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. 2024 Dec 28;14:31506. doi: 10.1038/s41598-024-83210-z

Discovery of fossil avian footprints from Late Holocene sediments of Allahbund uplift in Great Rann of Kachchh of Western India

Darshit Padia 1, Bhawanisingh Desai 2,✉, Suruchi Chauhan 2, Babulal Vaghela 1
PMCID: PMC11682152  PMID: 39732983

Abstract

The Great Rann of Kachchh is a sabkha terrain with a thick succession of Quaternary to Late Holocene sediments, deposited during high sea level after the Last Glacial Maxima. Geomorphologically, the Great Rann of Kachchh is subdivided into Bet Zone, Linear Trench Zone, Great Barren Zone, and Banni Plain. The Bet zone is a slightly elevated flat surface comprising a complex network of bets and interbet channels—the geomorphic entities developed as complex interplay of sea level and coseismic tectonic activity during the Holocene. Moreover, during the last 5.5 to 2 ka, the central part of the western Great Rann was under the influence of tidal flat sedimentation. The present study at Karimshahi Bet/ Allahbund (the western portion of the Bet zone) records the first group of avian tracks of shorebirds from Late Holocene sediment preserved at a depth of 15 cm below the present-day Rann surface. According to the morphological and preservational study, the footprints are classified as cf. Gruipeda, suggesting the trace maker as waders (Shorebirds) foraging on the shoreface. The discovery of avian trackways and abundant shallow marine ichnoassemblage within the studied unit suggests that the area was flooded by low-energy water, forming a favourable habitat for waders (shorebirds) to forage shoreline. Thus, the work describes the first-ever record of avian shorebird tracks from the Late Holocene mudflat succession of Great Rann of Kachchh, Western India.

Subject terms: Palaeoecology, Behavioural ecology

Introduction

The Great Rann of Kachchh (~ 300 km in length and 80–100 km in width, Fig. 1a and b) is a vast saline flat terrain preserving thick Quaternary succession from a post-Last Glacial Maxima high sea level onwards1–4.These fine-grained sediments are inferred to be deposited in shallow marine especially in estuarine and intertidal marshy deposition environments4–8. The area also houses several Bronze Age sites of Harappan civilisation that flourished in and around Great Rann of Kachchh, this includes the large town of Dholavira on Khadir Island. Multiple studies have suggested on high sea levels during the middle Holocene4,9,10 and a navigable route through the Rann. Additionaly, the Early Iron Age to Medieval (~ 3100 − 900 year B.P.) period archeological sites provided evidence for cultural collapse and human migration from west to east due to monsoon failure11. Thus, the Quaternary Great Rann of Kachchh sediments are the repositories of Holocene paleoenvironmental changes archiving the transition of depositional gradient from shallow marine to fluvial environment8,12,13.

Fig. 1.

Fig. 1

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(a) Simplified outline map of India showing Gujarat state and Kachchh district marked with box (outline maps are downloaded from free maps-open source (https://d-maps.com). (b) SRTM-DEM of Kachchh showing geomorphic units bordered by major faults (fault positions are inferred from26), geomorphic subdivisions of Great Rann of Kachchh, and studied location marked by a yellow colored circle. Abbreviation: GRK: Great Rann of Kachchh, PU: Pachham Uplift, KU: Khadir Uplift, BU: Bela Uplift, CU: Charor Uplift, WU: Wagad Uplift. KMF: Kachchh Mainland Fault, KHF: Katrol Hill Fault, IBF: Island Belt Fault, ABF: Allahbund Fault, GF: Gedi Fault, NPF: Nagar Parkar Fault, SWF: South Wagad Fault. The SRTM-DEM data was downloaded from the USGS Earth Explorer Interface (http://earthexplorer.usgs.gov/) and the figure was generated with QGIS v. 3.34.1 (https://www.qgis.org), and licensed CorelDRAW v.22.1.1.523 (https://www.corelindia.co.in/).

