Hukam Singh

Biogeographic and evolutionary implications of a diverse paleobiota in amber from the early Eocene of India

For nearly 100 million years, the India subcontinent drifted from Gondwana until its collision with Asia some 50 Ma, during which time the landmass presumably evolved a highly endemic biota. Recent excavations of rich outcrops of 50–52-million-year-old amber with diverse inclusions from the Cambay Shale of Gujarat, western India address this issue. Cambay amber occurs in lignitic and muddy sediments concentrated by near-shore chenier systems; its chemistry and the anatomy of associated fossil wood indicates a definitive source of Dipterocarpaceae. The amber is very partially polymerized and readily dissolves in organic solvents, thus allowing extraction of whole insects whose cuticle retains microscopic fidelity. Fourteen orders and more than 55 families and 100 species of arthropod inclusions have been discovered thus far, which have affinities to taxa from the Eocene of northern Europe, to the Recent of Australasia, and the Miocene to Recent of tropical America. Thus, India just prior to or immediately following contact shows little biological insularity. A significant diversity of eusocial insects are fossilized, including corbiculate bees, rhinotermitid termites, and modern subfamilies of ants (Formicidae), groups that apparently radiated during the contemporaneous Early Eocene Climatic Optimum or just prior to it during the Paleocene-Eocene Thermal Maximum. Cambay amber preserves a uniquely diverse and early biota of a modern-type of broad-leaf tropical forest, revealing 50 Ma of stasis and change in biological communities of the dipterocarp primary forests that dominate southeastern Asia today.

High bat (Chiroptera) diversity in the Early Eocene of India

The geographic origin of bats is still unknown, and fossils of earliest bats are rare and poorly diversified, with, maybe, the exception of Europe. The earliest bats are recorded from the Early Eocene of North America, Europe, North Africa and Australia where they seem to appear suddenly and simultaneously. Until now, the oldest record in Asia was from the Middle Eocene. In this paper, we report the discovery of the oldest bat fauna of Asia dating from the Early Eocene of the Cambay Formation at Vastan Lignite Mine in Western India. The fossil taxa are described on the basis of well-preserved fragments of dentaries and lower teeth. The fauna is highly diversified and is represented by seven species belonging to seven genera and at least four families. Two genera and five species are new. Three species exhibit very primitive dental characters, whereas four others indicate more advanced states. Unexpectedly, this fauna presents strong affinities with the European faunas from the French Paris Basin and the German Messel locality. This could result from the limited fossil record of bats in Asia, but could also suggest new palaeobiogeographic scenarios involving the relative position of India during the Early Eocene.

An ailuravine rodent from the lower Eocene Cambay Formation at Vastan, western India, and its palaeobiogeographic implications

A new ailuravine rodent, Meldimys musak sp. nov. (Mammalia: Rodentia, Ischyromyidae), is recorded from the lower Eocene lignites of western India. It is the oldest record of Rodentia from India. M. musak is more derived than the earliest Eocene ailuravine Euromys cardosoi from Portugal and more generalized than late early Eocene E. inexpectatus and Ailuravus michauxi from France. Its dental morphology closely corresponds to the middle early Eocene species M. louisi, which lived about 52 Ma in Western Europe. Meldimys was previously known only from Europe, and ailuravines were previously reported only from Europe and North America. Its occurrence in India allows the first direct correlation between the early Eocene land mammal horizons of Europe and India, and raises the possibility of a terrestrial faunal exchange between India and Eurasia close to the Palaeocene—Eocene transition.

A Diverse Snake Fauna from the Early Eocene of Vastan Lignite Mine, Gujarat, India

