2021 | 74/2 | 121–126 | 4 Figs. | 2 Tabs. | www.geologia-croatica.hr Journal of the Croatian Geological Survey and the Croatian Geological Society 1. INTRODUCTION The ichnogenus Cardioichnus SMITH & CRIMES, 1983 embraces heart-shaped structures interpreted as a resting trace (cubichnia), produced by deposit-feeding, spatangoid irregular echinoids (known as heart urchins or potatoes) (SMITH & CRIMES, 1983; GIBERT & GOLDRING, 2008). It is common in deep- and shallow -marine deposits throughout the Meso-Cenozoic, from the Kimmeridgian (BOUCHEMLA et al., 2020) to the Pleistocene (MAYORAL et al., 2013; CARUSO & MONACO, 2015). This trace fossil is preserved as a convex or concave form, ovoid or sub- quadrate in shape, in positive hyporelief or negative epirelief on sandstone beds, and formed by two symmetrical broad lateral lobes connected at the depressed V-shaped region (SMITH & CRIMES, 1983; GIBERT & MARTINELL, 1995; KAPPUS & LUCAS, 2019). Five ichnospecies of Cardioichnus are distinguished, includ- ing C. planus SMITH & CRIMES, 1983, the type ichnospecies; C. ovalis SMITH & CRIMES, 1983; C. foradadensis PLAZIAT & MAHMOUDI, 1988; C. reniformis MAYORAL & MUÑIZ, 2001 and C. biloba KAPPUS & LUCAS, 2019. Furthermore, this ichnogenus is found in association with locomotion traces, and it may co-occur with Scolicia (MAYORAL & MUÑIZ, 2001; UCHMAN, 2007a), and Bichordites (BERNARDI et al., 2010; MAYORAL et al., 2013; KAPPUS & LUCAS, 2019). Here, the ichnospecies C. planus is recorded for the first time from Algeria and in the African continent. The interpretation of abundant C. planus from the lower Miocene Tiaret Marl Forma- tion in western Algeria is presented and discussed. Furthermore, the spatangoid echinoid Echinocardium nummuliticum PÉRON & GAUTHIER, 1885 has been proposed as the probable producer of the studied C. planus specimens. Finally, a stratigraphic, pala- eogeographic and palaeoenvironmental review of Cardioichnus and its ichnospecies is presented. 2. GEOLOGICAL SETTING AND LOCALITY In the northwesternmost part of Algeria, the lower Chelif (e.g., NAIMI & CHERIF, 2021a; NAIMI et al., 2020), Tafna (e.g., NAIMI The trace fossil Cardioichnus planus from the lower Miocene of Algeria: the first record from Africa and a probable endemic tracemaker Mohammed Nadir Naimi1, Amine Cherif1,* and Mourad Belaid2 1 Université de Kasdi Merbah, Département des Sciences de la Terre et de l'Univers, FHERSTU, Ghardaïa Road, Ouargla, 30000, Algeria; (*corresponding author: mohammednadirnaimi@gmail.com (Mohammed Nadir Naimi); acherif11@gmail.com (Amine Cherif)) 2 Université de Kasdi Merbah, Laboratoire des Réservoirs Souterrains, Pétroliers, Gaziers et Aquifères, Ghardaïa Road, Ouargla, 30000, Algeria; (mouradgeo14@gmail.com) doi: 10.4154/gc.2021.09 Abstract Abundant Cardioichnus planus SMITH & CRIMES, 1983 have been reported for the first time from Africa. They occur in lower Miocene deep-marine deposits near Tiaret in northwestern Al- geria. Morphological features of the trace, coupled with the spatial interrelationships between Cardioichnus and the echinoids documented in this area, provide evidence that the endemic spatangoid Echinocardium nummuliticum PÉRON & GAUTHIER, 1885 represents the trace- maker of the studied C. planus. Cardioichnus is known from the upper Jurassic to the Pleisto- cene and it has a wide geographic and environmental distribution. et al., 2021), the Tiaret (CHERIF et al., 2021) Neogene basins are located. They belong to the Tell Atlas (NAIMI & CHERIF, 2021b). The studied material comes from the Tiaret Marl Formation (CHERIF et al., 2021) of Sidi Ali Mellal section in the Tiaret basin (POLVÊCHE, 1960). The section is located some kilometeres to the northwest of the town of Tiaret (Fig. 1A–C). The lower Miocene Tiaret Marl Formation crops out widely in the Tiaret area. It