Trace fossils, such as tracks, trails, and burrows, are invaluable records of ancient life, offering unique insights into trace makers’ behaviours, interactions, and adaptations14. They serve as a window into sediment records, helping to unravel paleoenvironmental mysteries. Among these, avian tracks and trackways are essential, providing a snapshot of avian behaviour and revealing insights into locomotion, social dynamics, and environmental interactions. The occurrences of Avian, Mammals and Human footprints within Quaternary, especially the Pleistocene and Lower Holocene sediments are globally reported15–25. However, avian footprints within upper Holocene sediments seldomly reported owing to its proximity to the recent environment. In this context, our research presents a significant finding: the first-ever record of avian shorebird tracks from the Late Holocene mudflat succession in the Great Rann of Kachchh, Western India are also contemporary with Harappan Civilisation.

Geological setting of the Great Rann of Kachchh

The Great Rann of Kachchh is structurally part of an east–west-trending paleo rift graben, bounded by the Nagar Parker Fault in the north and the Kachchh Mainland Fault and South Wagad Fault in the south. Its eastern extent is limited to the West Cambay Basin Margin Fault. The basin opens into the Arabian Sea and the Indus Delta region to the west. The gentle gradient of the Rann surface allows for inundation by seawater from the Arabian Sea through Kori Creek (Fig. 1). The region is also flooded by rivers from the east and surrounding highlands.

Due to heterogeneity in topography and inundation patterns, the Great Rann of Kachchh can be subdivided into four geomorphic facets: Bet Zone, Linear Trench Zone, Great Barren Zone, and Banni Plain. The Bet zone is a slightly elevated flat surface comprising a complex network of bets (islands) and interbet channels occupying the northwestern portion of the Great Rann of Kachchh. It consists of several bets of different sizes, with a Rann surface occupying the interbet depression. The area is bounded by Kuar and Bediya bets in the east; towards the north, it is limited by sand ridges of Sind. The Allahbund Fault scarp restricts the Bet zone and marks the southern boundary, while in the west, it merges to the Indus Delta plains. Historically, the 1819 Allah Bund earthquake was the most significant event, creating a 80–90 km long fault scarp (known as Allahbund Fault Scarp) with variable height of 4–6 m27–31. The uplifted fault scarp is today preserved as geomorphic high with ravines, its effect of sudden uplift which is well documented in historic accounts caused blocking of the distributary of the Indus River and submergence of the Sindri Fort in the Great Rann of Kachchh27–30. Moreover, Rann sediments consists abundant fossils traces in the form of burrows, track and trails, with dominance of ichnogenera like Arernicolites, Palaeophyscus, Planolites, Psilonichnus, Skolithos, Teichichnus and Thalassinoides8.

Materials and methods

Our research employed a meticulous approach to identify the paleoenvironmental and paleo-ecological conditions during the Mid-Late Holocene period. Multiple field visits were conducted in the Western Great Rann of Kachchh during summers of 2019 to 2023 for understanding ichnological variation and associated ichnoassemblages. The Karimshahi site in Allahbund uplift (24° 4′ 23.36′′ N; 69° 18′ 31.31′′ E) offers an abundance fossil traces prominently visible on the top surface. The site’s sedimentary composition is unconsolidated and hence, trenching technique was the most effective approach for examining subsurface sediments and their associated trace fossils. Trenches were excavated using traditional hand tools, including mattocks/pickaxes, spades, and shovels. A vertical and horizontal scraping of mm to cm thick layer technique was adopted to delineate burrow dimensions in cross-section accurately (Fig. 2). This enabled us to determine the depth of penetration of fossil traces and reveal the burrow’s outline, enabling the detailed assessment of its dimensions. These observations were meticulously recorded to reconstruct the complete burrow morphology, essential for identifying the associated ichnogenera and ichnospecies (Fig. 3). Vertical and horizontal scraping and slicing were conducted concurrently for inclined burrows to obtain a comprehensive view of the burrow structure. Consequently, the morphological characteristics of the trace fossils were documented in the field through continuous observation and photographic documentation. The photographs of the trace fossils were later subjected to detailed morphological analysis using ImageJ, an open-source software widely utilized for image processing and analysis. During horizontal slicing at an approximate depth of 15 cm, approximately 10 partially to moderately preserved avian foot prints, produced by a bipedal trackmaker represented by concave epirelief were encountered on the surface consisting of micaceous sandy silt were discovered (Fig. 2D). Out of which 8 footprints KA-1, KA-2, KA-3, KA-4, KA-5, KA-6, KA-7 and KA-8 were selected for the measurement purpose (Fig. 4; Table 1). False color images for two moderately preserved avian tracks were processed and developed using ImajeJ software for detail understanding of footprint digits morphology (Fig. 5).Moreover, to determine the nature of sediments, the samples were collected and analysed using pipette method. The calculated proportion of Sand, silt and clay were plotted using SEDICLASS (Sediment classification software) developed by USGS. Moreover, to determine the nature of sediments, the samples were collected and analysed using pipette method. The calculated proportion of Sand, silt and clay were plotted using SEDICLASS (Sediment classification software) developed by USGS.

