Annales Societatis Geologorum Poloniae (2014), vol. 84: 143–151.
RUSOPHYCUS INEXPECTUS ISP. NOV. FROM THE FURONGIAN
(UP PER CAM BRIAN) OF THE HOLY CROSS MOUN TAINS
(PO LAND)
Grzegorz SADLOK
In sti tute of Paleobiology, Pol ish Acad emy of Sci ences, ul. Twarda 51/55, PL- 00-818 Warszawa, Po land;
e-mail: gregsadlok@gmail.com
ad dress for cor re spon dence: 77 Leadside Road, Aberdeen AB25 1RX, Aberdeenshire, Scot land, United King dom
Sadlok, G., 2014. Rusophycus inexpectus isp. nov. from the Furongian (Up per Cam brian) of the Holy Cross Moun -tains (Po land). Annales Societatis Geologorum Poloniae, 84: 143–151.
Ab stract: A new trace fos sil Rusophycus inexpectus isp. nov. is de scribed from the Furongian (Up per Cam brian)
strata of the Holy Cross Moun tains. This ichnospecies is prob a bly non-tri lo bite in or i gin and is com monly pre served as an undertrace. This pres er va tion style re sem bles that of Rusophycus ramellensis Legg, an in dex fos sil of the Cam brian Se ries 3. There fore, pre vi ous work ers mis in ter preted ma te rial from the Wiœniówka Sand stone For ma tion as Rusophycus ramellensis Legg (re corded by them as Cruziana barbata) and put the Cam brian Se ries 3 and Furongian bound ary within this unit. Rusophycus inexpectus isp. nov dif fers from Rusophycus ramellensis Legg: 1) in hav ing a smooth trap e zoidal area be hind the lobes; 2) in the pres ence of a di rect con tact be tween the endopodal lobes and ce phalic mar gin im prints and 3) in the re stric tion of the oc cur rence of the endopodal lobes and scratches to the ce phalic re gion of the trace fos sil. Care must be taken, when deal ing with the undertrace pres er va tion style of Rusophycus ramellensis Legg in other as sem blages, as it may rep re sent Rusophycus inexpectus isp. nov.
Key words: Rusophycus ramellensis, Rusophycus inexpectus, Furongian, Cam brian, ichnostratigraphy, new ichnotaxa.
Manu script re ceived 14 November 2013, ac cepted 3 May 2014
IN TRO DUC TION
Wiœniówka Du¿a (or Wiœniówka Wielka in older lit er a
-ture) is a well known site with Cam brian trace fos sils
(Or-³owski et al., 1970, 1971; Or(Or-³owski, 1992; Or(Or-³owski and
¯yliñska, 1996; Sadlok, 2010, 2011; Sadlok and Machalski,
2010). Or³owski (1992) used the ichnostratigraphical
scheme pro posed by Seilacher (1970, 1994, 2007) for strati
graphi cal sub di vi sion of the Wiœniówka Sand stone For ma
-tion as ex posed at this quarry (see the crit i cal opin ion in Ko-
walczewski, 1995). The Furongian (Up per Cam brian) trace
fos sils (Cruziana semiplicata Salter and Rusophycus
polo-nicus Or³owski, Radwañski and Roniewicz) were well
known from the Wiœniówka Sand
stone For
ma
tion
(Or-³owski et al., 1970, 1971; Seilacher, 1970; Radwañski and
Roniewicz, 1972). Sub se quently, a trace fos sil re sem bling
Rusophycus ramellensis Legg was re corded from this unit
un der the name Cruziana barbata Seilacher, 1970 by
Or-³owski (1992). Rusophycus ramellensis Legg is a typ i cal
Cambrian Se
ries 3 ichnospecies (Seilacher, 1970;
Mac-Naughton, 2007; see Cam brian sub di vi sion in Bab cock and
Peng, 2007). This ma te rial came from the low er most part of
the suc ces sion ac ces si ble in the Wiœniówka Du¿a Quarry
and it was the ba sis for rec og ni tion of the Cam brian Se ries 3
– Furongian bound
ary within the Wiœniówka Sand
stone
Formation (Or³owski, 1992). Later, ¯yliñska et al. (2006)
stud ied acritarchs and trilobites from the Wiœniówka Sand
-stone For ma tion and con cluded that the strata with R.
rame-llensis (their C. barbata) (Fig. 1) be longed to the lower part
of the Furongian. This most re cent view is fol lowed herein.
