

MA TE RIAL AND METH ODS

Mor phol ogy and in ter nal struc tures

Mi crostro mato lites were docu mented in the Tri an gu lar Cham ber (Fig. 2). Sam pling was lim ited to the mini mum due to the cave pro tec tion. Sam ples were col lected from

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both sides of the cham ber, pref era bly from hid den places. A thin sec tion was made from one sam ple sunken in a box filled with AR ALDITE. In ter nal struc tures of the mi crostro -mato lites were ob served un der pet ro graphic and scan ning elec tron mi cro scope (SEM) JEOL 5410, cou pled with a mi cro probe (EDS) Voy ager 3100 (No ran prod uct). The sam -ples were mounted on SEM hold ers with sil ver glue and coated with C or Au. Sam ples were ly ophi lized prior to coat ing to pre vent col lapse of the or ganic struc ture.

Min eral com po si tion

Sam ples were ana lysed by the pow der X ray dif frac to me try (XRD) us ing a ver ti cal XPert APD Phil ips gonio me -ter (PW 1830). In fra red ab sorp tion spec tra (IR- FT) were ob tained at am bi ent tem pera ture and with 2.0 cm–1 reso lu -tion us ing a BIO- RAD Fou rier Trans form Spec trome ter (FTS 135).

Mi cro bi ol ogy

Sam ples were asep ti cally col lected to ster ile glass flasks, trans ported to labo ra tory and sus pended in physio -logi cal salt so lu tion, shaken and in ocu lated in liq uid and agar me dia. They were in cu bated at 20 °C and 35 °C from 1 to 21 days. The growth of micro organisms was sys tem ati -cally moni tored. The fol low ing mi cro bio logi cal me dia were used for iso la tion: Beaf Ex tract – Nu tri ent Broth – Merck, Tryp ti case Soy Broth (Soybean- Casein Di gest Me dium) – BioMer ieux, Nu tri ent Agar – Merck, TSA (Tryp case Soy Agar) – BioMer ieux, Soil Ex tract Agar (At las & Parks,

1997), Iron Bac te ria Iso la tion Me dium (At las & Parks, 1997) and Actino my ce tes Iso la tion Agar (At las & Parks, 1997). Af ter in cu ba tion the clean cul tures of bac te ria were iso lated on agar me dia (Pep per & Gerba, 2004). Mor phol -ogy, Gram stain and bio chemi cal pro prie ties of bac te ria were ana lyzed to iden tify mi croor gan isms. Spe cies iden ti -fi ca tion was based on Ber gey’s Man ual of De ter mi na tive Bac te ri ol ogy and Ber gey’s Man ual of Sys tem atic Bac te ri ol ogy (Holt, 1989, 1994). Since there are no stan dard bio chemi cal tests for the ma jor ity of iso lated gen era, the bio chemi cal tests were in di vidu ally se lected ac cord ing to di ag -nos tic manu als.

RE SULTS

Re lief

The mi crostro mato lites dis play a soft, pasty con sis tency and con tain a sub stan tial amount of wa ter. They cover the ver ti cal and over hang ing walls of the Tri an gu lar Cham ber (Fig. 3A, B, D). The dip of rocky walls cov ered with the mi crostro mato lites ranges from 90° (ver ti cal) to 110° (over -hang ing). The mi crostro mato lites oc cur also on ver ti cally ori ented con vex bends of a cave wall, which over hangs with an an gle be tween 93° and 120° (Fig. 3A, B). Re cently, they coat around 0.5 m2 of the south ern wall of the cham ber (A in Fig. 2) and around 1 m2 of its op po site wall (B in Fig. 2). How ever, it is very plau si ble that they for merly oc cu pied a much big ger area and have been de stroyed by visi tors. On the ver ti cal walls, they form more or less hori zon tal rip ples with re lief (dis tance be tween the wall and the crest), up to

Fig. 2. Sim pli fied map of W Sopotni Wielkiej Cave (af ter Klassek, 1997),;sam pling sites are in di cated (white ar rows), big black ar row shows the cave en trance