Fig. 2.

Fig. 2

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Field photographs showing (A) The view of the study area, (B) Studied Site, (C) Removal of surface using horizontal Slices techniques to understand burrow morphology and (D) Occurrences of avian foot print while performing horizontal slicing technique.

Fig. 3.

Fig. 3

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Photographs (A and B) showing trace fossils associated with studied site; Arenicolites (Ar), Palaeophycus (Pa), Skolithos (Sk) and Thalassinoides (Th); (C) closeup view of horizontal burrow; Palaeophycus (D) isolated vertical burrows; Skolithos (Sk).

Fig. 4.

Fig. 4

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On site photographs of the avian tracks (A and B) and the sketches of the avian footprints.

Table 1.

Measurements of the bird tracks of Karimshahi Bet/Allahband:

Sr. No TL FW II III IV II^III III^IV II^IV FL/FW
KA-1 67 75 31 41 31 66 67 133 0.89
KA-2 65 74 31 42 30 63 66 129 0.87
KA-3 32 47 14 23 15 64 71 135 0.68
KA-4 - - - 26 18 - 70 - -
KA-5 54 61 31 41 32 68 71 139 0.88
KA-6 - - - 35 - - - - -
KA-7 42 44 13 29 16 63 66 129 0.95
KA-8 43 - - 28 17 - 67 - -
Mean 51 60 24 33 27 65 69 134 0.85
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Fig. 5.

Fig. 5

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Close up view of avian tracks and false color images of two moderately preserved footprints (all three forward digits (I, II, III) with moderately preserved hallux impression can be observed).

Ichnology

Karimshahi site sedimentary exposures contain trace fossils such vertical and horizontal burrows, three-dimensional burrows (boxwork) (Fig. 3), and bird footprints (Fig. 4). The vertical burrows are identified as Skolithos, a thin, unbranched, vertical or oblique burrow. While Arenicolites occurs as vertical U-tube burrows without sperite. Palaeophycus occurs as horizontal burrows while the three-dimensional burrow systems were identified as Thalassinoides8. Slender Skolithos burrows are made by polychaetes commonly associated with low-water lines of an intertidal zone32 in addition, Skolithos can also be found in shallow marine to marginal marine environments33,34. Arenicolites is a dwelling trace produced by suspension-feeding worms35 or by deposit-feeding worms, including polychaetes36, and similar burrows are produced by suspension-feeding polychaetes in a recent intertidal environment32. Palaeophycus and Thalassinoides are common in intertidal flats and other marginal marine environments37,38. The authors were fascinated by the unique occurrence of the bird footprints coupled with shallow marine/marginal marine burrows.

