1. INTRODUCTION The Carboniferous System and especially the Pennsylvanian Subsystem provides fossil evidence of the first major radia- tion of the winged insects, the dominant subclass of the In- secta (GRIMALDI & ENGEL, 2005). The record comprises various body- and some trace fossils found in finer-grained deltaplain sediments (JARZEMBOWSKI, 1987). Insect fos- sils are, however, generally uncommon when compared with the abundance of plant compressions. Nevertheless, exami- nation of the latter can reveal evidence of insect activity. In this contribution, the first discovery from the Croatian Penn- sylvanian of the ichnospecies Phagophytichnus ekowskii VAN AMEROM, 1966, on the leaf of the cycadopsid Taeni- opteris carnotii ZEILLER, 1888, is described and discussed. Animal folivory on leaves is known from the Croatian Mio- cene (ÐEREK & JAPUNDŽIĆ, 2010), but this find is con- siderably older (c.~290 Ma). 2. GEOLOGY The tectonic belt of Mt. Velebit and Lika is the best known and most completely developed Upper Palaeozoic outcrop in Croatia, showing more or less continuous sedimentation from the Middle Pennsylvanian (Moscovian) to the end- Permian (Changhsinghian) (Jasenka Sremac pers. comm., 2011; JAPONDŽIĆ, KRIZMANIĆ, POLJAK & MJEDA, 2005). Upper Kasimovian-Gzhelian sediments are exten- sive, recording the rhythmic oscillations of sea-level fluc- tuation. Fossiliferous marine sediments predominate over sporadic beds of continental origin, the shoreline having never been far away. Plant compression fossils occur in grey mudstones in the Lika Region from which the trace fossil is described below. The Lika find is compared with Upper La- dinian material from Germany found in fluvial sediments and associated with a low sea-level stand (GRAUVOGEL- STAMM & KELBER, 1996). 3. SYSTEMATIC PALAEONTOLOGY Ichnogroup Phagophytichnidea VASILENKO, 2007b Ichnosubgroup Phagophytichnida VASILENKO, 2007b Ichnofamily Phagophytichnidae VYALOV, 1975 Ichnosubfamily Phagophytichninae VYALOV, 1975 Ichnogenus Phagophytichnus van AMEROM, 1966 Remarks. Phagophytichnus is a long-ranging ichnoge- nus of marginal leaf-biting species and usually considered to be the result of insect chewing (as opposed to gastropod rasping (by land snails/slugs), RETALLACK, 2001: p. 137). Ab STRA CT The feeding trace Phagophytichnus ekowskii VAN AMEROM, 1966 is recorded on Taeniopteris carnotii ZEILLER, 1888, a cycadopsid leaf, from the Lika mudstone (Upper Kasimovian-Gzhelian) of Croatia. The distribution and oc- currence of this ichnogenus and species are discussed and comparison made with occurrences on Taeniopteris an- gustifolia (SCHENK, 1927) in the Upper Landinian of Germany. Recent analogues are considered of the possible producer and an orthopteroid insect is suggested. Keywords: Pennsylvanian, Ladinian, Phagophytichnus ekowskii, cycadopsid, Taeniopteris carnotii, chewing Geologia CroaticaGeologia Croatica The oldest plant-insect interaction in Croatia: Carboniferous evidence  Ed A. Jarzembowski Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, Nanjing 210008, China; (jarzembowski2@live.co.uk) doi: 104154/gc.2012.28 Geologia Croatica 65/3 387–392 1 Pl. Zagreb 2012 Geologia Croatica 65/3Geologia Croatica 388 Originally referred to the Cibichnia or feeding (eating) traces, Phagophytichnus was then included in Praedichnia, trace fossils of predation, but more recently moved to Phagophy - tichnidea, feeding traces on plants (VASILENKO, 2007b), and even been given its own family, Phagophtyichnidae (VYALOV, 1975). ZHERIKHIN (2003) previously placed Pha gophytichnus in Trogichnia reserved for chew marks on plants. Evidence that the living plant was predated is provided by a pronounced thickening (ridge) sometimes seen along the damaged margin and interpreted as callus (‘scab’) formation (see figure of damaged leaf of the pteridosperm Macroneu- ropteris scheuchzeri (Hoffmann) Cleal, Shute & Zodrow from the late Asturian of England in ATTENBO ROUGH, 2005: p.167). This was previously interpreted as leaf mining (MÜL- LER, 1982). Distribution. Phagophytichnus has been described from the Late Mississippian to the Pliocene and has a worldwide distribution (IANNUZZI & LABANDEIRA, 2008; BRUS- TUR, 1997). It also represents the most common type of ex- ternal foliage feeding in the fossil record (LABANDEIRA, 2006). The inclusion of small holes in ginkgoalean foliage from the Lower