2 mm (Fig. 3D). The length of par ticu lar ripples ex ceeds 35 cm. The neigh bour ing rip ples bi fur cate. With the wall dip chang ing to wards more over hang ing, the rip ples be -come more sinu ous and form tongue- shaped stepped mi cro- terracettes (Fig. 3A, B, D). The mi croter racet tes over hang and their up per parts dip out ward at a maxi mum an gle of 45°. In some cases, a mi crorim is de vel oped along their crest with a mi cro pool formed be hind it. Mi croter racet tes are par ticu larly well de vel oped along a ver ti cally ori ented con vex bends of the cave wall (Fig. 3B). The dis tance be -tween the microterracette crest and rock wall reaches 1.5 cm. The ver ti cal dis tance be tween neigh bour ing rip ples ranges be tween 2 and 5 mm, whereas be tween mi croter acet -tes it is slightly higher and maxi mally reaches 8 mm.

Mi cro bi ol ogy

Mi cro bi o log i cal anal y sis re veals the oc cur rence of sev -eral taxa which are listed in Ta ble 1. It seems rea son able to ac cept that Arthrobacter, Ba cilli and Actinomycetes be long to in dig e nous microflora of the ana lysed microstromato-lites. A scarce pres ence of Micrococcus and Staphyloccus in the stud ied sam ples im plies that they should be con sid ered allochthonous. A pre ferred en vi ron ment of their growth sup ports the above view (Table 1).

Min eral com po si tion

Quartz is the ma jor con stit u ent of the microstromato-lites; the al kali feld spars and clay min er als have a smaller

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Fig. 3. Siliciclastic microstromatolites and their mor pho log i cal coun ter parts; if no oth er wise stated pho to graphs are taken in W Sopotni Wielkiej Cave: A – Con vex cor ner of cave wall cov ered with microstromatolite, microterrces are de vel oped along the cor ner (M), flat wall to the left is cov ered with rip ples (R); B – Microteracettes with microrim fring ing the micropool; C – Miroteracettes com posed of trav er tine, Sivá brada, (Spiš, Slovakia); D – rip ples cov er ing cave walls, bi fur ca tion of rip ples is vis i ble; E – Microterracettes and rip ples com -posed of moonmilk de pos its, Szczelina Chocho³owska cave (West ern Tatra Mts., Po land)

con tri bu tion. The clay min eral con tent, al though in sig nif i -cant, is slightly higher in the microstromatolites than in the host rock. Sheet sil i cates show dif frac tion peaks at 14.5  and 10.0  as well as a mi nor peak cen tered at 12.0  (Fig. 4A). The 14.5  and 10.0  peaks cor re spond prob a bly to min er als pres ent in the host rocks (smectite or chlorite and micas), whereas the 12.0  peak re flects most likely the pres ence of mixed-layer clay min er als. This is sue re quires fur ther in ves ti ga tion.

Ad di tional in for ma tion as to the com po si tion of the microstromatolite sam ple is brought by IR ab sorp tion spec -trum (Fig. 4B). A weak ab sorp tion band at 1405 cm–1 may point to the pres ence of only very small amounts of car bon ates. The weak in ten sity of this ab sorp tion bend and the ab -sence of ap pro pri ate dif frac tion peaks, that is 3.03  (Fig. 4A) in di cate al most neg li gi ble amount of car bon ates. The microstromatolites have sim i lar com po si tion to the sandsto- ne which hosts the cave (Fig. 5A). The XRD pat terns are given for com par i son of the min er al ogy of both rocks (Fig. 4A).

In ter nal struc tures

The sur face of the microstromatolites is un even. It dis -plays el e vated clumps sur rounded by de pres sions. The clumps show lower po ros ity than the de pres sions. Also the microrim crest is char ac ter ized by a lower po ros ity than the micropool fringed by it (Fig. 5B).