The morphological study of the footprints suggests the possible trace maker as bird waders. Waders, often known as shorebirds/wetland birds that search for food by crawling or burrowing in the mud/sand and have diverse foraging tactics that depend on food sources39. Bird footprints are difficult to identify to a specific species level, but they can provide valuable information such as the organism’s behavior and the habitat in which it was living40. Such footprints are formed when birds walk across soft (likely wet) mud or sand that hardens enough to retain the impressions before the next layer of sediment is deposited. During fieldwork at the fringe of Karimshahi bet/Allahbund of the Great Rann of Kachchh, such bird tracks were discovered. The Trace fossils produced due to the locomotion activity of organisms are ethologically classified as repichnia41,42 These traces are basically bedding-plane trace fossils created at the sediment–water interface or along lithological interfaces. The birds mostly feed on tiny amphibians, arthropods such as aquatic insects or crustaceans found in shallow wetlands. Wading birds have different feeding strategies, so their bills are adapted to variable shapes and functions, for example, filter feeding, pecking on sediment and foraging inshore sediments are the main feeding strategies43–45. These birds typically have long, slender legs that allow them to walk slowly and effortlessly through the water and the beak is mostly used for finding and catching food, filtering food from water and carrying or killing prey. Waders have three toes pointing forward and one pointing backwards. The size and shape of the foot helps the waders walk through mud/sand without losing their balance. Several paleoichnological avian tracks also records similar behaviour40,46–50.

Ichnotaxonomy of bird-like footprints

Following the ICZN (1999, Art. 10.3), the use of ichnofamilies is explicitly permitted, and they are recognized as valuable tools for identifying groups of tracks that share significant morphological characteristics. Despite their potential utility, the classification of avian footprints into ichnofamilies is relatively uncommon. The initial attempts to do so can be attributed to51, who introduced two ichnofamilies: Ignotornidae and Trisauropodiscidae. Following this52,53 established five morphofamilies for avian tracks (Gruipedidae, Charadriipedidae, Avipedidae, Anatipedidae, and Culcitapedidae), a classification later supported by54 and55. Although morphofamilies appear synonymous with ichnofamilies cf56. , the latter term is preferred in this discussion despite not being explicitly defined by52. The family Ignotornidae was recently revised56, and a new family, Koreanornipodidae, was introduced57.

According to a revised definition of ichnofamilies for avian footprints, there are two main approaches: (a) Lockley and coworkers use an extensive range of features and detailed descriptions (digit count, morphology, and orientation; footprint symmetry and size; number and presence of pad impressions and claw marks; presence and morphology of webbing; rotation of footprint, and stride length); and (b) Sarjeant and coworkers employ a few key features (digit count, relative position, and proportionate length of digits; and the presence or absence of webbing impressions) to distinguish ichnofamilies. At the ichnogeneric and ichnospecific levels, certain features have been more frequently utilized as ichnotaxobases51,52,54,58–63. For example, the number, morphology, relative length, and orientation of digit impressions or the total divarication are commonly used as ichnogeneric taxobases. Some authors have also considered features such as the footprint length ratio51,61, divarication between the impressions of digits II-III and III-IV52, the presence or absence of sole62 or heel impressions52,54,63,] webbing52,54,63 and the alignment of digit I with digit III52,63. Based on this, the most useful ichnogeneric taxobases for distinguishing avian-like footprints are proposed to be; (1) the number, morphology, relative length, and orientation of digit imprints, specifically of digit I; (2) the length ratio of footprints; (3) the number of phalangeal pad marks; (4) the divarication between the impressions of digits II-III, III-IV, and II-IV; and (5) the presence or absence of webbing.

The described avian tracks are comparable with the ichnofamily Gruipeda due to their tetradactyl form, with three forward-facing digits (II-IV) and a smaller, backward-facing hallux (I) and due to its relative digit lengths (I < II < IV < III). In addition, the present forms have slender digit impressions along with divarication angles between digits II and IV less than 140°.

Analysis of avian footprints

The Ichnotaxobases proposed in this study include the length ratios of footprints, footprint size range, the proportion of digit lengths I: II: III, and the range of angles between digit impressions (Table 1). All measurements on these tracks follow the norms and methodologies established by64,65 (Fig. 6). The measurements from identified avian tracks are summarized in Table 1.

Fig. 6.