Cretaceous, however, overstretches the generic limits of Phagophytichnus (WATSON, LYDON & HARRI- SON, 2001). These traces are Trogichnia or Nygmichnia (chewing or piercing marks respectively) following ZHE- RIKHIN (2003, tab. III), the latter not included in VYALOV (1975). Like Phagophytichnus, these marks are both in- cluded in Zherikhin’s ‘redundant’ Phagophytichnia: Phyllo- phagichnia (leaf and petiole damage). For ease of reference, I propose to call them porichnid (from the ancient Greek po- ros, hole, and Latin ichnus, trace) within Vasilenko’s Phago- phytichnida pending a more detailed study. Phagophytichnus ekowskii VAN AMEROM, 1966 (Pl. 1, Figs. 1–2) = Cuniculonomus (Arcophionomus) undulatus MÜL LER, 1982 = Damage Type 12 (LABANDEIRA, WILF, JOHNSON & MARSH, 2007) Diagnosis: An approximately semicircular, isolated exci- sion (cut out) of the leaf margin which is shallow or deep but less than 180 degrees of arc (LABANDEIRA et al., 2007). Description and Measurements: The Lika trace fossil is cuspate excisions (overall length c. 2 cm, width 3 mm) from the margin of the leaf of Taeniopteris carnotii ZEIL- LER, 1888 (NĔMEJC, 1936) preserved as a compression in mudstone (Pl. 1, Figs. 1A–C). It comprises 2 to 3 semicir- cular, asymmetrically curved, successive and separate exci- sions, each of which is up to 9 mm long, and all approaching the midpoint of the distance between the leaf margin and midvein. Material (figured), locality and date: Specimen 1 (Pl. 1, Figs. 1A–C). Registration number: 10776. Repository: Ge- ology-Palaeontology Department, Croatian Museum of Nat- ural History. Locality Lika Region, east of Mt. Velebit, Cro- atia. Age: Stephanian C (late Kasimovian to Gzhelian). Specimen 2 (Pl. 1, Fig. 2). Registration number: SCHL- 061a. Repository: Kelber Collection. Locality: Scheerich, Franconia, Germany. Age: Lower Keuper, Upper Landinian. Remarks. The Lika ichnofossil (Upper Pennsylvanian, c. 305 Ma) is of a similar size to marginal excisions on Taen- iopteris angustifolia (SCHENK, 1927) from the late Middle Triassic (c. 230 Ma) of Scheerich, Germany (cf. GEYER & KELBER, 1987: fig. 7, lower left and KELBER & GEYER, 1989: pl. 2, figs 3, 5; also KELBER & GAYER, 1989: figs 4, 6, another specimen). One of these Triassic leaves, how- ever, figured herein is extensively eaten (Pl. 1, Fig. 2) with successive marks (KELBER & GEYER, 1987), although a new find shows two widely spaced excisions, one compara- tively small (STEINKERN, 2011). Both Carboniferous and Triassic specimens are similar in that their location is away from the midvein of the leaf and occurrence along one mar- gin. The Triassic material has been only tentatively referred to Phagophytichnus (GRAUVOGEL-STAMM & KELBER, 1996), but like the Lika specimen, can be referred to P. ekowskii, as currently diagnosed. In the absence of cuticular studies, Taeniopteris carnot- tii may be a cycad or a bennettite within the Class Cycadop- sida Barnard & Long (cf. CLEAL & REES, 2003), and pos- sibly a synonym of Taeniopteris multinervis Weiss, 1869 (Chris Cleal, pers. comm., 2012). Distribution (range and occurrence): Phagophytich- nus was originally based on P. ekowskii found on the pteri- dosperm Mixoneura Weiss (Mixoneura wagneri Lorenzo nec Neuropteris praedentata GOTHAN; CASTRO, 1997), from the Spanish Stephanian B (VAN AMEROM, 1966). The stratigraphic range of P. ekowskii was subsequently extended from the Late Mississippian to the late Triassic and the host range extended to various other pteridosperm genera including: Autunia Krasser, Dicroidium Gothan, Glossopteris Brongniart, Macroneuropteris Cleal, Shute & Zodrow, Neu- ropteris Brongniart Paripteris Gothan, Odontopteris (Brong- niart) Sternberg, Pursongia ZALESSKY, Triphyllopteris SCHIMPER, and the ginkgopsid Dejerseya Herbst (IAN- NUZZI & LABANDEIRA, 2008; LABANDEIRA, 2006; LABANDEIRA & ALLEN, 2007; TROUT, LABANDEIRA & CHAPMAN, 2000; VASILENKO, 2007a). VASILENKO (2007b) recorded it (as P. ekovskii) on ferns and pteridosperms as late as the Cretaceous (Cenomanian), but he referred bite (chew) marks on other gymnosperms (conifers and gingkos) to the ichnogenus Pinovulnus VASILENKO, although the smaller Pinovulnus serpentiformis Vasilenko resembles P. ekowskii and there may be some overlap (cf. VASILENKO, 2006, fig. 5 and Pl. 1, Fig. 1). He was evidently following ZHERIKHIN’s (2003) classification based on