Journal of Earth Science, Vol. 26, No. 2, p. 192–202, April 2015 Printed in China DOI: 10.1007/s12583-015-0525-z ISSN 1674-487X Exceptionally Preserved Caddisfly Larval Cases (Insecta) from the Lower Cretaceous of the Liupanshan Basin, Western China Xin He1, 2, Zhong-Qiang Chen*1, Zongsheng Lu1, Jun Li3, Wei Hu3, Shengfu Li2, Zhitao Xu2 1. State Key Laboratory of Biogeology and Environmental Geology, China University of Geosciences, Wuhan 430074, China 2. Guangxi Geological Exploration Institute of China Metellurgical Geology Bureau, Nanning 530000, China 3. School of Earth Sciences and Graduate School, China University of Geosciences, Wuhan 430074, China ABSTRACT: Abundant well-preserved tubular fossils of caddisfly (Insecta: Trichoptera) larval cases are reported from the Early Cretaceous Madongshan and Naijiahe formations of the Liupanshan Basin, Ningxia Province, western China. Most cases were mainly preserved in life position and densely packed in various layers. Individual cases in each layer tended to be same in size and were erect and parallel to one another and open at both ends. In a transverse section cut perpendicular to the long axis of the cases, individual case appears to form a rounded ring. Small cases are elliptic in a cross-section oblique to the long axis of the cases. Tube walls are nearly subparallel to one another in longitudinal section with both ends being open. The caudal end of the case slightly tapers and usually points downward. The cases were closely packed, almost touching with one another and lacking bifurcate or connecting structure. The overwhelming majority of cases were partially or fully filled with calcite. The case wall embraces a medium particle layer flanked by inner and outer organic layers. Individual particles are ovate in outline and comprise cryptocrystalline or ganic pellets. SEM imaging shows that those pellets are sub-cylindrical in outline and elliptic in cross section, and are made primarily of calcium carbonate. All features observed justify the assignment of the Liupanshan caddisfly cases to ichnogenus Coprindusia. The extinct insect Ningxiapsyche fangi was found in association with the Liupanshan caddisfly larval cases, and thus could be the candidate of the potential trace-maker. KEY WORDS: caddisfly larval case, tube wall microstructure, Early Cretaceous, Liupanshan Basin, western China. 0 INTRODUCTION Both the Trichoptera and Lepidoptera of the Ampliesmenoptera are most distinct among the holometabolous insects (Huang et al., 2009; Kjer et al., 2001). They are also the second largest lineage of aquatic insects (Gallego et al., 2011). Compared with extremely diverse extant Trichoptera, which contains 13 574 species of 608 genera belonging to 47 families, fossil Trichoptera is much less diverse and includes 680 species, 127 genera and 12 families. The larval and pupal stages of extant Trichoptera caddisflies are mainly aquatic and widespread in all freshwater ecosystems ranging from mountain streams, ponds, large rivers, lakes, wetlands, and even brackish water, while the adults inhabit mostly terrestrial environments (Daniel et al., 2009; Moor and Ivanov, 2008; Greenwood et al., 2003). Most caddisfly larvae obtain materials from the surrounding *Corresponding author: [email protected] © China University of Geosciences and Springer-Verlag Berlin Heidelberg 2015 Manuscript received October 21, 2014. Manuscript accepted January 20, 2015. habitats to construct their tubular cases. Their portable cases were attached as dwelling and protection in foraging food (Brian et al., 2011; Boyero and Barnard, 2004; Williams and Penak, 1980; Mackay and Wiggins, 1979). Since Cockerell (1923) reported the first example, the fossil caddisfly cases have been reported from the Mesozoic and Cenozoic strata worldwide. The known oldest fossil larval case was found in Middle Jurassic (Huang et al., 2009; Ponomarenko et al., 2009; Ivanov and Sukatsheva, 2002), although fossil caddisflies are known from the Permian. To date, total 13 ichnogenera have been proposed to accommodate fossil larval cases based on their various morphologies, including Conchindusia, Piscindusia, Ostracindusia, Pelindusia, Terrindusia, Molindusia, Folindusia, Secrindusia, Scyphindusia, Charindusia, Indusia, Coprindusia, and