Fig. 6

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Diagram illustrating the footprint and trackway measuring conventions used in this paper. A, parameters related to isolated tracks; B, detail of the measured angles between the impression of the digits I, II, III, and IV. (Abbreviation: TL stands for total footprint length, which is the distance between the distal points of digit III and the point where the hallux projection and the digit III axis prolongation intersect; W, the footprint width, is the measurement made perpendicular to the footprint axis between the distal tips of digits II and IV; I, length of digit I; II, length of digit II; III, length of digit III; IV, length of digit IV; II-IV, angle formed by the axis of digits II and IV. II-III, angle formed by the axis of digits II and III; III-IV, angle formed by the axis of digits III and IV).

Ichnofamily: Gruipeidae Panin and Avram66.

cf. Gruipedaisp.

Description

Avian tracks consist of four digits, three of which (II to IV) are directed forward and larger, and the fourth (I) is directed backwards, spur-like and short (Fig. 5). The tracks range from 44 to 75 mm in width and 32 mm to 67 mm in length. Digit impressions are slender, with a maximum width of 6 mm and may exhibit a tapered outline. The average length of the digits III, IV, and II imprints are decreasing. The average divarication of the impression of digits II and IV is 134°. The angles formed by the digits II and III, and by the III and IV, have an average difference of about 4°, about 65° and 69°, respectively. Two tracks resemble tetradactyl, showing a small hallux impression placed 5–8 mm behind digit III’s sole or proximal end. The remaining measured tracks are tridactyl, lacking the hallux impression. Thus, the interdigital angles between digits II and III and between digits III and IV are commonly less than 70°. The axis of digit I does not correspond with that of digit III, the interdigital angle between digits I and II being smaller than that between digits I and IV. Relative digit length is I < II < IV < III. Identification at the ichnospecies level was not performed because the tracks are not well-preserved.

Remarks

The present forms (cf. Gruipeda) have slender digit impressions along with divarication angles between digits II and IV. The tracks’ dimensions (width: 44–75 mm; length: 32–67 mm), slender digit impressions, and divarication angles (e.g., 134° between digits II and IV). However, due to lack of full preservation we consider it as similar to Gruipeda. Moreover66, described many new avian ichnotaxa from the Miocene of Romania, classifying them based on the possible tracemaker52 amended66 systematic approach to making the ichnotaxa independent of distinct avian groups and modified their original ichnogeneric diagnosis. The ichnogenus Gruipeda66 can be comprehended from other ichnotaxa having birdlike characteristics67. Redefined the ichnogenus Plesiornis58, from the Triassic of the United States as a monospecific ichnogenus, P. pilulatus, to include additional tracks from the Late Triassic-Early Jurassic of Poland. It can be differentiated from Gruipeda by having a U-shaped appearance, thicker digit imprints, less total divarication, and footprints in the forward direction.

Archaeornithipus62, from the Late Jurassic-Early Cretaceous of Spain, shares some morphological similarities with Gruipeda, but its hallux impression is proportionally shorter and the length-to-width ratio is higher, making it unique. The ichnotaxa Ignotornis68, from the Cretaceous of the US, as well as Jindongornipes51,69, and Koreanornis69, both from the Early Cretaceous of Korea, are quite similar in general appearance, but their ichnotaxonomic status needs a revision due to their unique features such as more pad impressions in the digits.

Aquatilavipes61, initially described from the Cretaceous of Canada and later identified in various other regions (e.g., The Cretaceous of the U.S.A51. , China70, Japan71), includes tracks with a general morphology similar to Gruipeda but lacks a hallux impression. The ichnogenus Yacoraitichnus72, 73, from the Late Cretaceous of Argentina, represents avian-like footprints with high digit divarication (II-IV) and clear claw impressions, this was revised by50 as Gruipeda. However in many reports, poor preservation of the type material and the absence of a detailed description and illustration make comparisons with other avian footprints difficult. Another ichnogenus from the Late Cretaceous of Argentina is Barrosopus73, distinguished from Gruipeda by the separation of digit II from the impressions of digits III and IV and the absence of a hallux imprint.