This pa per pro vides new data on trace fos sil from the
Wiœniówka Sand stone For ma tion that pre vi ously was as
-signed to Rusophycus ramellensis Legg (Or³owski, 1992).
New data show that the ma te rial from the Wiœniówka Sand
stone For ma tion rep re sents a new ichnospecies. The im pli
-ca tions of this dis cov ery for lo -cal strati graphic study and
palaeographical con sid er ations are briefly dis cussed.
LO CA TION AND GEO LOG I CAL
SET TINGS
The Wiœniówka Sand stone For ma tion be longs to the
Furongian, as in di cated by body fos sils (¯yliñska et al.,
2006). From a strati graphic point of view, the unit is lo cated
be tween the Pep per Moun tains For ma tion (Cam brian Se ries
3 to Furongian) and the Klonówka Shale For ma tion (Furon-
gian, see ¯yliñska et al., 2006). The strata of this for ma tion
are best ex posed at the Wiœniówka Du¿a Quarry and a few
smaller quar ries in the wes tern most part of the Holy Cross
Moun tains (Wiœniówka Ma³a and Podwiœniówka), as well
as in the Opatów area (Kowalczewski et al., 2006 and ref er
-ences therein; Fig. 1A). The es ti mated thick ness of the
Wiœ-niówka Sand stone For ma tion ranges from 80–200 m, up to
400–1400 m, de pend ing on the tec tonic model ap plied
(Ko-walczewski et al., 2006 and ref er ences therein).
The Wiœniówka Sand stone For ma tion com prises mainly
sand stones, heterolithic in ter vals and mudstones (Fig. 1B, C).
Sand stones dis play per va sive sili ci fi ca tion that ob scures the
orig i nal tex tural char ac ter is tics (Sikorska, 2000). These de
pos its ap pear to be well sorted and both tex tur ally and min
-er al og i cally ma ture (close to quartz arenites sensu
Nagte-gaal, 1978; thin sec tions were stud ied, but no point-count
modal anal y sis was done and ac tual per cent age val ues are
not avail able). The heterolithic beds dis play nu mer ous and
var i ous cur rent- and wave-rip ple struc tures (Radwañski and
Roniewicz, 1960; Fig. 1B, C) and flaser to len tic u lar bed
-ding. The char ac ter is tic fea ture of the sand stone interbeds is
their com mon amal gam ation and un du la tion of bed ding pla-
nes, with lat eral changes in the thick ness of beds (pos si bly
due to ?hummocky cross strat i fi ca tion). The strata of the
Wiœniówka Sand stone For ma tion have been in ter preted as
shal low ma rine by most au thors (D¿u³yñski and ¯ak, 1960;
Radwañski and Roniewicz, 1960; Jaworowski and
Sikor-ska, 2006).
MA TE RIAL AND METH ODS
This study was based on: 1) col lec tion of trace fos sils,
in or
der to test the pres
ence of Rusophycus ramellensis
Legg in the part of the sec tion with Furongian body fos sils
(see ¯yliñska et al., 2006), and 2) ob ser va tions on trace fos
-sil mor phol ogy; ma te rial from the Wiœniówka Sand stone
Fig. 1. Lo ca tion of Wiœniówka Du¿a (“Wielka” in older lit er a ture) Quarry and gen eral pro files of strata, ex posed on north ern and southern walls. A. Lo ca tion of Wiœniówka Du¿a Quarry. B. The gen eral pro file of north ern wall. C. Gen eral pro file of south ern wall. A ques -tion mark for the Furongian age in de scrip -tion of south ern wall pro file co mes from the lack of di rect body in dex fos sils for this in ter val (see text).
For ma tion was com pared with the type ma te rial and with
other sim i lar ichnospecies of Seilacher (1970), on the ba sis
of ma te rial from the Mu seum of Eberhard Karls Uni ver sity
in Tübingen; Palaeontological Col lec tion of Tübingen Uni
ver sity, Sigwartstraße 10, 72076 Tübingen, Ger many (ab
-bre vi a tion GPIT).
The ma te rial stud ied from the Wiœniówka Sand stone
Formation is a part of a trace fos sil col lec tion stored at the
In sti tute of Paleobiology of Pol ish Acad emy of Sci ences
(War saw, Po land; ab bre vi a tion ZPAL Tf. 4). The col lec tion
com prises ma te rial amassed by the Au thor (80%) and by
Marcin Machalski (20%) from the In sti tute of Paleobiology
of Pol ish Acad emy of Sci ences, War saw.