Tektonargus (Gallego et al., 2011; Ponomarenko et al., 2009; Ivanov, 2006; Sukatsheva, 2005; Hasiotis et al., 1998; Jarzembowski, 1995; Lewis, 1970a, b). The possible case-makers still remain disputed, although several candidates have been proposed (Paik, 2005; Leggitt and Cushman, 2001; Bradley, 1974). This is because no smoke-gun organisms were captured in the cases. Moreover, the preliminary classification schemes have also proposed to describe the wall components of those tubular cases (Boyero and Barnard, He, X., Chen, Z. Q., Lu, Z. S., et al., 2015. Exceptionally Preserved Caddisfly Larval Cases (Insecta) from the Lower Cretaceous of the Liupanshan Basin, Western China. Journal of Earth Science, 26(2): 192–202. doi:10.1007/s12583-015-0525-z Exceptionally Preserved Caddisfly Larval Cases (Insecta) from the Lower Cretaceous of the Liupanshan Basin, Western China 2004; Loewen et al., 1999; Bradley, 1924). However, microstructures of caddisfly case walls still remain unclear. More recently, we have obtained abundant, exceptionally preserved caddisfly larval cases from the Liupanshan Basin, Ningxia Province, western China (Fig. 1). Such well-preserved larval cases not only provide additional examples of fossil caddisfly cases, but also enable us to insight into microstructure of case walls, in particular, the arrangement pattern of cemented pellets in medium layer of case wall. The latter characteristics are crucial in ichnology of those fossil caddisfly cases. 1 GEOLOGICAL AND STRATIGRAPHIC SETTINGS The studied larval cases were collected from the Lower Cretaceous succession in the Yaoshan Section of the Liupanshan Basin, approximately 20 km northeast of the Tongxin Town, eastern Ningxia Province, western China (Figs. 1 and 193 2a). The Liupanshan Basin (105°E–107°E, 34 40'N–37 10'N) is a reverse triangular inland basin and located at the junction among the Shaanxi, Gansu and Ningxia provinces, western China (Fig. 1). Tectonically, the basin lies at the northeastern margin of the Tibet-Qinghai Plateau and is bounded by the Eerduosi Block in the east, the Alashan Block in the north, and the Qilian-Qinling orogenic fold belt in the southwest (Tang et al., 2004; Fu and Li, 2002), respectively. The lacustrine basin was uplifted during the Yanshan orogeny of Late Jurassic age. Since then, the Liupanshan Basin was filled with terrestrial sediments of alluvial fan, deltaic, braided river, and lacustrine facies of the Liupanshan Group, which comprises the Sanqiao, Heshangpu, Liwaxia, Madongshan, and Naijiahe formations in an ascending order (Zhou et al., 2005). Both palynology and paleomagnetic correlation suggest an Early Cretaceous age for this group (Dai et al., 2010, 2009; Li and Du, 2006). Figure 1. Geographical map showing the location of the studied section in the Liupanshan Basin, Ningxia Province, western China (left figure) and columnar section of the Lower Cretaceous succession showing fossil horizons. Of these, the early Early Cretaceous Sanqiao and Heshangpu formations are dominated by reddish coarse sandstone and conglomerate of alluvial fan and deltaic facies and indicate the dry and hot sub-tropical temperate climatic conditions (Li and Du, 2006; Zhou e al., 2005). The Liwaxia Formation and lower Madongshan Formation of middle Early Cretaceous age are composed mainly of grey find sandstone and mudstone of deltaic and lacustrine facies, indicating a relatively warm, humid climatic condition (Zhou et al., 2005). The upper Madongshan Formation and Naijiahe Formation of late Early Cretaceous age consist chiefly of fine sandstone, mudstone and dolomites and characterized by abundant pseudomorphs of gypsum and halite, indicating high-evaporatic, dry climatic condition (Jin and Cao, 2006). Climatic varations through the Liupanshan Group are also reinforced by palynologic analysis. The Liupanshan Group contains abundant spores Schizaeoisporites and pollens Ephedripites, Jugella, and Classopollis (Li and Du, 2006; Liu, 1983), characteristics of hot, arid microfloral assemblages worldwide (Moor and Ivanov, 2008). The environmental and climatic inference of the Liupanshan Group is also strengthened