Tracks from the Upper Eocene of the western Pyrenees, Europe, have been categorized into six morphotypes (1 to 6)74. Some specimens in Charadriipeda ichnosp. 1 and ichnosp. 2 exhibit tridactyl and tetradactyl footprints with slender digit imprints and wide divarication. Given the emended diagnosis of Charadriipeda by52, these specimens are not comparable with this ichnogenus due to the presence of hallux impressions. Based on general morphology, they are considered to have an affinity with Gruipeda. The ichnogenus Carpathipeda, from the Neogene of Romania, represents tridactyl footprints with slender digit impressions, distinguished from Gruipeda by the absence of a hallux impression and greater asymmetry relative to the axis of digit III. Aviadactyla75, from the Miocene of Hungary, exhibits slender digit impressions, occasional claw and pad marks, and a total divarication exceeding 95°, but lacks a hallux impression. Tetraornithopedia75, also from the Miocene of Hungary, is a tetradactyl avian footprint that could potentially be included within Gruipeda, though the published illustrations do not provide enough detail for a conclusive comparison. Detailed description and illustration are crucial for making accurate comparisons.

The ichnogenus Ardeipeda66, from the Miocene of Romania, resembles Gruipeda but has a more extended digit I impression and an angle between digits I and III approaching 180°. Iranipeda76, was established to include the ichnospecies Urmiornis abeli76,77, from the Pliocene of Iran52. Subsequently synonymized it with Gruipeda, however it was later rejected. A new species of Gruipeda, G. lambrechti were described from Eocene of Birjand area of Iran, distinguished as large size tracks with digit -II, III and IV having identical size and digital impressions having recognizable phalanges78. Finally, Alaripeda53, from the Miocene of California, differs from Gruipeda by having curved digit II-IV impressions and a large divarication between digits II and IV (180°). Uvaichnites are enormous mesaxonic tridactyl bird track with a prominent central pad and not proximally connected digits and angle of divarication between digits II–III is more than 100 degrees79. Tracks are asymmetrical due to angle between digits III and IV is larger than the angle between digits II and III. Webbing and claw impression are usually absent. Thus the present specimens differs from Uvaichnites in having presence of hallux and connected digits to central pad.

Ichnological association

The avian trackways co occurs with other invertebrate trace fossils like Skolithos, Palaeophycus and Thalassinoides. The occurrence of these invertebrate trace fossils suggests that the environment was colonized by annelids and crustaceans. Polychaetes and Crustaceans are common trace makers of Skolithos, Palaeophycus and Thalassinoides in a shallow marine intertidal environments8. The occurrence of avian trackways along with these invertebrate trace fossils suggests avian gregarious behavior, including “shuffling” and “stop-start” progression of foraging activity80 probably these shorebirds visited the area for their foraging activity.

Description of track bearing unit

Karimshahi site consist two distinct units; the bottom unit-1 consist of gray coloured clayey silt, overlain by 135 cm thick unit-2 composed of micaceous sandy silt with numerous thin intercalated clay bands in the form of flaser and wavy bedding (Fig. 7A). Grain size distribution is dominated by (1) Clayey silt and (2) Sandy silt. However, the silt remains the dominant fraction throughout the section (Fig. 7B). According to4, micaceous sand was deposited by southern draining streams (Himalayan source) in western Great Rann of Kachchh is through Indus River channel (Nara River Channel). Moreover, the silty-clay sediments were deposited during tidal flat environment, supported by grain size analysis and suggested by4,8.

Fig. 7.

Fig. 7

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(A) field photos showing falser and wavy bedding, and (B) Grain size Plot (Shepard Plot) showing dominating Sandy Silt/Clayey Silt fractions.