plants rather than functional-feeding groups. As for geographic range, Phagophytichnus ekowskii is now worldwide (Laurasia + Gondwana), being known from Euramerica to Australasia (LOBUE & HASIOTIS, 2010; SRIVASTAVA & AGNI- HOTRI, 2011; PREVEC et al., 2009). In addition, it is for- mally recorded on cycadopsid leaves (Taeniopteris species) from the Upper Pennsylvanian of Croatia and Middle Triassic of Germany herein. Ed A. Jarzembowski: The oldest plant-insect interaction in Croatia: Carboniferous evidence Geologia Croatica 389 Attribution: LABANDEIRA et al. (2007) attributed this isp. to protorthopteran/stem-orthopteroid insects, whilst al- lowing that other mandibulate (jaw-bearing) arthropods might be involved (diplopod millipedes), whereas GU, BÉTHOUX & REN (2011) stressed a preference for carnivory in these in- sect groups. GEYER & KELBER (1987) and KELBER & GEYER (1989) considered that holometabolous insects, es- pecially the caterpillars of Lepidoptera (moths and butterflies) could also be responsible for this isp. as well as orthopterans (grasshoppers and crickets). Body fossils of Lepidoptera are known by the Lower Jurassic and Hymenoptera (sawflies, ants, bees and wasps), which also have a caterpillar larva, are known by the Upper Triassic although Orthoptera are known from the Pennsylvanian onwards (JARZEMBOWSKI, 2003). Millipedes are not considered to be a significant defoliator to- day but orthopterans are a large and diverse order with her- bivorous and carnivorous forms (KEY, 1970). A bush cricket- like insect may therefore have been responsible for the feeding damage on European Taeniopteris, although it is unlikely to be always the same species due to a significant time gap (~c. 75 Ma) between the Lika and Scheerich leaves, and an un- known stem-holometabolan might be involved. BECK & LA- BANDEIRA (1998) also recognised orthopteroid damage on Taeniopteris sp. from the Early-Middle Permian of the USA differing from P. ekowskii in including leaf holes as well as marginal feeding, but supported by the presence of veinal strands as produced by recent short-horned grasshoppers. The latter are also present in chew marks on leaflets of Anomo- zamites villosus POTT, McLOUGHLIN, WU & FRIIS, 2012, a newly described cycadopsid from the Middle Jurassic of PLATE 1 Phagophytichnus ekowskii on Taeniopteris carnotii, Lika. 1a – part, middle right. The undulation in the leaf margin (top right) is sedimentary. Scale: 20 lipa coin,18 mm diameter; 1b – close up of part in a; 1c – close up of counterpart. 2 – P. ekowskii on Taeniopteris angustifolia, SCHE ER ICH. Note callus formation. 3 – Psophus stridulus on Taraxacum officinale, recent. Scale line, 1 cm. Redrawn after KAZAKOVA (1985). 1a 1b 2 1c 3 Geologia Croatica 65/3Geologia Croatica 390 China (pers. obs.). Examination of the Lika leaf under mag- nification was however inconclusive, revealing irregularities along the margin, but also embedding in an obscuring, grainy matrix (but see gross cf. below). 4. RECENT ANALOGUE Grasshoppers and caterpillars are unrelated insects but share mandibulate (basic chewing) mouthparts. Modern lepi- dopteran (butterfly) caterpillars are known to defoliate some cycads, such as the Cycad Blue (Theclinesthes onycha HEWITSON) in Australia. Cycads are no longer native to Europe, but I have often observed caterpillars feeding on an- giosperms in southeast England. The caterpillar’s numerous paired legs are used to hold on to the leaf, often tenaciously. As in other mandibulate insects, the paired jaws are orien- tated beneath the anterior part of the head capsule and the edge of the leaf is harvested in a sideways cutting motion. The head moves progressively forwards and downwards, sometimes obliquely, to obtain purchase and then access to fresh tissue. This would produce a curved excision as seen on the fossil leaves. Slight body movements coupled with a break in feeding, often due to a disturbance, result in an ex- cision with subordinate cuspules evident (cf. KELBER & GEYER, 1989: fig. 4; also, GRAUVOGEL-STAMM & KELBER, 1996: fig. 6). The insect may resume feeding, or move on allowing wound reaction (callus) tissue and subse- quent drying out to form a thickening inside the damaged margin as seen on fossils (e. g. Pl. 1, Fig. 2). My observa- tions on bullace (Prunus insititia L.) point to this thickening of the callus being a gradual process, taking