overlies Visean rhyolites (POLVÊCHE, 1960), or Jurassic marine deposits (CHERIF et al., 2015, 2018; HALAMSKI & CHERIF, 2017; BELAID et al., 2020), and is overlain by the upper Miocene Tiaret Sandstone Formation (CHERIF et al., 2021). The Tiaret Marl Formation has been attributed to the lower Miocene on the basis of a foraminiferal assemblage including Cibicides mexicanus, C. pseudoungerianus, Globigerinoides sac­ culiferus irregularis, G. triloba, Nonion pompilioides, Orbulina universa, Pullenia bulloides and Uvigerina rustica (POLVÊCHE, 1960; CHERIF et al., 2021). In Sidi Ali Mellal section, the Tiaret Marl Formation overlies the Tardi-Hercynian rhyolites and it has been subdivided into two main units (CHERIF et al., 2021). Ichnological and sedimentological studies allowed the attri- bution of this formation to a deep-sea fan system, dominated by graphoglyptid trace fossils which characterize the turbidite suc- cessions and the Nereites ichnofacies (CHERIF et al., 2021). Moreover, these trace fossils have been subdivided into three ich- noassemblages corresponding to the Nereites, Paleodictyon, Ophiomorpha rudis ichnosubfacies. The trace fossil assemblage reported from this formation includes Cardioichnus isp., cf. Chondrites isp., Cosmorhaphe isp., Desmograpton isp., Gordia isp., Helminthorhaphe isp., Helminthopsis isp., cf. Megagrapton isp., Nereites isp., Ophiomorpha isp., O. annulata, O. rudis, Pala­ eophycus isp., Paleodictyon majus, P. maximum, P. minimum, P. miocenicum, P. strozzii, cf. Planolites isp., P. montanus, Scoli­ cia isp., S. prisca, S. vertebralis, Tisoa siphonalis, Tubulichnium cf. incertum, Urohelminthoida appendiculata and Zoophycos isp. In the Sidi Ali Mellal section, the ichnoassemblage of the first unit contains the trace fossil Cardioichnus (Fig. 1D). Article history: Manuscript received November 17, 2020 Revised manuscript accepted March 05, 2021 Available online June 30, 2021 Keywords: Cardioichnus planus, resting trace, Echinocardium nummuliticum, lower Miocene, Algeria G eo lo gi a C ro at ic a Geologia Croatica 74/2122 Figure 1. A) Location of Orania (Northern Algeria) in the Western Mediterranean. B) The main structural domains of Western Algeria. C) Geological map of the study area. Table 1. Length, width and l/w ratio of the studied 65 Cardioichnus planus (in mm). Sample Length Width L/W Ratio Sample Length Width L/W Ratio CAR-SAM-001 24 21 1.14 CAR-SAM-034 23 19 1.21 CAR-SAM-002 24 18 1.33 CAR-SAM-035 28 22 1.27 CAR-SAM-003 23 16 1.43 CAR-SAM-036 26 20 1.3 CAR-SAM-004 26 24 1.08 CAR-SAM-037 32 24 1.33 CAR-SAM-005 21 20 1.05 CAR-SAM-038 27 24 1.12 CAR-SAM-006 27 22 1.22 CAR-SAM-039 25 22 1.13 CAR-SAM-007 26 22 1.18 CAR-SAM-040 27 24 1.12 CAR-SAM-008 27 22 1.22 CAR-SAM-041 26 23 1.13 CAR-SAM-009 23 19 1.21 CAR-SAM-042 33 25 1.32 CAR-SAM-010 21 20 1.05 CAR-SAM-043 24 21 1.14 CAR-SAM-011 24 20 1.2 CAR-SAM-044 37 25 1.48 CAR-SAM-012 22 20 1.1 CAR-SAM-045 25 19 1.31 CAR-SAM-013 26 22 1.18 CAR-SAM-046 32 24 1.33 CAR-SAM-014 23 19 1.21 CAR-SAM-047 26 22 1.18 CAR-SAM-015 23 16 1.43 CAR-SAM-048 32 26 1.23 CAR-SAM-016 21 18 1.16 CAR-SAM-049 26 21 1.23 CAR-SAM-017 21 18 1.16 CAR-SAM-050 22 18 1.22 CAR-SAM-018 22 20 1.1 CAR-SAM-051 24 19 1.26 CAR-SAM-019 27 22 1.22 CAR-SAM-052 25 23 1.08 CAR-SAM-020 27 20 1.35 CAR-SAM-053 25 18 1.38 CAR-SAM-021 22 18 1.22 CAR-SAM-054 21 18 1.16 CAR-SAM-022 25 21 1.19 CAR-SAM-055 30 26 1.15 CAR-SAM-023 21 17 1.23 CAR-SAM-056 27 21 1.28 CAR-SAM-024 28 21 1.33 CAR-SAM-057 29 23 1.26 CAR-SAM-025 24 21 1.14 CAR-SAM-058 33 25 1.32 CAR-SAM-026 23 17 1.35 CAR-SAM-059 30 21 1.42 CAR-SAM-027 24 17 1.41 CAR-SAM-060 22 21 1.04 CAR-SAM-028 22 19 1.15 CAR-SAM-061 31 25 1.24 CAR-SAM-029 24 21 1.14 CAR-SAM-062 25 23 1.08 CAR-SAM-030 22 18 1.22 CAR-SAM-063 30 22 1.36 CAR-SAM-031 27 20 1.35 CAR-SAM-064 24 19 1.26 CAR-SAM-032 25 22 1.13 