The microstromatolite is finely lam i nated, with the lamina thick ness ca. 50–200 µm vis i ble due to dif fer en ti a -tion in min eral com po si -tion, grain size, and prob a bly changes in microporosity (Fig. 5C). The laminae are con vex out -ward, ir reg u lar. Some of them have con fused bound aries. They are com posed of par ti cles of clay and fine silt frac tion. Spo radic quartz and mus co vite grains reach up to 150 µm across (Fig. 5C). Out sized quartz and mus co vite grains are con cen trated in thicker laminae.

Ta ble 1

List of de ter mined mi cro-or gan isms with their ma jor char ac ter is tics (af ter Holt, 1989, 1994)

Bacteria Occurrence Morphology Properties

Arthrobacter sp. sample 1, sample 2 Gram positive irregular, nonsporing rods, in old cultures cocci

Aerobic, chemoorganotrophic; grows on simple media, widely distributed, principally in soils; psychrothropic arthrobacters have been reported to predominate in subterranean cave silts Amycolata autotrophica sample 1, sample 2 Gram positive, Actinomycetes, branched

vegetative hyphae

Aerobic, chemoorganotrophic - facultatively autotrophic; isolated from diverse habitats

Bacillus alcalophilus sample 1 Endospore forming, gram positive rods Aerobic, chemoorganotrophic; alcali tolerant, isolated from various material in media at pH 10

Bacillus megaterium sample 1, sample 2 Endospore forming, gram positive rods, cell diameter over 1 µm

Aerobic, chemoorganotrophic; found in wide range of habitats; grows in low temperatures

Bacillus mycoides sample 1 Endospore forming, gram positive rods, cell diameter over 1 µm, forms filaments

Aerobic but can grow anaerobically, chemoorganotrophic; forms rhizoid colonies; widely distributed

Micrococcus varians sample 2 Gram positive cocci, nonosporing Aerobic; micrococci are common on mammalian skin, soil, air Staphylococcus xylosus sample 1 Gram positive cocci, nonsporing

Facultatively anaerobic; staphylococci are mainly associated with skin, but often isolated from food products, dust and water

Streptomyces sp. sample 2 Gram positive, Actinomycetes, extensively branched vegetative hyphae

Aerobic; chemoorganotrophic; widely distributed and abundant in soil

Fig. 4. A – XRD pat tern of host rock and microstromatolite; Q – quartz, Al – al bite, Kf – K feld spar, C14 – clay min er als with main re flec tion 14.5 ; C12 – clay min er als with main re flec tion 12.5 , C10 – 10.0  micas or illite, B – IR ab sorp tion spec trum of the microstromatolite

Ob ser va tion un der SEM re veals the in ter nal struc tures of the microstromatolites. They are com posed of min eral grains, mi cro-or gan isms and their extracellular poly meric se cre tions (EPS).

Clay min er als com pose clumpy ag gre gates, up to 200 µm across (Figs 5B, D, 6A). The ag gre gates also com prise some ad mix tures of big ger min eral grains. Mica grains dis play an gu lar shape and tab u lar habit (Fig. 6B). They con tain Al,

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Fig. 5. A – Host sand stone: quartz and al kali feld spar grains ac com pa nied by bi o tite and mus co vite flakes pres ent in the host sand stones, Q – quartz, Kf – po tas sium feld spar, M – mus co vite, B – bi o tite, Cl – clays, thin sec tion, crossed polars; B – Ir reg u lar clumps form -ing a microstromatolite sur face, more com pact arch struc ture to the right side is crest of microrim (ar row) seen from above, gran u lar, more po rous ma te rial fill the micropool, SEM photo; C – Finely lam i nated crust of the microstromatolite is com posed mainly of clay min er als (Cl), with larger mus co vite flakes (M) and vari able size quartz (Q) grains, laminae rich in coarsergrained quartz are in di cated with big ar -rows, thin sec tion, par al lel polars; D – Clumps de void of mi cro-or gan isms from deeper part of microstromatolite, SEM photo