Discussion

Avian tracks

Bird and Mammal footprints were revised by66 and were later redefined in to families by52,53. Tracks of shorebird footprints, including Gruipeda dominguensis and cf. Alaripedia isp, have been described in the fine-grained sediments deposited within the low-gradient fluvio-lacustrine environment of the San Domingo Formation (Late Triassic-Early Jurassic) of Argentina46. However, these track bearing sediments are later dated to be of Late Eocene81. Shorebirds tracks along with feeding and invertebrate traces were described from Early Cretaceous of Haman Formation in the Gyeongsang Basin of Republic of Korea, the results documents feeding behaviour of the shorebirds, by pecking and probing for foraging40. The Yacoraite Formation (Maastrichtian-Danian) of Argentina contains a diverse assemblage of shorebird tracks, the results suggest shorebird tracks dominate moderate to low energy ephimeral ponds, and shorebird ichnosubfacies are subset Scoyenia ichnofacies50. Cretaceous shorebird tracks from Upper Cretaceous Dakota Formation of Utah also provides similarity between Mesozoic and Cenozoic avian ichnotaxa and early evolution of shorebird species82. Cretaceous avian birds tracks from Dakota Group of USA occur along with dinosaur and crocodile dominated track assemblages, representing small herons and plover birds, suggests gregarious behavior, including “shuffling” and “stop-start” progression of foraging activity82.

Avian ichnotaxa (Avipeda) of shorebirds visiting saline playa lake shorelines were described from the fine-grained dolomitic carbonates of Copper Canyon Formation (Neogene) from Death Valley National Park83. The results suggested that the ancient saline lake was frequented by several birds, equids, felids and other mammals, diversity and density of the tracks fluctuated with the salinity of the lake. Further, four different categories of substrates preservation based on water content were identified, namely basic categories of water saturation were identified and includes water saturated, water unstaturated, moist-damp and dry83. A small wader bird imprints with long hind toe were also described from Early Eocene of Messel of Germany84. Recurvirostid-like bird with long legs and short toes (Similar to Flamingoes) were descrbied from early Eocene fossil track locality in Utah85. Shorebird footprints were described from upper Eocene tidal flat deposits of Liedena Sandstone, Navarre74. A tracksite with bird footprints is described from Lower Miocene of Lerin Formation outcropping at Bardenas Reales de Navarra Natural Park (Navarre, Spain), the tracks were formed along the shores of paleo perilacustrine deltaic systems and distal alluvial systems in Ebro Basin86. Solitary and group avian tracks of shorebirds were extensively described from the Late Miocene of the Sorbas Basin, Spain, considering all the tracks in shorebird ichnofacies63. Ichnogenus Aquatilavipes were described from the Oligocene Sawahlunto Formation of Sumatra, suggesting birds occupied the intertidal flats for foraging purposes87. Two avian footprints of intermediate-sized modern gulls were reported from Pleistocene oolitic-bioclastic grainstone of end regressive cycle from San Salvador island of Bahamas88.

Neoichnological studies of avian/shorebird tracks along coastal freshwater pond (the Bajo de los Huesos, Chubut, Argentina) were compared with the Late Triassic-Early Jurassic Santo Domingo Formation tracks46. The extensive shorebird track studies along the freshwater ponds converge twenty one behaviour into four morpho-types of modern footprints, additionally suggesting bird tracks forms parallel to the shoreline and distinction between modern and shallower part of pond89. Spatial analysis of the shorebird trackways using low-level drone photogrammetry was conducted for compiling neoichnological dataset for paleoenvironment interpretation89. The detailed neoichnological studies allowed surface orthomosaics and elevation models for correlating track morphology (track density, stride orientation, stride length) with ecological factors.

The co-occurrence of Gruipeda with invertebrate traces suggests a dynamic interplay between avian and invertebrate activities in coastal ecosystems. Gruipeda footprints indicate the presence of wading birds foraging in shallow waters, while the invertebrate traces reflect burrowing and feeding behaviours in the same sediments. This ichnoassociation is typically characteristic of shallow marine, lagoon, and tidal flat environments, where both avian and invertebrate activities are prevalent. Such Ichnoassemblage has been documented from Late Triassic–Early Jurassic46? , Cretaceous50,90, Eocene91–94, Early to middle Eocene95, Oligocene96, Miocene97, Pleistocene98, Late Quaternary24 and from Holocene (Present paper). The results suggested direct correlation between macrobenthic invertebrate prey availability and shorebird track.