by organic carbon isotopic geochemistry (Bai et al., 2003) and changes of chromaticity and magnetic suscep- 194 Xin He, Zhong-Qiang Chen, Zongsheng Lu, Jun Li, Wei Hu, Shengfu Li and Zhitao Xu tibility (Dai et al., 2010, 2009), all of which indicate that the Liupanshan Group successions were deposited in the dry and hot sub-tropical temperate climatic conditions, as the similar climatic conditions of most landmass in the world during that times (Moor and Ivanov, 2008). The caddisfly cases are preserved in several horizons between the upper Madongshan and basal Naijiahe formations (Fig. 2a). The former formation is a thick lacustrine siliciclasticcarbonate unit, resting conformably on the Liwaxia Formation of Hauterivian-Barremian age and overlain by the Naijiahe Formation which is characterized by evaporitic rocks of Albian age (Dai et al., 2009; Shi et al., 2006; Liu, 1983). The Madongshan Formation is dominated by mudstone in the lower part and composed of mudstone, siltstone and marl, with minor constituents of shale and calcareous mudstone in the upper part. The upper Madongshan Formation yields caddisfly larval cases at its basal part (beds 1–4; Fig. 1) and abundant fossils of ostracods, fishes, insects, plants, sporopollen, and conchostracans in the rest parts of the formation (Li and Du, 2006; Zhang, 2004; Li, 1995; Qi, 1988; Liu, 1983). These faunal and floral assemblages point collectively to an Early Cretaceous age (Li and Du, 2006). Both lithofacies and paleoecologic analyses indicate that the upper Madongshan Formation represents the sedimentation of lacustrine setting. The basal Naijiahe Formation also yields caddisfly cases and comprises mudstone and calcareous mudstone, from which Hong and Li (2004) reported insect wing of Ningxiapsyche fangi of Aptian–Albian age (Early Cretaceous). All lines of evidence indicate that the Liupanshan caddisfly cases are Aptian–Albian (Early Cretaceous) in age. Figure 2. Caddisfly case bioherms and taphonomy and morphologies of fossil caddisfly larval cases from the Lower Cretaceous of the Liupanshan Basin. (a) Caddisfly case bioherms on the outcrop (arrow indicated); (b) columnar colonies of caddisfly larval cases on the outcrop; (c) the planar structure of the caddisfly-dominated columns; (d) the autochthonous burial caddisfly larval cases, scale bar: 5 mm; (e) close-up of the boxed area in D showing one complete larval case; (f) the allochthonous burial caddisfly larval cases, bar: 5 mm; (g) close-up of F showing tiny larval cases, scale bar: 1 mm. Exceptionally Preserved Caddisfly Larval Cases (Insecta) from the Lower Cretaceous of the Liupanshan Basin, Western China 2 3 MATERIAL AND METHOD Number of specimens 75 (a) 200 (b) N=281 160 60 120 45 80 30 40 15 0 RESULTS 3.1 Taphonomy and Morphology of Caddisfly Larval Cases 3.1.1 Taphonomy On the outcrop, the caddisfly larval cases were mainly preserved in autochthonous and allochthonous burial states. The autochthonously preserved cases are characterized by the densely packed larval cases in various layers. Individual cases in each layer were erect and parallel to one another and open at both ends, exhibiting their life position (Fig. 2d). They appear to have a narrow size range with case length varying from 11 to 15 mm. Of these, 70% individuals are 13–14 mm long (Fig. 3a). Of 281 cases counted here, 75% individuals are 2.4–2.5 mm in diameter (Fig. 3b). Thus, the in situ larval cases exhibit strong size selectivity. In contrast, the allochthonously buried Both individual and colonial caddisfly case samples have been collected. Their taphonomic features were also observed in the field. Both petrographic and scanning electron microscope (SEM) imaging analyses have been undertaken to probe into microstructures of caddisfly case walls. Prior to observation under a Quanta 200 FEI SEM, cemented pellets of tube walls were coated with 5–7 nm gold using a SCD 005 BAL-TEC auto sputter coater. Thin-section micrographs were taken using a Nikon Eclipse E200 digital camera mounted on an E200 POL Standard Set transmitted-light polarizing microscope. N=135 0 1 3 5 7 9 11 13 1.0 15 17 1.5 2.0 2.5 Number of specimens (c) 25 N=145 15 10 10 5 5 1 3 5 7 9 11 13 0 15 17 1.0 (e) Number of