Ta ble 2

Char ac ter is tics of dis tin guished morphotypes

Morphotype Shape Dimensions Branching Morphology Cell arrangement Comments Figure 1 coccoid to short

rods

Æ – 0.8 µm

length < 1.5 µm no smooth

chains or three-dimensional

colonies covered with EPS 6C, 7A-D 2 rods Æ – 0.8 µm

length - 2-3 µm no smooth straight or zig-zag chains 7E 3 spindle-shaped

rods

Æ – 1 µm

length - 4-5 µm no granulated chains 7F

4 coccoid Æ – 0.5–0.8 µm no smooth single or in linked pairs co-occur with no 6 8D 5 irregular ovoid Æ – 6 µm no irregular but

smooth single or in linked pairs co-occur with no 6 8C 6 filamentous Æ – 1.1–1.3 µm

length > 20 µm yes spinose filaments slightly curved

co-occur with no 4, 5, 7, forms dense mat of intertwinned filaments

8A

7 filamentous Æ < 0.5 µm

Si and K, which, along with the re sults of the XRD anal y sis, sug gests mus co vite (Fig. 6B). Quartz grains are more roun-ded and de void of cleav age planes with Si as their ma jor EDS de tect able com po nent. Min eral grains co-oc cur with, and are en tombed by, mi cro-or gan isms which in many cases are tightly clung to them (Figs 6C, D, 7).

Seven morphotypes of mi croor gan isms have been dis -tin guished. Their char ac ter is tics are listed in Ta ble 2. All morphotypes are built ex clu sively of or ganic mat ter. Nei -ther EDS anal y ses nor ob ser va tion un der SEM re veal any traces of cell min er al iza tion. Mi cro-or gan isms show three-di men sional morphologies, only some of them are collapsed. The mi croor gan isms and their EPS oc cur on the sur -face of microstromatolites and in their shal low subsur-face where they form an ac tive biofilm. Deeper on, their amount radically decreases (Fig. 5D).

Af fin ity of the par tic u lar morphotypes is hard to be de -ter mined even in the light of the re sults of mi cro bi o log i cal anal y ses. The morphotypes 1 to 3 most prob a bly rep re sent bac te ria (Fig. 7A–C). Chains of coccoid cells, such as those of the morphotype 1, are typ i cal two-di men sional col o nies of coccoid bac te ria de vel oped due to cell di vi sions (Fig. 7A–C). Sim i larly, chains of rods char ac ter is tic for the morphotypes 2 and 3 are the ef fect of cell di vi sion (Fig. 7D, E). The morphotype 2 may be as signed to Arthrobacter ge nus, tak ing into ac count the list of de ter mined mi croor gan

-isms and their shape and size (Ta ble 1). More over, it forms a zig-zag chain of cells typ i cal of this ge nus (cf. Carlile, 1979). The fil a men tous morphotypes 6 and 7 re sem ble Actinomycetes. The morphotype 6 is par tic u larly akin to ‘hyphae morphotype 5’ de scribed and il lus trated by Jones (2009b, fig. 7A–C) from the Grand Cayman speleothems. This morphotype forms ex ten sive three-di men sional, po -rous net work (Fig. 8A–D). The coccoid morphotype 4 and ovoid morphotype 5 may rep re sent both bac te ria and spores of Actinomycetes (Fig. 8C, D). How ever, their close spa tial re la tion ship with the morphotype 6 strongly sug gests the lat ter pos si bil ity. Al though the EPS form ir reg u larly twisted fil a ments or a dense layer which covers micro-organisms, none of the morphotypes is associated with a copious amount of EPS.