CAR-SAM-065 26 21 1.23 CAR-SAM-033 25 20 1.25 Average 25.58 20.87 1.22 G eologia C roatica Naimi et al.: The trace fossil Cardioichnus planus from the lower Miocene of Algeria: the first record from Africa and a probable endemic tracemaker 123 3. MATERIAL AND METHODS More than seventy specimens of C. planus have been observed and collected by the authors during several field trips in to the Tiaret basin between 2019 and 2020. The studied collection (N = 65) is deposited at the Laboratory of Geology of Sahara at Kasdi Merbah Ouargla University (CAR-SAM-001 to CAR-SAM-065). These sixty-five specimens were selected because of their pre- servation. They were photographed both in the field and the labo- ratory. The measurements were taken in the laboratory (Table 1). Also, the ichnotaxonomic identification of C. planus is in accor- dance with SMITH & CRIMES (1983). 4. RESULTS 4.1. Description Cardioichnus planus is very abundant in the Sidi Ali Mellal sec- tion. It is a hypichnion in the sandstone beds, and co-occurs with Scolicia vertebralis (Fig. 2). The studied traces (Fig. 3A-F) are heart-shaped mounds. They are ovoid or subquadratic in outline, bilobate and bilaterally symmetrical. Their two lateral lobes are generally rounded, curved, and merged at the central axial depressed V-shaped de- pression. Some specimens show a well developed posterior im- pression with a sharp edge at the terminus, which extends from the V-shaped depression, and separates the two distinct lobes. The mounds are 20–37 mm long and 16–26 mm wide, with a length/width ratio of about 1.05–1.42. The mean length (N = 65) is 25.58 mm, whereas the mean width is 20.87 mm (Table 1). 4.2. Remarks The selected samples have been attributed to Cardioichnus planus due to their dimensions (l= 20–37 mm; w= 16–26 mm), and the length/width ratio which are similar to that described by SMITH & CRIMES (1983) and BRUSTUR (2005). The ovoid to subquadratic form of the studied specimens, their preservation as a convex body and the presence of two curved lateral lobes merged at the median longitudinal V-shaped depression and pos- terior impression characterize C. planus (SMITH & CRIMES, 1983; UCHMAN, 2007a), and confirm their determination as this ichnospecies. 5. DISCUSSION 5.1. Probable local tracemakers of the Algerian C. planus Spatangoid echinoderms or heart urchins are known from the early Jurassic (BUATOIS & MÁNGANO, 2018). They are the producers of Bichordites, Cardioichnus and Scolicia. Bichordites and Scolicia are attributed to the burrowing of spatangoids. Scoli­ cia is the best known among these ichnogenera. Its oldest occur- rence is from the Tithonian of Bulgaria (TCHOUMATCHENCO & UCHMAN, 2001). All Cardioichnus ichnospecies are considered to be resting traces produced by spatangoid echinoids (SMITH & CRIMES, 1983; PLAZIAT & MAHMOUDI, 1988; MAYORAL & MU- ÑIZ, 2001; KAPPUS & LUCAS, 2019). The determination of tracemaker is problematic. In the Oligocene of Trentino (north- eastern Italy), Cardioichnus isp. and the associated Bichordites have been attributed to Eupatagus ornatus (BERNARDI et al., 2010). Furthermore, Bichordites recorded from the middle Mio- cene of Spain has been attributed to the grazing activity of spa- tangoid echinoids assigned to the genus Maretia (GIBERT & GOLDRING, 2008). Recently, the spatangoid Heteraster has been considered as the tracemaker of the ichnospecies C. biloba, described from the mid-Cretaceous of New Mexico in the United States of America (KAPPUS & LUCAS, 2019). The studied outcrop belongs to the southern Tellian border of the Ouarsenis range which constitutes the northern edge of the Tiaret Miocene basin. The most important palaeontological in- Figure 2. The stratigraphic succession and vertical distribution of trace fossils of the lower Miocene Tiaret Marls Formation in the Sidi