specimens N=113 20 15 100 3.0 (d) 25 20 0 195 N=280 250 1.5 2.0 2.5 3.0 (f) N=394 200 80 60 150 40 100 20 50 0 1 3 5 7 9 11 Length (mm) 13 15 17 0 1.0 1.5 2.0 2.5 Diameter (mm) 3.0 Figure 3. Measurements of larval cases from the Liupanshan Group. (a) Lengths of the autochthonously preserved larval cases; (b) diameters of the autochthonously preserved larval cases; (c) lengths of the allochthonously preserved larval cases; (d) diameters of the allochthonously preserved larval cases; (e) lengths of all caddisfly larval cases; (f) diameters of all caddisfly larval cases. Xin He, Zhong-Qiang Chen, Zongsheng Lu, Jun Li, Wei Hu, Shengfu Li and Zhitao Xu 196 larval cases have a much broader size range with case length varying from 2 to 17 mm. Of these, the individuals having 10–11 mm in length are most frequently present, but only occupy 17.2% of all cases. Moreover, 13% individuals are 5–6 and 8–9 mm in length, respectively and 10% individuals are 9–10 mm and 13–14 mm in length, respectively (Fig. 3c). Of 113 cases counted here, 20% individuals are 2.5–2.6 mm in diameter, but individuals having 2.0–2.1, 2.1–2.2, 2.3–2.4, 2.4–2.5, and 2.6–2.7 mm, respectively are also commonly present, each group occupying around 10% (Fig. 3d). Thus, the allochthonously buried larval cases do not show any size preference. Furthermore, they were usually not orientated and randomly scattered on sediments, indicating transportation (Figs. 2f, 2g). The single cases are usually coated by thin calcite laminae comprising microsparite. Single lamina is about 0.1 mm thick. Blue-green algal mats, i.e., Girvanella (Fig. 4c) encrusted on surfaces of single larval cases to aggregate pellets, shell fragments and small microbial encrusts, like the modern counterparts (Fig. 4a) and aggregated single or colonial cases to form caddisfly bioherms. Single caddisfly case, the overwhelming majority of larval cases were partially or fully filled with sparite cement, microbial encrusts and pellets (Fig. 4b). 3.1.2 Morphology Caddisfly cases usually colonized to build bioherm, which is 3–5 m high. Caddisfly bioherms extend laterally up to 400–500 m, and thus are spectacular in the field (Fig. 2b). Each bioherm comprises case columns, which were constructed by 4–6 layers of caddisfly cases. Individual columns, up to 2– 2.5 m in height, are cylindrical and slightly taper towards the top in vertical outline (Fig. 2b). They are about 1 m in diameter and dome-like in plan view (Fig. 2c). In the columnar colonies the layered caddisfly cases are separated from one another by thin lime laminae of possible microbial origin. Erect caddisfly cases form case bands, which are parallel to bedding plane (Fig. 2d). Single caddisfly case-concentrating bands are 10–13 mm thick, while single laminar layers are about 2 mm thick. The incumbent caddisfly cases are scarce (Fig. 2e). In addition, some caddisfly larval cases were dispersedly preserved in calcareous siltstones (Figs. 2f, 2g). Of the caddisfly cases observed herein, about 33% individual cases are 13–14 mm in length and 60% cases are 2.4–2.5 mm in diameter (Figs. 3e–3f). In a transverse section cut perpendicular to the long axis of the case, individual case appears to be a rounded ring (Figs. 5d, 5f). In a cross-section oblique to the long axis of the case, individual case is elliptic in outline (Figs. 5e, 5g). In the cross section cut parallel to the long axis of the case, individual case is characterized by two almost subparallel walls with both ends being open (Fig. 5h). The caudal end of the tube often slightly tapers and points downward (Figs. 5a–5b, 5h). The cases are closely packed, open at two ends, nearly touching with one another and lacking bifurcate or connecting structures. 