DIS CUS SION

Trap ping and bind ing of de tri tal par ti cles cause the growth of the stud ied microstromatolites. Two mech a nisms are evoked to en trap de tri tal grains into a stromatolite – ad he sion by sticky EPS and baf fling by com pli cated threedi men sional mi cro bial com mu nity (Rid ing, 1991). In the dis -cussed case the lat ter mech a nism seems to be de ci sive, since the SEM ex am i na tion of the stud ied sam ples has not re

-Fig. 6. A – Ir reg u lar clumps com posed of clay min er als, fil a ments of EPS are vis i ble (ar row), subsurafce part of stromatolite; B – Mus -co vite flakes in -cor po rated into microstromatolite; C – quartz grain -cov ered with partly -col lapsed mi cro bial fil a ment, -coc-coid bac te ria (morphotype 1) are vis i ble to the left, D – min eral grain en tombed by mi cro-or gan isms, biofilm sur face seen from above; all pho to graphs un der SEM, in A–C EDS chem i cal com po si tion is in di cated

vealed co pi ous amounts of EPS. Highly branched cells of Actinomycetes Amycolata and Streptomyces spe cies act as a dense threedi men sional net work ca pa ble of baf fling the de -tri tal par ti cles. Nev er the less, Arthrobacter and Ba cil lus cells can ex crete some slimy sub stances, hence the for mer mechanism may also work, but only to a limited extent.

Newly pre cip i tated min eral phases seem not to con trib -ute to the microstromatolite growth. Bear ing in mind their growth mech a nism, the microstromatolites in ques tion rep -re sent ag glu ti nated stromatolites sensu Rid ing (1991). In this as pect they bear a strong re sem blance to some mod ern stromatolites in ma rine (Schwarz et al., 1975) and lac us trine set tings (Squyres et al., 1991) as well as to sev eral fos sil ex am ples (Mar tin et al., 1993; Braga & Mar tin, 2000 and ref -er ences th-erein). Mar tin et al. (1993) coined the t-erm

siliciclastic stromatolite which ad e quately re flects the com po si -tion and mode of growth of the stud ied ex am ples. The size of en trapped siliciclastic grains dif fer en ti ates cave mi crostromatolites from the hith erto de scribed ma rine and lac us -trine ones. Non-spelean forms are com posed mainly of sand grains with some ad mix tures of coarser ma te rial (Mar tin et

al., 1993); how ever, some silt-rich siliciclastic stromatolites

are also known (Bertrand-Sarfati, 1994). Jones and Kahle (1985) in tro duced the term microstromatolites in or der to de scribe microbolites, formed of fine car bon ate par ti cles and dis play ing fine lam i na tion, rec og nized in cave sed i -ments from the Cayman Is lands. The term microstroma-tolite seems to be ap pro pri ate to the stud ied de pos its. The mode of growth dis tin guishes the de scribed forms from any other hith erto known mi cro bial cave de pos its be ing con

-310

Fig. 7. A–D – Morphotype 1 rep re sent ing bac te ria, elon gated chains de vel oped due to cell di vi sions; E – Chain of rod-shaped cells (morphotype 2) prob a bly rep re sent ing Arthrobacter sp., F – Morphotype 3 formed elon gated chains of spin dleshaped cells with gran u -lated wall; all pho to graphs un der SEM

structed pre dom i nantly by min er als pre cip i tated from so lu -tion. Al though the dis cussed de pos its mimic the re lief of some speleothems (see be low and Fig. 3), they can not be in -cluded into this ge net i cally de fined group, be cause they con tain ex tremely small amount of, if any, sec ond ary min -eral phases, that is min er als pre cip i tated within a cave (cf. Hill & Forti, 1997, p. 45). The microstromatolites bear mor -pho log i cal re sem blance to speleothems de scribed from other sand stone caves in the Pol ish Outer Carpathians (Ur -ban et al., 2007). Many of the latter forms are composed of detrital grains cemented with opal, which is not a case in the discussed microstromatolites.