Ali Mellal section. Figure 3. Cardioichnus planus from the lower Miocene of the Sidi Ali Mellal sec- tion. A) CAR-SAM-043. B) CAR-SAM-044. C) CAR-SAM-063. D) CAR-SAM-055. E) CAR-SAM-056. F) CAR-SAM-057 (scale: 1 cm). G eo lo gi a C ro at ic a Geologia Croatica 74/2124 vestigations which yielded a rich echinoid fauna in the southern border of the Ouarsenis range have been carried out at the Kef Ighoud section, located 50 km to the ENE of our study area. Kef Ighoud echinoid-rich deposits have been attributed to the Eocene (POMEL, 1885), the Oligocene (DALLONI, 1936; MATTAUER, 1958) and finally to the lower to middle Miocene on the basis of their planktonic foraminifera (DERKAOUI, 2017). Several spatangoid echinoids have been described from the lower-middle Miocene of the Kef Ighoud section (COTTEAU et al., 1885; POMEL, 1885). The most important ones are Echino­ cardium dubium PÉRON & GAUTHIER, 1885; E. nummuliticum PÉRON & GAUTHIER (Fig. 4), 1885; Euspatangus cruciatus PÉRON & GAUTHIER, 1885; E. hangenmulleri PÉRON & GAUTHIER, 1885; E. subrostratus PÉRON & GAUTHIER, 1885; Pericosmus nicaisei POMEL, 1885; P. subœquipetalus POMEL, 1885; Sarsella mauritanica POMEL, 1885; Schizaster mac carthyi POMEL, 1885; Spatanus (Pseudopatagus) cruciatus POMEL, 1885 and Tuberaster tuberculatus PÉRON & GAUTHI ER, 1885. Some taxa such as S. mauritanica and T. tuberculatus have been placed into the spatangoid genus Hemipatagus (KROH, 2007). This echinoid fauna is important due to its endemism, and has never been reported out of this area. Probable producers of Cardioichnus are proposed on the ba- sis of the similarities in size and morphology between the spa- tangoids and Cardioichnus (KAPPUS & LUCAS, 2019). How- ever, among the spatangoid assemblage of Kef Ighoud, three species have similar sizes to the studied Cardioichnus specimens. They include Echinocardium dubium (l= 26 mm; w= 25 mm), E. nummuliticum (l= 26 mm; w= 23 mm) and Euspatangus hangen­ mulleri (l= 25; w= 25). According to the l/w ratio, the two Echi­ nocardium species resemble Cardioichnus planus. However, the morphology of Echinocardium nummuliticum allows proposing it as the probable tracemaker (Fig. 4). The size decreases near the posterior part of the test where a prominent rostrum is present. Accordingly, the posterior impression at the terminus of the trace could be interpreted as the imprint of this rostrum. Alternatively, the ambitus of this species is heart-shaped, whereas it is rounded in E. dubium. This confirms that C. planus most closely resem- bles E. nummuliticum in size and morphology. Echinocardium species occur generally in shallower envi- ronments. POMEL (1887) indicated that the attribution of E. nummuliticum to this genus by PÉRON & GAUTHIER is incor- rect, and these specimens should be revised. However, it co-oc- curs with other sea urchins such as Schizaster (POMEL, 1885) Figure 4. Echinocardium nummuliticum PÉRON & GAUTIER (1885) holotype, MNHN.F.J01420. Photography Peter Massicard, program RECOLNAT (ARN-11-IN- BS-0004). A) Aboral view. B) Oral view. C) Posterior view. D) Side view. E) Anterior view (scale: 0.5 cm). Table 2. Stratigraphic and geographic record of the ichnogenus Cardioichnus. Age Ichnotaxa Country References Upper Jurassic Cardioichnus isp. Algeria BOUCHEMLA et al. (2020) Lower Cretaceous Cardioichnus isp. Spain MONACO et al. (2005); GIANNETTI et al. (2014) C. biloba United States of America KAPPUS & LUCAS (2019, 2020) C. foradadensis United States of America KAPPUS & LUCAS (2020) Upper Cretaceous C. planus Poland SMITH & CRIMES (1983) C. planus Spain SMITH & CRIMES (1983) C. planus Italy UCHMAN (2007a) Cardioichnus isp. Poland KSIĄŻKIEWICZ (1977); RAJCHEL & UCHMAN (2012) Cardioichnus isp. Iran BAYET-GOLL et al. (2016) Paleocene Cardioichnus isp. Italy UCHMAN (2007b) Eocene C. ovalis Switzerland