3.1.3 Figure 4. (a) Modern caddisfly case with insect inside (after Holzenthal et al., 2007), scale bar is 3 mm; (b) microphotograph in longitudinal section of larval case showing that case inside is filled with sparite cement (sc), microbes (m), and microbial encrusts (me), and that surfaces were encrusted by microbial mats (mm) (i.e., Girvanella), microbial encrusts, pellets (p), and fossil fragments (sf), Scale bar is 1 mm. Case wall microstructure and components The case wall embraces a distinct three-layer architecture with a medium particle layer being flanked by the inner and outer layers, both of which comprise dark organic matters (Figs. 4d–4h). The inner dark organic layer is discontiguous, 0.02 mm in thickness and is composed mainly of micrites. The outer organic layer shares the same compositions with the inner layer, but is thicker, about 0.04 mm in thickness and continuous. Individual particles are principally ovate in outline and comprise cryptocrystalline organic pellets, which are similar to the Late Jurassic−Early Cretaceous fecal pellets reported from the Bavaria (Flügel, 2004) and modern fecal pellets from the Coorong Lake in Australia (Scholle and Ulmer-Scholle, 2003). These oval particles are arranged regularly along the case walls with similar sizes and shapes. They, however, usually appear out-of-order on the bottom of the tubes. The particles appear baculiform shapes with rounded ends in cross section and are nearly circular in longitudinal section (Fig. 6). SEM imaging analysis shows that these pellets are sub-cylindrical in outline and elliptic in cross section, and comprise primarily calcium carbonates (Fig. 7). 4 DISCUSSION 4.1 Comparison with Other Fossil Caddisfly Larval Cases Elsewhere in the World A few collections of the Meosozic–Cenozoic caddisfly (Trichoptera) cases have been reported worldwide. The Liupanshan cases are significantly smaller than the Early Cretaceous counterparts reported from southern England (Jarzembowski, 1995), Siberia of Far East, Russia (Ivanov, 2006) and Liaoning of northeastern China (Huang et al., 2009). The morphologically similar caddisfly cases have also been described from the Eocene Green River Formation of the western US (Leggitt and Cushman, 2001), but they are significantly smaller than the Liupanshan cases. Paik (2005) also reported a Exceptionally Preserved Caddisfly Larval Cases (Insecta) from the Lower Cretaceous of the Liupanshan Basin, Western China 197 Figure 5. Morphology and micro-structures of the Early Cretaceous caddisfly larval cases. All came from Bed 4 of the Yadongshan Formation in the Yaoshan Section in Tongxin, Ningxia (scale bar is 1 mm). (a) Individual coniform larval case; (b) larval case colonies showing no connecting structures between the single cases; (c) larval case colonies showing both open ends; (d)–(g) cross sections showing that case tube wall comprises three layers: inner dark organic layer (IL), medium pellet layer (ML) and outer dark organic layer (OL); (h) longitudinal section showing that the case colony comprises two subparallel rows. Note the pellets in tube walls have a similar size and shape to one another. Figure 6. Diagrams showing arrangements of cemented particles in case wall. (a) Pellets arranged in planar cross section of caddisfly cases; (b) pellets arranged in longitudinal section; (c) 3-demension view of caddisfly cases showing pellet arrangement, scale bar is 1 mm. microbial caddisfly bioherms from the Early Cretaceous Jinju Formation of the Jahyeri area, Korea. The Jahyeri caddisfly bioherms are similar to the Liupanshan caddisfly bioherms in terms of dome outline and size. Single cases in both bioherms were stuck together by microbes and form layers to construct high-relief bioherms. However, the Liupanshan caddisfly bioherms are better preserved with more distinguished layers than Korean bioherms. The pronounced tiny particles of cryptocrystalline pellets filled in the medium layer of wall tube of single cases were not observed in Korean caddisfly cases. Moreover, the tubular portable cases and cemented pellets were controlled by the habitats of the caddisfly larvae. There are principally three types of the pellets cemented in larval case walls (Table 1). The first type, with caddis larvae cases dispersed in streams and lakes, is constructed of sand grains. The second type has the case walls constructed by leaf pieces of both angiosperm and gymnosperm fragments, fragmented ostracod valves, and fish bones/scales. Their case-building caddisfly larvae usually inhabit the lentic ecosystems such as pond and lake. The third type, with the larval cases being constructed by organic pellets, is usually preserved in some restricted