The growth of the dis cussed microstromatolites is in flu -enced by sev eral fac tors. The in flux of de tri tal par ti cles is pos si ble due to rel a tively close dis tance to the Earth sur face and the weath er ing zone (see Ur ban et al., 2007). It is ad di tion ally fa cil i tated by the pres ence of grav i ta tion ally wid -ened fis sures which guide the wa ter trans port ing min eral par ti cles down to the cave. The fine-grained na ture of microstromatolites more prob a bly de pends on the pref er en -tial re moval of such grains from soils. The par ti cles must be trans ported only dur ing the wet pe ri ods, that is dur ing thaw or af ter heavy rains, since dur ing the vis its to the cave its walls were cov ered only by drops of con den sa tion wa ter and were de void of seep ing wa ter. The ep i sodic sup ply of ma te rial re sults in a vis i ble lam i na tion of microstromatolite

(Fig. 5C). The mi cro bial biofilm cov er ing the ver ti cal or even over hang ing cave walls trap and bind the trans ported de tri tal par ti cles (Figs 6C, D, 8D). The ca pac ity of bac te ria to sta bi lize sand and finer grains has been ex per i men tally con firmed (Mead ows et al., 1994; Wes tall & Rincé, 1994; Dade et al., 1996). The par ti cles are sta bi lized on the sur face of microstromatolite and then cov ered with a newly de -vel oped biofilm (Fig. 6D). Si mul ta neously, an older part of the biofilm dis in te grates due to se nes cence, which is prob a -bly fa cil i tated by cov er ing min eral par ti cles. The laminae which com prise out sized quartz grains mark the es pe cially wet ep i sodes when rel a tively coarse-grained ma te rial could have been remobilized from soils and washed down into the cave (Fig. 5C). The microterracettes form in zones of more in tense wa ter flow (Fig. 3A–C). It is con firmed by their pre- ferential for ma tion along ver ti cally ori ented con vex bends of the over hang ing cave wall, thus the part where water flow is con cen trated due to the sur face ten sion. The down ward in clined shape of par tic u lar microterracettes most prob a bly re -sults from plas tic de for ma tion and creep ing of soft microstro- matolite un der the ac tion of grav ity (Fig. 3C).

The in flux of seep age wa ter is also im por tant for the mi cro-or gan ism as sem blage form ing the microstromatoli-tes, which most prob a bly de pends on the in put of or ganic mat ter from the sur face (cf. Groth & Saiz-Jimenez, 1999). On the one hand, the lim ited en ergy of seep ing wa ter sorts

Fig. 8. A – Three-di men sional po rous net work com posed mainly of morphotype 6 (Actinomycetes); B – De tailed view of the net work, spinosewalled morphotype 6 dom i nate, smoothwalled morphotype 7 oc cur subordinately (ar rows), C – ovoid morphotype 5 (ar rows) as so ci ated with spinosewalled fil a men tous morphotype 6; D – min ute coccoid bod ies of morphotype 4 (spores of Actinomycetes; small ar -rows) as so ci ated with fil a men tous morphotypes 6 and 7 (big ar row), in the cen tre platy min eral grain en trapped within or ganic fil a ments (EDS chem i cal com po si tion is in di cated); all pho to graphs un der SEM

min eral grains and con trols the fine-grained com po si tion of microstromatolites. On the other hand, it al lows del i cate mi -cro bial biofilm to ex ist on cave walls; the higher en ergy of flow would cause de struc tion and scrub bing off of the mi -crobial biofilm.

The lack of pre cip i ta tion of min er als is most prob a bly con nected with the chem is try of the seep ing wa ter. Al though di rect data are lack ing, we can hy poth e size that the wa ter in W Sopotni Wielkiej Cave is sim i lar to the wa ter in other non-karst caves in the flysch rocks of the Outer Carpathians. Zawierucha et al. (2005) re ported that the vadose wa ter in those caves is only slightly more min er al ized than the rain wa ter. The mean con cen tra tion of the Ca ion is only 9.5 mg/l, that is def i nitely lower than in karst caves. For ex am -ple, the Ca con tent in wa ter from se lected Slo vak karst caves ranges be tween 44.1 and 132.1 mg/l (Motyka et al., 2005) whereas in the vadose zone wa ter of karst caves on the Kraków–Wieluñ Up land it is from 52 to 137 mg/l (Ró¿-kowski, 2006). More over, one can sup pose that the wa ter which quickly per co lates down af ter heavy rains, as it is in the dis cussed case, has lower con cen tra tion of ions due to its lower res i dence time (see dis cus sion in Musgrove & Ban ner, 2004). Thus, the wa ter in the stud ied cave most prob a -bly does not achieve the ap pro pri ate sat u ra tion to pre cip i tate car bon ate min er als.