weeks to form a feature prominent enough to be readily seen on compres- sion leaves. This could explain the apparent absence of cal- lus in P. ekowskii in JARZEMBOWSKI (2004: pl. 1, fig. 3). Bush crickets are less easy to watch, but such a great green ‘grasshopper’ fed on lettuce showed broadly similar chewing behaviour (MICHAËL, 2011). GANGWERE (1966) discussed feeding in more derived orthopteroids (acri- diid grasshoppers): interestingly, some of the chew marks produced by the rattle grasshopper (Psophus stridulus (L.)), a recent forbivorous European species, show a comparable outline to the Lika fossil (Pl. 1, Fig. 3). Modern land snails, such as Helix aspersa (Müller), can also form marginal inci- sions and holes in leaves by rasping downwards and side- ways with their radulae. Terrestrial molluscs are scarce in the Carboniferous-Triassic of Europe, but have been found in Middle Pennsylvanian nodules with plants and insects in the Upper Silesian Coalfield and in the Middle/Upper Penn- sylvanian of the English Midlands, investigated during IGCP 469 (STWORZEWICZ, SZULC & POKRY SZKO, 2009). Where known, the affinities of Carboniferous European snails (ellobioid and Cerion-like, loc. cit.) suggest that they fed on detritus or non-vascular plants by analogy with recent thorn and peanut snails rather than the leaves discussed herein. The situation is similar in the Late Pennsylvanian of North America, such as at Joggins, in Newfoundland, also investigated during IGCP 469 (FALCON-LANG, BENTON, BRADDY & DAVIES, 2006; molluscan detritivory rather than folivory (leaf feeding) was also suggested by SOLEM & YOCHELSON, 1979). In the absence of slime trails and coprolites, stereo-electron microscopy on well-preserved fossil plants may help to distinguish the work of biting man- dibles from rasping radulae in future studies. 5. CONCLUSION Phagophytichnus ekowskii provides fossil evidence for a long- and wide-ranging, generalised feeder with low host specific- ity, principally on pteridosperm leaf margins. GRAUVOGEL- STAMM & KELBER (1996) considered Phagophytichnus on Taeniopteris, however, as part of a novel insect-plant interac- tion on Mesozoic Cycadopsida following the Permo-Triassic extinction event. The Lika find shows that the Phagophytich- nus-Taeniopteris association is considerably older, dating back to the late Palaeozoic (Upper Pennsylvanian). Insects are otherwise unknown from the Croatian Carboniferous and this discovery also helps to fill a gap in the Pennsylvanian record of southeast Europe. Its rarity is, however, not surprising be- cause Taeniopteris was likely more resistant than pterido- sperms to insect attack due to the cycadopsid’s tougher cuticle and well-developed resin glands (BECK & LABANDEIRA, 1998). In contrast, the hirsute leaves of Macroneuropteris may have only slowed down chewing (ATTENBOROUGH, 2005, figure p. 167). IGCP 575 is concerned with environmental change at the Middle/Upper Pennsylvanian transition. The Lika Flora needs to be examined further to determine the extent of in- sect herbivory, the current data predicting higher rates of pteridosperm folivory by insects unless carnivory was wide- spread. A search for associated fauna may provide clues to a more precise identification of the consumer. ACKNOWLEDGEMENT I wish to thank: the Geological Society and Royal Society of London for financial support to visit Croatia; the staff at the Croatian Museum of Natural History and Geology De- partment, University of Zagreb, for their kindly hospitality; Conrad LABANDEIRA (Smithsonian Institution) for help- ful comments; Klaus-Peter KELBER (Würzburg) for copies of his papers and use of Pl. 1, Fig. 2; James JEPSON (Man- chester University), Chris CLEAL (National Museum of Wales), Wendy CAWTHORNE (Geological Society of Lon- don), Xavier DELCLÒS (Barcelona University), Jason DUNLOP (Museum für Naturkunde), André NEL (Muséum national d’Histoire naturelle), Jörg SCHNEIDER (Ber- gakademie Freiberg) and library staff at the Natural History Museum (London) for help with papers; Fred CLOUTER (Isle of Sheppey) and Prof. Tihomir MARJANAC (Faculty of Science, Geology Department) for improving my Figures; my wife, Brigid, for garden ecology; Conrad LABAN- DEIRA and Olivier BÉTHOUX (Commonwealth Scientific and Industrial Research Organisation) for their reviews. 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