Terms, such as “cephalon”, “endopod” and “exopod”
(and re spec tive ad jec tives), cor re spond to terms ap plied
pre vi ously to var i ous parts of the trace fos sils dis cussed (see
Seilacher, 1970). How ever, all of these terms are in ter pre ta
-tive. The va lid ity of their ap pli ca tion to Rusophycus
inexpectus isp. nov., which is thought to rep re sent a nontri lo
-bite trace fos sil, is even more prob lem atic, but for the sake
of sim plic ity these terms are re tained herein. The ap pli ca
-tion of other terms would be also in ter pre ta tive.
SYS TEM ATIC DE SCRIP TION
Ichnogenus Rusophycus Hall, 1852
Di ag no sis: Short, bilobate, rarely multilobate traces. Lobes pre
-dom i nantly bi lat er ally sym met ri cal. Con vex forms (hypichnia) with dis tinct me dian fur row; con cave forms (epichnia) with me -dian ridge. Out line ovate to cof fee-bean-shaped; sculp tured with oblique to trans verse or lon gi tu di nal striae in var i ous ar range -ments, or al most smooth (Schlirf and Uchman in Schlirf et al., 2001).
Dis cus sion: Seilacher (1970) pro posed that all trace fos sils of pre
-sumed tri lo bite or i gin should be treated un der one ichnogenus Cruziana. How ever, Seilacher’s ap proach to ichnotaxonomy is not ac cepted by most work ers (e.g., Keighley and Pickerill, 1996; Schlirf et al., 2001 and ref er ences therein) and is not fol lowed here. Most work ers dif fer en ti ate be tween an elon gated, rib bon-like trace fos sil (Cruziana) and a short, ver ti cally bur rowed trace fos sil (Rusophycus; see dis cus sion in Jensen, 1997). The dif fer en -ti a -tion is not al ways straight for ward, but sim ply cri te ria, such as length-to-width ra tio, can be used (Keighley and Pickerill, 1996).
Rusophycus inexpectus new ichnospecies
Figs 2, 3A–C, 3E, F
1992 Cruziana barbata Seilacher, 1970 – Or³owski: p. 24, fig. 7.
Ma te rial and holotype: holotype – ZPAL Tf. 4/1329; other ma te
-rial – ZPAL Tf. 4/1, 2, 8, 13, 104, 112, 122, 145, 150, 161, 166, 231, 237, 250, 265, 275, 292, 350–351, 366, 395, 368, 389, 445, 450, 478, 831, 895–896, 898, 907, 910, 912, 914, 917–918, 958, 963–964, 673, 975, 998–999, 1023, 1029, 1067, 1081, 1239, 1331, 1328, 1329, 1330, 1335, 1359, 1361.
Et y mol ogy: “inexpectus” form Latin means un ex pected and re fers
to the un ex pected mor phol ogy of the well pre served, full ver sions of the ichnospecies.
Di ag no sis: Rusophycus with two heart-shaped lobes and with a
smooth trap e zoidal area be hind those lobes. The lobes are con nected with elon gated im prints lat er ally and meet me di ally, form -ing a groove which be comes a deeper de pres sion to ward one end of the trace fos sil. The de pres sion forms a V-shaped gap in the
fron tal part of the lobes. The ex tent of the lobes cor re sponds with the ex tent of elon gated lat eral im prints. The lobe-cov er ing ridges (scratches) have di ver gent pat terns and in deep undertraces a “mous-tache-like” pat tern is ap par ent.