The early Eocene (Ypresian) Cambay Formation of Vastan Lignite Mine in Gujarat, western India, has produced a diverse assemblage of snakes including at least ten species that belong to the Madtsoiidae, Palaeophiidae (Palaeophis and Pterosphenus), Boidae, and several Caenophidia. Within the latter taxon, the Colubroidea are represented by Russellophis crassus sp. nov. (Russellophiidae) and by Procerophis sahnii gen. et sp. nov. Thaumastophis missiaeni gen. et sp. nov. is a caenophidian of uncertain family assignment. At least two other forms probably represent new genera and species, but they are not named; both appear to be related to the Caenophidia. The number of taxa that represent the Colubroidea or at least the Caenophidia, i.e., advanced snakes, is astonishing for the Eocene. This is consistent with the view that Asia played an important part in the early history of these taxa. The fossils come from marine and continental levels; however, no significant difference is evident between faunas from these levels. The fauna from Vastan Mine includes highly aquatic, amphibious, and terrestrial snakes. All are found in the continental levels, including the aquatic palaeophiids, whereas the marine beds yielded only two taxa. Vastan Mine is only the second locality in which the palaeophiids Palaeophis and Pterosphenus co-occur. The composition of the fauna from Vastan is on the whole similar to that of the early Eocene of Europe; however, comparisons with early Eocene faunas of other continents are not possible because they are poorly known or unknown.

Ectomycorrhizas from a Lower Eocene angiosperm forest.

The development of mycorrhizal associations is considered a key innovation that enabled vascular plants to extensively colonize terrestrial habitats. Here, we present the first known fossil ectomycorrhizas from an angiosperm forest. Our fossils are preserved in a 52 million-yr-old piece of amber from the Tadkeshwar Lignite Mine of Gujarat State, western India. The amber was produced by representatives of Dipterocarpaceae in an early tropical broadleaf forest. The ectomycorrhizas were investigated using light microscopy and field emission scanning electron microscopy. Dissolving the amber surrounding one of the fossils allowed ultrastructural analyses and Raman spectroscopy. Approx. 20 unramified, cruciform and monopodial-pinnate ectomycorrhizas are fossilized adjacent to rootlets, and different developmental stages of the fossil mycorrhizas are delicately preserved in the ancient resin. Compounds of melanins were detectable in the dark hyphae. The mycobiont, Eomelanomyces cenococcoides gen. et spec. nov., is considered to be an ascomycete; the host is most likely a dipterocarp representative. An early ectomycorrhizal association may have conferred an evolutionary advantage on dipterocarps. Our find indicates that ectomycorrhizas occurred contemporaneously within both gymnosperms (Pinaceae) and angiosperms (Dipterocarpaceae) by the Lower Eocene.

Crown Group Lejeuneaceae and Pleurocarpous Mosses in Early Eocene (Ypresian) Indian Amber

Cambay amber originates from the warmest period of the Eocene, which is also well known for the appearance of early angiosperm-dominated megathermal forests. The humid climate of these forests may have triggered the evolution of epiphytic lineages of bryophytes; however, early Eocene fossils of bryophytes are rare. Here, we present evidence for lejeuneoid liverworts and pleurocarpous mosses in Cambay amber. The preserved morphology of the moss fossil is inconclusive for a detailed taxonomic treatment. The liverwort fossil is, however, distinctive; its zig-zagged stems, suberect complicate-bilobed leaves, large leaf lobules, and small, deeply bifid underleaves suggest a member of Lejeuneaceae subtribe Lejeuneinae (Harpalejeunea, Lejeunea, Microlejeunea). We tested alternative classification possibilities by conducting divergence time estimates based on DNA sequence variation of Lejeuneinae using the age of the fossil for corresponding age constraints. Consideration of the fossil as a stem group member of Microlejeunea or Lejeunea resulted in an Eocene to Late Cretaceous age of the Lejeuneinae crown group. This reconstruction is in good accordance with published divergence time estimates generated without the newly presented fossil evidence. Balancing available evidence, we describe the liverwort fossil as the extinct species Microlejeunea nyiahae, representing the oldest crown group fossil of Lejeuneaceae.

The termites of Early Eocene Cambay amber, with the earliest record of the Termitidae (Isoptera)

Abstract The fauna of termites (Isoptera) preserved in Early Eocene amber from the Cambay Basin (Gujarat, India) are described and figured. Three new genera and four new species are recognized, all of them Neoisoptera – Parastylotermes krishnai Engel & Grimaldi, sp. n. (Stylotermitidae); Prostylotermes kamboja Engel & Grimaldi, gen. et sp. n. (Stylotermitidae?); Zophotermes Engel, gen. n., with Zophotermes ashoki Engel & Singh, sp. n. (Rhinotermitidae: Prorhinotermitinae); and Nanotermes isaacae Engel & Grimaldi, gen. et sp. n. (Termitidae: Termitinae?). Together these species represent the earliest Tertiary records of the Neoisoptera and the oldest definitive record of Termitidae, a family that comprises >75% of the living species of Isoptera. Interestingly, the affinities of the Cambay amber termites are with largely Laurasian lineages, in this regard paralleling relationships seen between the fauna of bees and some flies. Diversity of Neoisoptera in Indian amber may reflect origin of the amber deposit in Dipterocarpaceae forests formed at or near the paleoequator.