SMITH & CRIMES (1983) C. foradadensis Spain PLAZIAT & MAHMOUDI (1988) C. ovalis Romania BRUSTUR (1996) C. cf. planus Romania BRUSTUR (1996) Cardioichnus isp. Romania BUATOIS et al. (2001) Cardioichnus isp. Argentina LÓPEZ-CABRERA et al. (2008) Cardioichnus isp. Italy MONACO et al. (2017) Oligocene Cardioichnus isp. Italy BERNARDI et al. (2010) Oligocene/Miocene C. planus Romania BRUSTUR (2005) Cardioichnus isp. Tunisia RIAHI et al. (2014) Miocene Cardioichnus isp. New Zealand MANLEY & LEWIS (1998) C. reniformis Spain MAYORAL & MUÑIZ (2001) C. planus Spain MAYORAL & MUÑIZ (2001) Cardioichnus isp. Spain MAYORAL & MUÑIZ (2001) Cardioichnus isp. Spain GIBERT & ROBLES (2005) Cardioichnus isp. Argentina LÓPEZ-CABRERA et al. (2008) Cardioichnus isp. Algeria CHERIF et al. (2020) Cardioichnus planus Algeria This paper Pliocene Cardioichnus isp. Spain GIBERT & MARTINELL (1995) C. planus Spain GIBERT & MARTINELL (1999) Pleistocene Cardioichnus isp. Cape Verde MAYORAL et al. (2013) Cardioichnus isp. Italy CARUSO & MONACO (2015) G eologia C roatica Naimi et al.: The trace fossil Cardioichnus planus from the lower Miocene of Algeria: the first record from Africa and a probable endemic tracemaker 125 which could be found in deep basins (MCKINNEY, 1986). Also, echinoids of the Echinocardium group are responsible for traces with one drain such as Bichordites, which could be continuous with Cardioichnus and produced by the same organism. Bichordi­ tes occurs from shallow-marine settings (UCHMAN & KREN- MAYR, 1995; BERNARDI et al., 2010; CARUSO & MONACO, 2015) to slope deposits in soft substrates (VILLEGAS-MARTÍN et al., 2014). 5.2. Stratigraphy, palaeoenvironments and palaeogeography The oldest Cardioichnus were observed in the Kimmeridgian of northwestern Algeria, near this study area (BOUCHEMLA et al., 2020) (Table 2). It occurs in the lower to transitional offshore en- vironment characterizing the archetypal Cruziana ichnofacies. This area belongs to the south Tethyan Ocean. This constitutes the only record of this ichnogenus in the Jurassic. In the lower Cretaceous, Cardioichnus occurred in similar environments to those of the upper Jurassic of Algeria. It has been reported in seve ral domains from Spain (Tethyan Realm) (MONACO et al., 2005; GIANNETTI et al., 2014) to North America (KAPPUS & LUCAS, 2019, 2020). Cardioichnus is common in flysch deposits from as early as the upper Cretaceous. It has been found in Iran, Italy, Poland and Spain (KSIĄŻKIEWICZ, 1977; SMITH & CRIMES, 1983; UCHMAN, 2007a; RAJCHEL & UCHMAN, 2012; BAYET-GOLL et al., 2016). The Cenozoic records seem to be more numerous. This ichnogenus is known from the Palaeo- cene siliciclastic flysch deposits of Italy (UCHMAN, 2007b). It had a larger distribution during the Eocene, generally in the cir- cum-Tethyan region (SMITH & CRIMES, 1983; PLAZIAT & MAHMOUDI, 1988; BRUSTUR, 1996; BUATOIS et al., 2001; MONACO et al., 2017) and South American (LÓPEZ-CA- BRERA et al., 2008) deep-sea systems, within ichnoassemblages assigned to the Zoophycos and Nereites ichnofacies. Cardioich­ nus also has been reported from the Oligocene and the Oligo- Miocene of the Mediterranean Tethys in shallow (BERNARDI et al., 2010) to deep-sea deposits (BRUSTUR, 2005; RIAHI et al., 2014). Miocene records of Cardioichnus are widespread, e.g., it occurs in New Zealand (MANLEY & LEWIS, 1998), Argen- tina (LÓPEZ-CABRERA et al., 2008), and the western Mediter- ranean basin, especially in Spain (MAYORAL & MUÑIZ, 2001; GIBERT & ROBLES, 2005) and Algeria (CHERIF et al., 2021; this paper). Miocene Cardioichnus have been reported from in- fralittoral shelf sands to deep-sea deposits. In the Pliocene, Car­ dioichnus is known only in Spain from sandy storm beds, typical of the Cruziana ichnofacies (GIBERT & MARTINELL, 1995, 1999). Finally, the newest records of this trace fossil are from the Pleistocene of the Tropical Eastern Atlantic (Cape Verde), in a moderate to high-energy shallow marine environment, where it constitutes part of the proximal Cruziana ichnosubfacies (MAY- ORAL et al., 2013), and from the shallow deposits of southern Italy (CARUSO & MONACO, 2015). 