De scrip tion: Rusophycus oc cur ring most com monly as a
hypich-nion with two large lobes, cov ered with ridges (scratches). This trace fos sil ap pears to be strongly bi ased to ward undertrace pres er -va tion and there fore its di men sions are con trolled by taphonomy. Well pre served spec i mens have lengths of be tween ~30 and ~50 mm and widths of be tween ~20 and ~40 mm. There is a V-shaped gap oc cur ring within the lobes. The lobes at one end of the trace fos sil are at a higher an gle to the cen tre-line in deeper spec i mens; the an gle ranges from ~25° to ~50° (Fig. 2B, C). Un ex pect edly, the well pre served ver sion of Rusophycus inexpectus isp. nov. also has a smooth rear area, which is trap e zoidal and oc curs just be hind the main lobes. This smooth area ta pers back wards. Well pre -served Rusophycus inexpectus isp. nov. also dis plays the pres ence of elon gated lat eral im prints, in con tact with and bor der ing the lobes; the lobes and lobecov er ing scratches ex tend across the en -tire width of the trace fos sil (Figs 2C, 3A–C). In lat eral view, the ex tent of the lobes cor re sponds with the ex tent of the elon gated lat -eral im prints (Fig. 3B, C). The lobe-cov er ing ridges (scratches) have a typ i cal “mous tache-like” pat tern (Fig. 2B), in which the fron tal ridges (scratches) change their di rec tion from trans verse to di ag o nal with re spect to the long axis of the trace fos sil (with a typ i cal di ver gent an gle of ~50°, Fig. 2A–C) and the rear ones form a nar row in ci sion (acute an gle), cut ting into the smooth trap e -zoidal area be hind the lobes (Figs 2, 3A, B, E, F). The trace fos sil stud ied dis plays sets of lobe-cov er ing ridges, com posed of two ridges (scratches) each. The dis tance be tween the ridges within a set is typ i cally 1–3 mm (Fig. 2D). The ap pear ance of lobecov er -ing ridge sets (scratches) de pends on pres er va tion and lo cally they may ap pear to be com posed of a sin gle ridge (Fig. 2E). A ridge is typ i cally ~1 mm wide (Fig. 3A–C). The ichnospecies is U-shaped in lon gi tu di nal cross-sec tion with the fron tal and rear parts fused with the over lay ing bed. This ichnospecies dis plays sig nif i cant vari a tion in pres er va tion style and the ob served range has been il -lus trated in Fig ures 2 and 3.
Re marks: In ear lier pa pers, spec i mens from Wiœniówka Wielka
site as signed herein to Rusophycus ramellensis Legg, were re -ferred to as Cruziana barbata Seilacher, 1970 by Or³owski (1992: p. 24, fig. 7.) The lat ter ichnospecies was cre ated by Seilacher (1970) on the ba sis of ma te rial from the Cam brian Se ries 3 of Spain. Later, Legg (1985) re lo cated the short Rusophycus-like vari ants of this ichnospecies to Rusophycus ramellensis Legg. The char ac ter is tic fea ture of Rusophycus ramellensis Legg is its di ver -gent pat tern of endopodal scratches, which has been re ferred to as a “mous tache-like” pat tern by Seilacher (1970). How ever, in Rusophycus ramellensis Legg the ce phalic im prints do not con tact di -rectly the endopodal lobes and in this ichnospecies, the ex tent of the endopodal lobes is not re stricted to the ce phalic area in the la-teral view, and the smooth trap e zoidal ta per ing back wards area is miss ing (Seilacher, 1970; Legg, 1985; Fig. 3D). The di ver gent pat -tern of endopodal scratches oc curs also in Rusophycus dispar Lin-narsson. How ever, in Rusophycus dispar the endopodal scratches are not re stricted to ce phalic area.
STRATI GRAPHI CAL DIS TRI BU TION
Rusophycus inexpectus isp. nov. was re cov ered from
the lower part of the pro
file of the Wiœniówka Du¿a
(Wielka) Quarry (south ern wall, Figs 1C, 2A, B, 3E, F).
Or³owski (1992) found in this in ter val an undertrace ver sion
of the trace fos sil, as signed by him to Cruziana barbata
Fig. 2. Rusophycus inexpectus isp. nov. from the Wiœniówka Sand stone For ma tion. Sym bols used on fig ures: esad – endopodal scratches an te ri orly di rected, espd –endopodal scratches pos te ri orly di rected, esapd – endopodal scratches an te ri orly to pos te ri orly di -rected, esatd – endopodal scratches an te ri orly di rected to trans verse, est endopodal scratches trans verse, ci – cephalon im print, Ri1 – first Rusophycus inexpectus, ?c/?p – ?ce phalic/?pleu ral im prints, el – endopodal lobe, sa – smooth area, exa – exopodal area, pi – pleurae im prints, cmi1, 2, 3 – ce phalic mar gin im print num ber 1, 2, 3. A. Shal low ex pres sion of the trace fos sil (ZPAL Tf. 4/1335). B. Deeper ex pres -sion of the trace fos sil (typ i cal “mous tache-like” pat tern of scratches; ZPAL Tf. 4/912). C. Full ver -sion of the trace fos sil (ZPAL Tf. 4/1329), note the shal low ver sion of the trace fos sil, lo cated close to the pos te rior mar gin of this spec i men (Ri1). D, E. Two shal low ex -pres sions of the trace fos sil (D: ZPAL Tf. 4/1331; E: ZPAL Tf. 4/1023).