Paleoenvironment of Great Rann of Kachchh sediments

The geomorphic setting of the Great Rann of Kachchh is attributed to the complex interplay of sea level and coseismic tectonic activity during the Holocene4,8,13,30,99–101. The study carried out by9 on the Core raised from Dhordo site and its analysis of Nd and Sr isotopic compositions suggest that the large Himalayan and sub-Himalayan Saraswati-like river may have discharged into the Arabian Sea until early Holocene. During the period of 5.5–2 ka, the major part of the western Great Rann was under the influence of flat tidal sedimentation (Fig. 8)4,99,101. However, the western Great Rann of Kachchh witnessed gradually fluctuating palaeoenvironmental conditions from the dominance of marine to brackish water to terrestrial during the Mid-Late Holocene period8.

Fig. 8.

Fig. 8

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(a) Block diagram showing various depositional environments and studied site, marked with a red-colored box. (b) Stratigraphy of the studied site along with the position of buried foot print with co-occurring ichnoassociation.

Based on the Stratigraphic correlation, and OSL dating of the sediments, the top of the Karimshahi bet is dated as 3.0 + 0.3 Ka age4.Our ichnological data including avian footprint is also derived from the same succession suggesting the synchronous formation of the tracks indicating its formation during Late Holocene period. The grain size, sedimentary structure and geochemistry of the Karimshahi sediment were envisaged to be deposited in active tidal flat environment during Late Holocene4.

At present Great Rann of Kachchh is a monotonous, salt-encrusted, vast mudflat, largely dried up during early summer (i.e., March–July) and inundated during the summer monsoon and winter season (i.e., July to February). Strong summer monsoon winds push seawater from the Arabian Sea into the Great Rann of Kachchh; usually the water does not evacuate or evaporate until the next summer. Hence, the low-lying western Great Rann opens into the Arabian Sea through a macro tidal dominated Kori creek, which serves as a conduit for sea water and storm-induced tidal surges to enter inland that result in the flooding of the Great Rann of Kachchh.

Conclusion

The Holocene sediment of Karimshahi Bet succession shows diverse trace fossil assemblage of Arenicolites, Skolithos, Palaeophycus, and Thalassinoides with co occurrence of bird foot print cf. Gruipeda at the depth of approx. 15 cm. At present the study site is above the present-day tidal inundation limit. Moreover, the abundance of invertebrate burrows attracted shoreface birds (especially waders) for foraging on these invertebrate trace makers. The present avian footprint from Late Holocene at Karimshahi indicates the low energy tidal environment favourable for shorebirds to forage on the invertebrates. These results emphasize the need for more research and preservation of paleontological sites, enhancing our understanding of both past and present environment. The Great Rann of Kachchh, with its diverse geological and paleoenvironmental history, demonstrations how environmental changes have shaped the land in human history. It highlights the importance of ongoing research in paleontology and ichnology to better understand the paleoenvieonmental and paleoecological history of the area.

Acknowledgements

The authors are thankful to Border Security Force (BSF) for providing necessary permission to carry out field visits and authors (DP, BKV) are thankful to Shri R. R. Lalan College for granting research approval and facilities. Authors (BGD and SC) gratefully acknowledges Pandit Deendayal Energy University, Gandhinagar. Authors are grateful to Dr. Ignacio Diaz Martinez; Dr. Andrew Rindsberg and one anonymous reviewer for improving the manuscript.

Author contributions

DP: Data collection, Fieldwork, result analysis, Manuscript textBGD: Data collection, Fieldwork, Result analysis, Manuscript text, SP: Result analysis, Manuscript textBV: Fieldwork, Manuscript textAll authors revieed the manuscript.

Data availability

The dataset used and/or analysed during the current study available from the corresponding author on a reasonable request.

Declarations

Competing interests

The authors declare no competing interests.

Footnotes

Publisher’s note

Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.

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