The mi cro-or gan isms de tected within the microstroma-tolites have been re ported from other caves and have been sup posed to in flu ence – ac tively or pas sively – pre cip i ta tion of min er als. Liv ing bac te ria be long ing to Arthrobater were de tected within grow ing moonmilk de pos its (Gradziñski et

al., 1997b) and cave pearls (Gradziñski, 2000).

Phosphati-zed mi cro bial cells as signed also to this ge nus are re ported from the speleothems of Grand Cayman Is land (Jones, 2009b). Min er al ized, com monly cal ci fied, Actinomycetes, in clud ing var i ous spe cies of Streptomyces, are known from cave de pos its of Spain (CaÔaveras et al., 2001), It aly (Groth

et al., 2001), USA (Melim et al., 2008) and the Cayman Is

-lands (Jones, 2009a, b). This leads to the sug ges tion that the min er al iza tion of mi croor gan isms in the cav ern en vi ron -ment is strongly de pend ent upon chem is try of the feed ing wa ter.

The de scribed or i gin of the stud ied siliciclastic mi cro-stromatolites may shed some light on the for ma tion of other cave de pos its. The un con sol i dated ac cu mu la tions of fine-grained clastics on cave walls, in clud ing ver micu la tions com mon in many caves, can be also formed by the trap ping and bind ing of de tri tal par ti cles by mi croor gan isms. Sev -eral forms of speleothems rec og nized in some caves of the Pol ish Outer Carpathians may also share their or i gin with the dis cussed microstromatolites. They are also com posed of siliciclastic ma te rial, dis play lam i na tion and con tain some laminae rich in coarser quartz sands (Ur ban et al., 2007).

The de scribed stromatolites have a low po ten tial for pres er va tion. The only one pos si bil ity for their pres er va tion is a quick ce men ta tion with cal cium car bon ates or opal act -ing as ce ment or cov er -ing a stromatolite as a youn ger flowstone (cf. Ur ban et al., 2007). Thus, clasticrich lay ers oc -cur ring within flowstones and other speleothems (see Dzia-dzio et al., 1993; Gradziñski et al., 1997a; Turgeon & Lund- berg, 2001) may be fos sil coun ter parts of

microstromato-lites formed dur ing the break of crys tal li za tion and in tense clastic sup ply. Since the mi cro-or gan isms en trapped de tri tal grains on the ver ti cal and over hang ing walls in the stud ied case, it seems plau si ble that they also play a role in sta bi li za -tion of such grains de pos ited on a flowstone sur face and later ce mented with cal cite or ar agon ite. Non-min er al ized mi cro-or gan ism cells have been de com posed and have not been pre served. Hence, the lay ers in ques tion lack any traces of mi cro-or gan isms.

The clos est mod ern coun ter parts of the de scribed forms, both in mor phol ogy and or i gin, are sand rip ples de -vel op ing on steep sand stone crags de scribed by Pen te cost (1999) from Kent. They are com posed of sand grains sta bi lized by al gae and mosses and dis play re lief strik ingly sim i lar to that of the de scribed microstromatolites. The most im -por tant dif fer ence be tween the re lief of sand rip ples and the cave microstromatolites is a slightly greater lat eral dis tance be tween neigh bour ing rip ples (5.6–8.0 mm for the sand rip -ples). Other sur face coun ter parts are lit ter dams oc cur ring on slopes. They owe their or i gin due to ac cre tion of small or ganic and min eral par ti cles and their sta bi li za tion by mosses (Eddy et al., 1999). The spa tial ar range ments and re lief of the lit ter dams are dif fer ent, mainly because of their formation on the inclined, not vertical slopes.