6. CONCLUSIONS The studied samples allow the following conclusions to be drawn: Abundant Cardioichnus planus has been found for the first time in Africa; The studied specimens are dated as being of the lower Mio- cene age, and co-occur with Scolicia vertebralis; The endemic spatangoid echinoderm Echinocardium num­ muliticum PÉRON & GAUTHIER, 1885 represents the probable tracemaker of the studied C. planus. ACKNOWLEDGMENT We thank the managing editor of Geologia Croatica T. FLUKSI and the reviewers A. UCHMAN (Jagiellonian University, Poland) and O. VINN (University of Tartu, Estonia) for their useful com- ments that have contributed to improving the original manuscript. The authors thank J. FALCONNET (MNHN of Paris, France), who permitted us the use of the E. nummnliticum holotype pho- tography. REFERENCES BAYET-GOLL, A., MONACO, P., JALILI, F. & MAHMUDY-GHARAIE, M.-H. (2016): Depositional environments and ichnology of Upper Cretaceous deep-ma- rine deposits in the Sistan Suture Zone, Birjand, Eastern Iran.– Cretaceous Re- search, 60, 28–51. doi: 10.1016/j.cretres.2015.10.015 BELAID, M., CHERIF, A., VINN, O. & NAIMI, M.N. (2020): First record of trace fos- sils from the Oxfordian Argiles rouges de Kheneg Formation (Tiaret, northwestern Algeria).– Geologia Croatica, 73/2, 85–94. doi: 10.4154/gc.2020.10 BERNARDI, M., BOSCHELE, S., FERRETTI, P. & AVANZINI, M. (2010): Echinoid burrow Bichordites monastiriensis from the Oligocene of NE Italy.– Acta Palae- ontologica Polonica, 55, 479–486. BOUCHEMLA, I., BENDELLA, M., BENYOUCEF, M., LAGNAOUI, A., FERRÉ, B., SCHERZINGER, A. & BEL HAOUZ, W. (2020): The Upper Jurassic Faïdja For- mation (Northwestern Algeria): Sedimentology, biostratigraphy and ichnology.– Journal of African Earth Sciences, 169, 103874. BRUSTUR, T. (1996): The stages of the paleoichnological studies in Romania.– Geo- Eco-Marina, 2, 205–216. BRUSTUR, T. (2005): The ichnogenus Cardioichnus from the Vineţişu Formation (Up- per Oligocene-Lower Miocene, Romania).– Proceedings of the Romanian Acade- my, Series B, 2, 51–53. BUATOIS, L.A., MÁNGANO, M.G. & SYLVESTER, Z. (2001): A diverse deep-marine ichnofauna from the Eocene Tarcau Sandstone of the Eastern Carpathians, Roma- nia.– Ichnos, 8, 23–62. doi: 10.1080/10420940109380172 BUATOIS, L.A. & MÁNGANO, M.G. (2018): The other biodiversity record: Innova- tions in animal-substrate interactions through geologic time.– GSA Today, 28/10, 4–10. doi: 10.1130/GSATG371A.1 CARUSO, C. & MONACO, P. (2015): Bichordites monastiriensis ichnofabric from the Pleistocene shallow-marine sandstones at Le Castella (Crotone), Ionian Calabria, southern Italy.– Rivista Italiana di Paleontologia e Stratigraphia, 121/3, 381–397. doi: 10.13130/2039-4942/6524 CHERIF, A., BERT, D., BENHAMOU, M. & BENYOUCEF, M. (2015): La formation des Argiles de Saïda (Jurassique supérieur) dans le domaine tlemcenien oriental (Takhemaret, Algérie): données biostratigraphiques, ichnologiques et sédimen- tologiques.– Revue de Paléobiologie, 34/2, 363–384. CHERIF, A., BENYOUCEF, M., FERRE, B. & BENHAMOU, M. (2018): Etude sédi- mentologique et ichnologique de la Formation des Argiles de Saïda (Jurassique supérieur) dans les monts de Frenda (Algérie nord-occidentale).– Revue de Paléo- biologie, 37/1, 121–135. CHERIF, A., NAIMI, M.N. & BELAID, M. (2021): Deep-sea trace fossils and deposi- tional model from the lower Miocene Tiaret Marl Formation (northwestern Alge- ria).– Journal of African Earth Sciences, 175, 104115. doi: 10.1016/j.ja fre- arsci.2021.104115 COTTEAU, G., PERON, A. & GAUTHIER, V. (1885): Echinides fossiles de l’Algérie: terrains tertiaires.