environments such as lagoons (especially supersaline lagoons). As stated above, the Liupanshan Group succession was referred to be deposited in dry and hot sub-tropical temperate climatic conditions by integrating sedimentary facies, palynofloral assemblages (Li and Du, 2006; Liu, 1983), organic carbon isotopic signals (Bai et al., 2003), and changes of chromaticity and magnetic susceptibility (Dai et al., 2010, 2009). Thus, the Liupanshan caddisfly cases possess tube walls constructed by organic pellets and are mainly preserved in the restricted, salty lake. They therefore share the similar living behaviors to those larval cases reported from the Early Cretaceous Baissa Formation of Siberia, Russia (Ivanov, 2006). Xin He, Zhong-Qiang Chen, Zongsheng Lu, Jun Li, Wei Hu, Shengfu Li and Zhitao Xu 198 Table 1 Cemented particle types of the caddisfly case tube walls Tube wall components Stratigraphic unit Age Locality References Ostracod valves Leaf pieces Green River Fm. Lataha Fm. Eocene Miocene Wyoming, US Washington, US Bradley, 1924 Lewis, 1970a Leaf pieces Windrow Fm. Cretaceous Minnesoda, US Lewis, 1970b Leaf pieces Peterson and Morman Creeks Fms. Oligocene Montana, US Lewis, 1972 Conchostracan valves, fish bones/scales, pellets, bivalve fragments Wealden Group Early Cretaceous South England, UK Jarzembowski, 1995 Sand grains of Morrison Fm. Late Jurassic Colorado, US Carbonate particles Green River Fm. Eocene Wyoming, US Carbonate particles, ooids, ostracod valves, quartz grains Green River Fm. Eocene Wyoming, US Other caddisfly cases, sand grains, vegetable material ? Modern Peñalara National Park, Spain Hasiotis et al., 1998 Loewen et al., 1999 Leggitt et al., 2007; Leggitt and Loewen, 2002; Leggitt and Cushman, 2001 Boyero and Barnard, 2004 Fine sands, wood fragments, limemudstone clasts Jinju Fm. Early Cretaceous Jahyeri, Korea Paik, 2005 Plant material Kundur Fm. Late Cretaceous Fish bones and scales Wealden Group Sukatsheva, 2005 Heads, 2006 Pellets (fecal particles) Baissa Fm. Sand grains, leaf pieces Daohugou Fauna Unit Early Cretaceous Early Cretaceous Middle Jurassic Amur Region, Russia Southern England, UK Siberia, Russia Pingquan, China Ostracod carapace, bivalve fragments Jehol Fauna Unit Early Cretaceous Yixian, NE China Huang et al., 2009 Huang et al., 2009 Sand grains, leaf pieces Lushangfen Horde Fm. Early Cretaceous Lushangfen, NE China Ivanov, 2006 Huang et al., 2009 Fm. Formation. 4.2 Ichnology of Caddisfly Cases Of the better defined 11 ichnogenera proposed for caddisfly cases in terms of various grain compositions in tubular case walls (Table 2), five ichnogenera Conchindusia Vyalov and Sukatsheva, 1976, Folindusia Berry, 1928, Indusia Sukatsheva, 1993, Pelindusia Vyalov and Sukatsheva, 1976, and Piscindusia Jarzembowski, 1995 possess tube walls that are made of fragments of plant leaves or invertebrate shells; four ichnogenera Scyphindusia Sukatsheva, 1985, Secrindusia Sukatsheva, 1982, Terrindusia Sukatsheva, 1993, and Tektonargus Hasiotis in Hasiotis et al., 1998 embrace tube walls that are comrpised with sand grains or secretory material. Both Ostracindusia Vyalov, 1973 and Coprindusia Ivanov, 2006 having tube walls that are made of ovoid pellets, and thus are most allied to the Liupanshan laval cases. These two ichnogenera, however, also differ from the Liupanshan cases in having a larger size and and components of pellets. Of these, Ostracindusia, 12−14 mm long and 4 mm in diameter, embraces wall pellets having a diameter up to 0.67 mm and compositions of decalcified ostracod valves and carbonate particles. Coprindusia is 8−14 mm long and 3−4 mm wide and its wall pellets are equal in size and shape (0.3−0.4 mm in length and 0.1−0.3 mm in diameter) and comprise fecal particles of aquatic animals, which are arranged perpendicular to or at a small angle to the case axis (Gorman et al., 2008; Ivanov, 2006). As described above, the Liupanshan caddisfly cases were constructed with organic pellets which were likely made of bacterial algae and fecal particles derived from aquatic invertebrates. They are mostly 4−17 mm long and 1.5−2.9 mm in diameter (Figs. 3e−3f). These characters therefore justify a tentative assignment of the Liupanshan caddisfly cases to Coprindusia, although some minor differences exist between the Russian ichnogenus and Liupanshan materials. 