Seilacher. How ever, in this study, a fully pre served
Ruso-phycus inexpectus isp. nov. also was re cov ered, in ad di tion
to the com mon est undertrace vari ant of Rusophycus
inex-pectus isp. nov., which is com pa ra ble to a typ i cal undertrace
of Rusophycus ramellensis Legg. Fig
ures 2C and 3A–C
show this full style of pres er va tion (see also Fig. 3G).
Rusophycus inexpectus isp. nov. was re cov ered also from the up
-per most part of the pro file (north ern wall, Figs 1B, 2C, D,
3A–C). This may con firm pre vi ous un doc u mented in for ma
-tion on the pres ence in the up per most Furongian part of the
pro file (Cruziana barbata Seilacher in ¯yliñska et al., 2006
and ref er ences therein) of a trace fos sil re sem bling
Ruso-phycus ramellensis Legg.
COM PAR I SON WITH RUSOPHYCUS
RAMELLENSIS LEGG
Rusophycus ramellensis, con sid ered as the in dex trace
fos sil for the Cam brian Se ries 3, is known from lo ca tions,
rep re sent ing the an cient shelves of Gond wana (Seilacher,
1970, 2007). The most com mon mode of oc cur rence is as a
deep undertrace, where only the endopodal scratches with
their char ac ter is tic “mous tache’-like” pat tern are pre served
(Seilacher, 1970). This type of oc cur rence, due to pres er va
-tion bi ases, is also most typ i cal of Rusophycus inexpectus
isp. nov. (Figs 2A, B, D, E, 3G). The more com plete
mor-phologies of Rusophycus ramellensis (Seilacher, 1985;
Sei-lacher, 2007; Fig ure 3D) dis play wide, exopodal, brushed
zones at the sides of the trace fos sil (Fig. 3D), in con trast to
ma te rial of Rusophycus inexpectus isp. nov. from the
Wiœ-niówka Sand stone For ma tion. The dif fer ence be tween these
trace fos sils in di cates that their trace mak ers prob a bly dif
-fered in:
1) the pres ence/ab sence or the size of exopodites; there
is no ev i dence for their pres ence in trace fos sils from the
Wiœniówka Sand stone For ma tion. The exopodal traces are
usu ally rec og nized on the ba sis of a reg u lar, par al lel pat tern
of scratches (Bergström, 1972, 1976; see Fig. 4C). No such
par al lel scratches are ob served in the ma te rial from the Wiœ-
niówka Sand stone For ma tion. This might be a
taphonomyre lated fea tutaphonomyre, but the fact that the ma te rial stud ied ap
-peared to be gen er ally well pre served may im ply that there
was an ac tual exopod size dif fer ence that could be re lated to
tax o nomic dif fer ences be tween the trace mak ers;
2) the de gree of trace maker tagmosis, re flect ing the de
gree of body dif fer en ti a tion into mor pho log i cally and func
-tion ally unique units, as re corded in Rusophycus
ramellen-sis Legg, is low as the change in ap pend age size (exopods
vs. endopods) was only in their pro por tions, as was
propo-sed by Seilacher (1985). The ma te rial from the Wiœniówka
Sand stone For ma tion shows that Rusophycus inexpectus
isp. nov. has endopodal scratches, which are re stricted to the
zone ad ja cent to the ce phalic im prints (see the lat eral view
in Fig. 3B, C). This may in di cate that the “ce phalic” part
was the “tagma” with the endopods. The smooth pos te rior
of Rusophycus inexpectus isp. nov. (Fig. 3) im plies an ab
-sence of ap pend ages, or a dif fer ent struc ture of them.