– Masson Edition, Paris, 273 p. DALLONI, M. (1936): Matériaux pour l’étude géologique du massif de l’Ouarsenis.– Bulletin du Service de la Carte Géologique de l’Algérie, 2, 1–41. DERKAOUI, S. (2017): Etude paléontologique, ichnologique et sédimentologique de la série Miocène inférieur/moyen de Kef Ighoud (Ouarsenis oriental, Algérie).– Unpubl. MSc Thesis, University of Oran 2, 65 p. GIANNETTI, A., MONACO, P., CORBÍ, H. & SORIA, J.M. (2014): Integrated tapho- nomy in an open-marine platform: The Lower Cretaceous of Sierra Helada (Betic Cordillera, SE Spain).– Cretaceous Research, 51, 274–284. doi: 10.1016/j.cre- tres.2014.07.001 GIBERT, J.M. DE & MARTINELL, J. (1995): Sedimentary substrate and trace fossil assemblages in marine Pliocene deposits in Northeast Spain.– Geobios, 28/supp.1, 197–206. doi: 10.1016/S0016-6995(95)80166-9 GIBERT, J.M. DE & MARTINELL, J. (1999): Proximal-distal variations of trace fossil assemblages in a Pliocene ria, Baix Llobregat, Northeastern Spain.– Revista de la Sociedad Geológica de España, 12/2, 209–214. GIBERT, J.M. DE & ROBLES, J.M. (2005): Firmground ichnofacies recording high- marine flooding events (Langhian transgression, Vallès-Pendès Basin, Spain).– Geologica Acta, 3, 295–305. GIBERT, J.M. DE & GOLDRING, R. (2008): Spatangoid-produced ichnofabrics (Bateig Limestone, Miocene, Spain) and the preservation of spatangoid trace fossils.– Pa- G eo lo gi a C ro at ic a Geologia Croatica 74/2126 laeogeography, Palaeoclimatology, Palaeoecology, 270/3–4, 299–310. doi: 10.1016/j.palaeo.2008.01.031 HALAMSKI, A.T. & CHERIF, A. (2017): Oxfordian brachiopods from the Saïda and Frenda mountains (Tlemcenian Domain, north-western Algeria).– Annales Soci- etatis Geologorum Poloniae, 87, 141–156. doi: 10.14241/asgp.2017.006 KAPPUS, E.J. & LUCAS, S.G. (2019): A New Ichnospecies of Cardioichnus from the Cretaceous (Albian) of New Mexico.– Ichnos, 26, 127–133. KAPPUS, E.J. & LUCAS, S.G. (2020): Ichnology of the Lower Cretaceous (Albian) Mesilla Valley Formation, Cerro de Cristo Rey, southeastern New Mexico, USA.– New Mexico Geology, 42, 3–30. KROH, A. (2007): Hemipatagus, a misinterpreted Loveniid (Echinodermata: Echinoi- dea).–Journal of Systematic Palaeontology, 5/2, 163–192. doi: 10.1017/ S1477201906002021 KSIĄŻKIEWICZ, M. (1977): Trace fossils in the Flysch of the Polish Carpathians.–Pa- laeontologia Polonica, 36, 1–208. LÓPEZ-CABRERA, M.I., OLIVERO, E.B., CARMONA, N.B. & PONCE, J.J. (2008): Cenozoic trace fossils of the Cruziana, Zoophycos and Nereites ichnofacies from Fuegian Andes, Argentina.– Ameghiniana, 45/2, 377–392. MATTAUER, M. (1958): Etude géologique de l’ouarsenis oriental (Algérie).– Bulletin du Service Géologique de l’Algérie, 17, 1–534. MANLEY, R. & LEWIS, D.E. (1998): Ichnocoenoses of the Mount Messenger Forma- tion, a Miocene submarine fan system, Taranaki Basin, New Zealand.– New Zea- land Journal of Geology and Geophysics, 41/1, 15–33. doi: 10.1080/00288306. 1998.9514787 MAYORAL, E. & MUÑIZ, F. (2001): New Ichnospecies of Cardioichnus from the Mio- cene of the Guadalquivir Basin, Huelva, Spain.– Ichnos, 8/1, 69–76. doi: 10.1080/10420940109380174 MAYORAL, E., LEDESMA-VAZQUEZ, J., BAARLI, B.G., SANTOS, A., ROMAL- HO, R., CACHÃO, M., DA SILVA, C.M. & JOHNSON, M.E. (2013): Ichnology in oceanic islands: case studies from the Cape Verde Archipelago.– Palaeogeogra- phy, Palaeoclimatology, Palaeoecology, 381–382, 47–66. doi: 10.1016/j.pal- aeo.2013.04.014 MCKINNEY, M.L. (1986): Ecological causation of heterochrony: a test and implications for evolutionary theory.– Paleobiology, 12/3, 282–289. doi: 10.1017/ S0094837300013786 MONACO, P., GIANNETTI, A., CARACUEL, J.E. & YÉBENES, A. (2005): Lower Cretaceous (Albian) shell-armoured and associated echinoid trace fossils from the Sácaras Formation, Serra Gelada area, southeast Spain.