Biting Midges (Diptera: Ceratopogonidae) from Cambay Amber Indicate that the Eocene Fauna of the Indian Subcontinent Was Not Isolated

India’s unique and highly diverse biota combined with its unique geodynamical history has generated significant interest in the patterns and processes that have shaped the current distribution of India’s flora and fauna and their biogeographical relationships. Fifty four million year old Cambay amber from northwestern India provides the opportunity to address questions relating to endemism and biogeographic history by studying fossil insects. Within the present study seven extant and three fossil genera of biting midges are recorded from Cambay amber and five new species are described: Eohelea indica Stebner & Szadziewski n. sp., Gedanohelea gerdesorum Stebner & Szadziewski n. sp., Meunierohelea cambayana Stebner & Szadziewski n. sp., Meunierohelea borkenti Stebner & Szadziewski n. sp., and Meunierohelea orientalis Stebner & Szadziewski n. sp. Fossils of species in the genera Leptoconops Skuse, 1889, Forcipomyia Meigen, 1818, Brachypogon Kieffer, 1899, Stilobezzia Kieffer, 1911, Serromyia Meigen, 1818, and Mantohelea Szadziewski, 1988 are recorded without formal description. Furthermore, one fossil belonging to the genus Camptopterohelea Wirth & Hubert, 1960 is included in the present study. Our study reveals faunal links among Ceratopogonidae from Cambay amber and contemporaneous amber from Fushun, China, Eocene Baltic amber from Europe, as well as the modern Australasian and the Oriental regions. These findings imply that faunal exchange between Europe, Asia and India took place before the formation of Cambay amber in the early Eocene.

Exceptional preservation of angiosperm markers in Miocene and Eocene ambers

The traditional organic geochemical view suggests that polar triterpenoids are degraded easily during fossilization of organic matter. For the first time, we show that dammarane-type tetracyclic polar triterpenoids are preserved in the geosphere. Hydroxydammarenone (20-hydroxy-24-dammaren-3-one), a typical constituent of extant Dipterocarpaceae (an angiosperm tree family) derived dammar resin, is preserved in Miocene amber. The oxidative products of hydroxydammarenone (i.e., 20,24-epoxy-25-hydroxydammaran-3-one, and hexakisnor-dammaran-3,20-dione) are found in both Miocene and Eocene ambers. The presence of hydroxydammarenone in Miocene amber clearly depicts that natural biotriterpenoid markers can be preserved in the geosphere for tens of millions of years. This result demonstrates that the dammarane-type oxygenated compounds, which are extensively used in archaeological studies, can also be utilized as diagnostic markers for paleochemosystematics.

The anamorphic genus Monotosporella (Ascomycota) from Eocene amber and from modern Agathis resin.

The anamorphic fungal genus Monotosporella (Ascomycota, Sordariomycetes) has been reco-vered from a piece of Early Eocene Indian amber, as well as from the surface of extant resin flows in New Caledonia. The fossil fungus was obtained from the Tarkeshwar Lignite Mine of Gujarat State, western India, and was part of the biota of an early tropical angiosperm rainforest. The amber inclusion represents the second fossil record of Sordariomycetes, as well as the first fossil of its particular order (either Savoryellales or Chaetosphaeriales). The fossil fungus is distinguished from extant representatives by possessing both short conidiophores and small two-septate pyriform conidia, and is described as Monotosporella doerfeltii sp. nov. Inside the amber, the anamorph is attached to its substrate, which is likely the degraded thallus of a cladoniform lichen. The extant New Caledonian species is assigned to Monotosporella setosa. It was found growing on semi-solidified resin flows of Agathis ovata (Araucariaceae), and is the first record of Monotosporella from modern resin substrates.