Tri-lobites do not show such tagmosis (Har ring ton, 1959; Berg-
ström, 1972; Whit ting ton, 1980; Hughes, 2003a, b), The
con trast with tri lo bite-made trace fos sils is vis i ble, when it
is com pared with Rusophycus arizonensis Seilacher, 1970
(Cam brian Se ries 3; Cruziana arizonensis in Seilacher,
1970), which bears dis tinct im prints of the ce phalic mar gin
(Fig. 4). In con trast with the Furongian ma te rial from the
Holy Cross Moun tains, Rusophycus arizonensis has
endo-podal scratches that ex tend be yond the rear mar gin of the
caphalic im print (Fig. 4B, D). This is a tagmosis pat tern ex
-pected from a tri lo bite trace maker, as a dor sally ex pressed
sub di vi sion does not cor re spond with ap pend age dif fer en ti
-a tion (cf. Hughes, 2003-a, b). There fore, Rusophycus
inex-pectus isp. nov. ap pears to be a good can di date for be ing a
non-tri lo bite trace fos sil, the trace maker of which could
dis play some be hav ioural con ver gence to Rusophycus
ramellensis trace maker, as the pat terns of ap pend age move
-ment ap pear to be sim i lar in both trace fos sils.
DIS CUS SION
Rusophycus inexpectus isp. nov. from the Wiœniówka
Sand stone For ma tion dif fers from the Cam brian Se ries 3
Rusophycus ramellensis Legg and rep re sents a new
ichno-spe cies (Fig. 5). The lim ited biostratigraphic data, col lected
from the up per part of the pro file, in di cates that the trace
fos sil de scribed in the pres ent ac count come from the
Fu-rongian se ries (¯yliñska et al., 2006). There fore, cur rently
there is no ev i dence that the lower part of the pro file ex
-posed at Wiœniówka rep re sents the Cam brian Se ries 3 se ries
(see Or³owski, 1992; Kowalczewski, 1995).
Rusophycus ramellensis Legg is still con sid ered to be a
Cam brian Se ries 3 in dex fos sil, but the pres ence of be hav
-ioural con ver gence with Rusophycus inexpectus must be
kept in mind, when deal ing with undertrace ma te rial. The
sim i lar ity be tween these two ichnospecies is a re sult of an
undertrace de fi ciency in mor pho log i cal de tails (cf.
Seila-cher, 1970, 2007).
Tak ing into con sid er ation the endemicity of the
Furon-gian Wiœniówka Sand stone For ma tion tri lo bite fauna
(¯y-liñska, 2002), it may well ap pear that sim i lar endemicity is
char ac ter is tic of the ar thro pods, re spon si ble for Rusophycus
in the re gion, and that the po ten tial for ap ply ing the
Cruziana ichnostratigraphy, an ichnostratigraphical scheme ap
-plied to Gond wana, is locally limited.
Trace fos sils from the Wiœniówka Du¿a Quarry (mainly
the pres ence of Cruziana semiplicata and Cruziana
bar-bata), were used as a one of the sup port ive ar gu ments for
link ing the £ysogóry Block, the struc tural unit form ing the
north ern part of the Holy Cross Moun tains (see Be³ka et al.,
2000; Jaworowski and Sikorska, 2006), with Gond
wana
(Seilacher, 2007). How ever, Cruziana semiplicata has been
re ported from other nonGondwanan ar eas (see the most re
cent over view in Jensen et al., 2011). There fore, the sig nif i
-cance of this trace for palaeogeographic re con struc tions is
lim ited. The dif fer ence be tween the Holy Cross Moun tains
ma te rial and the typ i cal Cam brian Se ries 3 Rusophycus
ramellensis Legg, as noted in the pres ent pa per, ap pears to
pre clude the pos si bil ity of ap ply ing the ma te rial from the
Wiœniówka Du¿a Quarry in palaeo geo graphi cal re con struc
-tion.
CON CLU SIONS
1) The pres ence of Rusophycus ramellensis Legg in the
Wiœniówka Sand stone For ma tion (re ported as Cruziana
barbata Seilacher by ear lier au thors) has not been
confir-med. The ma te rial rep re sents a new ichnospecies
Rusophy-cus inexpectus isp. nov.
2) Rusophycus inexpectus isp. nov. dif fers from the Cam
-brian Ser ies 3 Rusophycus ramellensis Legg: a) in hav ing a
smooth trap e zoidal area be hind the lobes; b) in the pres ence
of a di rect con tact be tween the endopodal lobes and the ce
phalic mar gin im prints; and c) in the re stric tion in the oc cur
-rence of endopodal lobes and scratches to the ce phalic re gion
of the trace fos sil. These mor pho log i cal dif fer ences be tween
the trace fos sils are rooted in dif fer ent body plans of the trace
mak ers, as ev i denced by well pre served spec i mens.