In ter est ingly, a re lief anal o gous to the de scribed mi cro-stromatolites char ac ter izes sur faces of sev eral ac tively grow ing con ti nen tal de pos its. The sur faces of some cave flowstones and dripstones are crenulated (Hill & Forti, 1997, pp. 71, 105) or rip pled (Ford, 1988). Sim i lar rip ples are formed on the grow ing ici cles (e.g., Ogawa & Furu-kawa, 2002). The cave drap er ies also dis play ser rated edges com posed of the stepped over hang ing terracettes com pa ra -ble to the terracettes formed by the de scribed microstromatolites. Iden ti cal terracettes are com posed of moonmilk de pos its (Gradziñski & Radomski, 1957). On the steeply in clined or over hang ing walls their up per sur faces are down -ward in clined sim i larly to the terracettes of the de scribed microstromatolites (Fig. 3E). Such ori en ta tion prob a bly re -sulted from some in sta bil ity of moonmilk hav ing a pasty con sis tency and from its ten dency to creep ing down, which is in com mon with the dis cussed microstromatolites. Also travertines form sim i lar reg u larly spaced terracettes. The lat eral dis tance be tween the neigh bour ing trav er tine ter race- ttes be comes shorter on steeper slopes (Ham mer et al., 2010). Hence, the trav er tine terrracettes de vel oped on ver ti cal and steep slopes may serve as a coun ter part of the de scribed rip -ples and terracettes formed by the microstromatolites. Nonetheless, in con trast with the microstromatolites and moonmilk ex am ples, the rim of trav er tine terracettes is al -most al ways per fectly hor i zon tal (Fig. 3C; Ham mer et al., 2010). It most prob a bly re sults from the ro bust con sis tency of trav er tine de pos its. Sur pris ingly, al though all the above men tioned cal car e ous de pos its orig i nated mainly by the pre -cip i ta tion of crys tals, not by the ac cu mu la tion of de tri tal par ti cles, they share their shape and geo met ric pat tern with the dis cussed microstromatolites whose growth is gov erned by dif fer ent fac tors. It adds a new di men sion to the dis cus sion on the fac tors in flu enc ing the shape and spa tial ar -range ments of terracettes in re cently grow ing travertines and speleothems.

CON CLU SIONS

1. The walls of W Sopotni Wielkiej Cave are cov ered with siliciclastic microstromatolites con structed by bac te ria and Actinomycetes, which trap and bind min eral par ti cles washed into the cave from over ly ing soil dur ing wet ep i -sodes.

2. Newly pre cip i tated min eral phases seem not to con -trib ute to the microstromatolite growth.

3. The microstromatolites form rip ples and microterra-cettes on ver ti cal and over hang ing cave walls. Their shape de pends upon the re lief of a cave wall. The microterracettes are lo cated where wa ter flow is more in tense, mainly along ver ti cally ori ented con vex bends of a cave wall.

4. It seems pos si ble that trap ping and bind ing mech a -nism in flu ences the or i gin of clastic-rich lay ers within cave flowstones. Clastic grains were sta bi lized by mi croor gan -isms and later were ce mented with cal cite or ar agon ite. The mi cro-or gan isms were sub se quently de com posed and have been not preserved till now.

5. Sand rip ples de vel op ing on al most ver ti cal sand -stone crags are a close ge netic and mor pho log i cal an a logue of the cave siliciclastic microstromatolites.

6. Rip ples (crenulations) on the speleothem and ici cle sur faces, as well as stepped microterracettes in speleothems and travertines share the same re lief with the de scribed microstromatolites in spite of their dif fer ent or i gin.

Ac knowl edge ments

MG was sup ported at the be gin ning of the study by the Foun -da tion for Pol ish Sci ence in the frame of Grant for Re search ers to Pro fes sor Józef KaŸmierczak. Renata Jach and Ryszard Gradziñ-ski as sisted in the field, Jaga Faber, Zuza Banach and Aleksandra WoŸnicka op er ated the SEM, Renata Jach also pre pared the fig -ures; the au thors grate fully ac knowl edge this help. The manu script bene fited from the re views of Tadeusz Peryt and Jan Ur ban.

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