– Lethaia, 38/4, 333–344. doi: 10.1080/00241160500355277 MONACO, P., RODRÍGUEZ-TOVAR, F. & UCHMAN, A. (2017): The ichnocoenosis of the bottom nepheloid layer (BNL) deposits: a case study from the Scaglia Tos- cana Formation (Paleogene, central Italy).– Bollettino della Società Paleontologi- ca Italiana, 56/2, 243–251. doi: :10.4435/BSPI.2017.13 NAIMI, M.N. & CHERIF, A. (2021a): Ichnological analysis of the late Miocene shallow marine diatomaceous deposits of the Lower Chelif basin (northwestern Algeria): Paleoenvironmental insights and comparison with deep diatomites.– Journal of African Earth Sciences, 104239. NAIMI, M.N. & CHERIF, A. (2021b): Inventory and assessment of significant scien- tific Algerian geoheritage: Case of remarkable geosites from Orania (Western Algeria). – International Journal of Geoheritage and parks, 9, 13–29. NAIMI, M.N., MANSOUR, B., CHERIF, A., CHEKKALI, M.C., BENKHEDDA, A. & BELAID, M. (2020): Lithostratigraphie et paléoenvironnements des dépôts mes- siniens de la terminaison nord-orientale des monts des Ouled Ali (bassin du Bas Chélif, Algérie nord-occidentale).– Revue de Paléobiologie, 39, 467–483. NAIMI, M.N., VINN, O. & CHERIF, A. (2021): Bioerosion in Ostrea lamellosa shells from the Messinian of the Tafna basin (NW Algeria).– Carnets de Géologie, 21/5, 127–135. PLAZIAT, J.C. & MAHMOUDI, M. (1988): Trace fossils attributed to burrowing echi- noids: A revision including new ichnogenus and ichnospecies.– Geobios, 21/2, 209–233. doi: 10.1016/S0016-6995(88)80019-6 POLVECHE, J. (1960): Contribution à l’étude géologique de l’Ouarsenis oranais.– Bul- letin du Service de la Carte Géologique de l’Algérie, 24, 1–577. POMEL, A. (1885): Les échinides du Kef Ighoud.– Matériaux pour la carte géologique de l’Algérie, Algiers, 31 p. POMEL, A. (1887): Echinodermes, fascicule 2.– Paléontologie ou description des ani- maux fossiles de l’Algérie, Algiers, 344 p. RAJCHEL, J. & UCHMAN, A. (2012): Ichnology of Upper Cretaceous deep-sea thick- bedded flysch sandstones: Lower Istebna Beds, Silesian Unit (Outer Carpathians, southern Poland).– Geologica Carpathica, 63/2, 107–120. doi: 10.2478/v10096- 012-0009-3 RIAHI, S., UCHMAN, A., STOW, D., SOUSSI, M. & BEN ISMAIL LATTRACHE, K. (2014): Deep-sea trace fossils of the Oligocene-Miocene Numidian Formation, northern Tunisia.– Palaeogeography, Palaeoclimatology, Palaeoecology, 414/15, 155–177. doi: 10.1016/j.palaeo.2014.08.010 SMITH, A.B. & CRIMES, T.P. (1983): Trace fossils formed by heart urchins – a study of Scolicia and related traces.– Lethaia, 16, 79–92. doi: 10.1016/j.pa laeo.2014.08.010 TCHOUMATCHENCO, P. & UCHMAN, A. (2001): The oldest deep-sea Ophiomorpha and Scolicia and associated trace fossils from the Upper Jurassic-Lower Cretaceous deep-water turbidite deposits of SW Bulgaria.– Palaeogeography, Palaeoclimato- logy, Palaeoecology, 169/1–2, 85–99. doi: 10.1016/S0031-0182(01)00218-8 UCHMAN, A. & KRENMAYR, H.G. (1995): Trace fossils from Lower Miocene (Ottnangian) molasse deposits of Upper Austria.– Paläontologische Zeitschrift, 69/3, 503–524. doi: 10.1007/BF02987810 UCHMAN, A. (2007a): Deep-sea trace fossils from the mixed carbonate-siliciclastic flysch of the Monte Antola Formation (Late Campanian–Maastrichtian), North Apennines, Italy.– Cretaceous Research, 28/6, 980–1004. doi: 10.1016/j.cre- tres.2007.01.005 UCHMAN, A. (2007b): Trace fossils of the Pagliaro Formation (Paleocene) in the North Apennines, Italy.– Beringeria, 37, 217–237. VILLEGAS-MARTÍN, J., NETTO, R.G., LAVINA, E.L.C. & ROJAS-CONSUEGRA, R. (2014): Ichnofabrics of the Capdevila Formation (early Eocene) in the Los Pa- lacios Basin (western Cuba): Paleoenvironmental and paleoecological implica- tions.– Journal of South American Earth Sciences, 56, 214–227. doi: 10.1016/j. jsames.2014.09.006