3) Rusophycus inexpectus isp. nov. ap pears to rep re sent
a trace fos sil prob a bly pro duced by a non-tri lo bite trace
maker, as in di cated by the non-tri lo bite-like ap pend age
tag-mosis pat tern.
RUSOPHYCUS INEXPECTUS ISP. NOV. FROM THE FURONGIAN
149
Fig. 3. More com plete mor pho log i cal ver sion of Rusophycus inexpectus isp. nov. from the Wiœniówka Sand stone For ma tion and com -par i son with typ i cal Mid dle Cam brian Rusophycus ramellensis Legg (Cruziana barbata Seilacher) A. Well pre served spec i men of the new ichnospecies (ZPAL Tf. 4/1328). B. Well pre served ver sion of the new ichnospecies, the same spec i men as in Fig. 2C, here a view from dif fer ent an gle shows ad di tional de tails (ZPAL Tf. 4/1329). C. The same spec i men as in A, here in a side view. Note an ap par ent tagmosis: endopodal lobes have ex tent com pa ra ble to cephalon mar gin. D. Full ver sion of typ i cal Mid dle Cam brian Rusophycus ramellensis Legg (Cruziana barbata Seilacher; see Seilacher, 1985, 2007; GPIT/IC/00147). E, F. Full ver sion of Rusophycus inexpectus isp. nov. (ZPAL Tf. 4/674). G. Sche matic il lus tra tion show ing cor re spon dence be tween depth of pres er va tion and undertrace de fi ciency: (a) the shal low est ver sion, (b) deeper ver sion and (c) the full, well pre served ver sion. Sym bols used: see cap tion of Fig ure 2.
Fig. 4. Rusophycus arizonensis (Seilacher) (Cruziana arizonensis Seilacher, 1970) dis play ing tri lo bite-like pat tern of tagmosis. A. Trace fos sils with all endopodal scratches di rected pos te ri orly; GPIT/IC/00149. B–D. Well pre served trace fos sils with ce phalic im prints (var i ous views of the same spec i men; GPIT/IC/00150). Note that endopodal scratches ex tend be yond rear mar gin of ce phalic im print (B and D: shown by dashed line). Sym bols used: see cap tion of Fig ure 2.
4) On the ba sis of avail able biostratigraphical data, the
Rusophycus inexpectus isp. nov. ap pears to come from the
Furongian strata (see ¯yliñska et al., 2006 and ref er ences
therein). There is no ev i dence at pres ent that the lower part
of the pro file ac ces si ble at the Wiœniówka Du¿a (Wielka)
Quarry rep re sents the Cam brian Series 3.
5) In gen eral, the ap pli ca bil ity of Rusophycus ramellen-
sis Legg to ichnostratigraphical sub di vi sion and
palaeogeo-graphic re
con
struc
tion is in need of care
ful re
ap
praisal,
since the pres ence of mor pho log i cally con ver gent non-trilo-
bite trace fos sils rep re sent ing Rusophycus inexpectus isp.
nov. also can not be ex cluded in other assemblages.
Ac knowl edge ments
I would like to thank Philipe Havlik (Eberhard Karls Uni ver -sity, Tübingen, Ger many) for his help dur ing my visit to Tübingen (Ger many) and my study of the rich tri lo bite trace fos sils ma te rial stored by the Uni ver sity. I would like to thank re view ers of ASGP: Duncan McIlroy (Me mo rial Uni ver sity, St. John’s, Can ada), Carlos Neto de Carvalho (Geopark Naturtejo Meseta Me rid i o nal, Idanha-a-Nova, Por tu gal) and Michal Stachacz (Jagiellonian University, Kraków, Po land) for their crit i cal and valu able re marks and lin guis tic im prove ments. I would like to thank also Al fred Uchman (Jagiellonian Uni ver sity, Kraków, Po land) for his re marks and help. I am grate ful to Frank Simpson (Wind sor, Canada) for lin guis tic im prove ments. This study is a part of a Ph.D. pro ject su per -vised by Marcin Machalski at the In sti tute of Paleobiology of the Pol ish Acad emy of Sci ences. It was fi nanced by a grant from the Min is try of Sci ence and Higher Ed u ca tion of Po land in years 2010 to 2012 (no. N N307 243539) to the Au thor and, par tially, by the In sti tute of Paleobiology of the Pol ish Acad emy of Sci ences.
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