4.3 Case-Makers A wide variety of candidates such as serpulids, vestimen- Exceptionally Preserved Caddisfly Larval Cases (Insecta) from the Lower Cretaceous of the Liupanshan Basin, Western China modern Oocardium possesses bifurcate tubes (Bradley, 1974). As a result, the tubular burrows built by the above organisms are readily different from the Liupanshan caddisfly larval cases. Modern caddisfly larval cases are open at both ends, although the caudal end may be partially closed, and composed of sand-size quartz grains of 2.5–30 mm in length (Leggitt and Loewen, 2002). They inhabit mainly springs, small streams, lakes, and wetlands, even brackish water (Daniel et al., 2009; Moor and Ivanov, 2008; Greenwood et al., 2003; Johansson, 1991). They aggregate when pupating or overwintering (Vvtenis, 1985). Total 29 of the extant and extinct caddisfly families contain taxa, which are capble of building cases. Of these, five families Phryganeidae, Brachycentridae, Lepidostomatidae, Calamoceratidae, and Leptoceridae are known to persist from the Early Cretaceous (Paik, 2005; Ivanov and Sukatsheva, 2002). Both the Phryganeidae and Leptoceridae possess lentic biotope (pond and lake) (Leggitt et al., 2007; Paik, 2005; Mackay and Wiggins, 1979). The larval cases of the Phryganeidae are generally larger than 20 mm in length and tiferans, dentaliid, tentaculitoid, siphonodendron, and some algae have been proposed for builders of such tubular traces, which are similar to the caddisfly larval cases. Of these, both tentaculitoid and siphonodendron are extinct marine invertebrates. The overwhelming majority of serpulids also inhabit marine niches, except Microconchus pussilus (Martin, 1809) which has been reported from terrestrial niches and possesses a rather small calcareous tube, 2–5 mm in length and 50–200 µm in diameter (Yang et al., 2015, 2011; Chen et al., 2008; Chen and Sun, 2001; Ding et al., 1993; Yu and Wang, 1983, 1981; Chen and Wu, 1979). The vestimentiferans, characteristics of the cold spring fluid faunas, are cylindric and crooked in outline and possess the tube walls constructed of cryptocrystalline calcium carbonate (Chen et al., 2007, 2006; Peckmann et al., 2005; Halanych et al., 1998). The tubes built by the dentaliids are usually preserved dispersely and their walls are variously ornamented. Oocardium is one of the few desmids growing only in highly calcareous waters, the cells of Oocardium secrete tubes of mucilage around which lime is precipitated, but Table 2 Key characteristics of selected caddisfly case ichnogenera Ichnogenus Species Size (mm) Tube wall composition References Conchindusia C. distans L: 14–23, D: 4–5 Cret.- Conchostracan valves Jarzembowski, 1995 P. sukachevae L: 17–30, D: 3–4 Cret.- Fish bones/scales Jarzembowski, 1995 O. vyalovi L: 12–14, D: 4 Cret.- Jarzembowski, 1995 Pelindusia P. percealleni L: 35, D: 5 Cret.- Pellets (ostracod valves, carbonate particles) Bivalve fragments Folindusia F. pinacea L: 23, D: 7 Jur.- Leaf fragments Lewis, 1970a L: 40, D: 8 Jur.- Secretory material Ponomarenko et al., 2009 Terrindusia S. hydroptiliformis T. notabilis L: 15–19, D: 5 Cret.- Terrigenous materials Ponomarenko et al., 2009 Coprindusia C. pallida L: 8–14, D: 3.1–4.1 L: 12–20, D: 3–4 Cret.- Pellets (fecal particles) of aquatic animals Plant material with ostracod shell fragments Sand grains Gorman et al., 2008 ; Ivanov, 2006 Ponomarenko et al., 2009 Secretary material with sand grains Ponomarenko et al., 2009 Piscindusia Ostracindusia Scyphindusia Indusia I. incredibilis Tektonargus T. kollaspilas Secrindusia S. admiranda 199 L: 11–14, D: 0.5–3 L: 23, D: 3.5 Age Cret.Jur.Cret.- Jarzembowski, 1995 Hosiotis et al., 1998 Figure 7. SEM image (left figure) and EDS analytical results (right figure) of the cemented pellets in case tube walls. White cross corresponds to analysing point. Element Au indicates gold coating of samples. 200 Xin He, Zhong-Qiang Chen, Zongsheng Lu, Jun Li, Wei Hu, Shengfu Li and Zhitao Xu composed mainly of wood fragments, whereas the larval cases of the Leptoceridae are generally 10−20 mm in length and are constructed of sand grains and some wood fragments (Paik, 2005). Judging by case size and outline as well as composition and construction pattern of cemented particles in case walls, the above two families can not accommodate the Liupanshan caddisfly larval cases. Moreover, Ponomarenko et al. (2009) suggested that a great number of lacustrine insect hordes became extinct in Cretaceous and they could also be candidates building such caddisfly cases. For example, fossil caddisfly cases such as Conchindusia, Piscindusia, Ostracindusia, and Pelindusia reported from the Early Cretaceous of southern England were believed to be built by the extinct Necrotauliidae, Vitimotauliidae, Dysoneuridae, and Plectrotarsidae (Sukatsheva and Jarzembowski, 2001; Jarzembowski, 1995). Similarly, the Liupanshan caddisfly cases may also have been built by some extinct insects. This inference is reinforced by the discovery of insect wing fossils in association with caddisfly larval cases in the studied section (Fig. 8). Hong and Li (2004) described insect wing fossils from the caddisfly case-bearing horizons in the Liupanshan Basin and assigned them to Ningxiapsyche fangi, which belongs to the Ningxiapsychidae (Trichoptera, Integripalpia). The forewing of the Ningxiapsychidae from the late Early Cretaceous (Aptian–Albian Stage) Naijiahe Formation in southern Ningxia, China, is 9.1 mm long and 3.5 mm wide (Hong and Li, 2004). Their larvae inhabited in clean lentic niches and could construct conoid, portable silk cases, while the adults live in land. They are holometabolous, typically unlvoltine (Leggitt and Cushman, 2001) and generally emergent in autumn. Pupation usually requires three weeks and the adult stage lasts 3–4 weeks (Leggitt and Cushman, 2001). As consequence, Ningxiapsyche fangi is the candidate for the potential trace-makers who produced the Liupanshan caddisfly cases (Fig. 8). They often inhabited nearshore environments, like other Trichopteriods (Paik, 2005; Leggitt and Cushman, 2001). Figure 8. The forewing of Ningxiapsyche fangi uncovered from the same horizon as caddisfly cases (after Hong and Li, 2004). 5 CONCLUSIONS Small, tubular and unique tube wall compositions indicate that the tubular fossils of Aptian–Albian age (Early Cretaceous) from the Liupanshan Basin, western China are assignable to fossil caddisfly larval case. They resemble both modern caddisfly cases and fossil counterparts. The case wall comprises inner, medium and outer layers. Both inner and outer layers are composed of organics, while the medium layer consists of tiny particles of cryptocrystalline pellets. The organic pellets were regularly arranged and perpendicular to the axis of the tube. Several major ichnologic features such as small size, tubular shape and tube walls constructed with ovoid pellets indicate that the Liupanshan caddisfly larval cases are tentatively assignable to ichnogenus Coprindusia. The co-occurrence of insect forewing fossils of Ningxiapsyche fangi and larval cases in Liupanshan suggests that the former insects could be the candidate for case-builders accountable for these exceptionally preserved caddisfly larval cases. They inhabited a highly evaporitc, salty lake during the Early Cretaceous. ACKNOWLEDGMENTS The authors are grateful to both Xing Huang and Lu Jia for preparing thin-sections, both Yongling Chen and Lei Shi for assistance in SEM imaging analysis, and Bin Chen for assistance in photographing these tiny fossils. Two anonymous reviewers are thanked for their constructive suggestions, which have improved greatly the quality of the paper. This study was supported by the 973 Program of China (No. 2011CB808800), the 111 Program of China, Ministry of Education of China, the National Natural Science Foundation of China (No. 41272023), and the research grant from the State Key Laboratory of Biogeology and Environmental Geology (No. GBL11206). It is a contribution to the special issue celebrating the 80th Birthday of Professor Hongfu Yin, China University of Geosciences (Wuhan). REFERENCES CITED Bai, S. M., Li, H. Q., Mao, Z. L., 2003. The Chemical Features of Cretaceous Earth Layer in Liupan Mountain Area in China. 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