Galling: the prevalent form of insect folivory in the latest Neogene monsoon-influenced tropical forests of the Chotanagpur Plateau,eastern India

Indian Cenozoic deposits contain well-preserved diverse angiosperm leaf assemblages, but galling, a common form of angiosperm leaf damage in modern tropical forests, has not been well-documented. Here we report insect herbivory on diverse angiosperm fossil leaf specimens from Pliocene (Rajdanda Formation) sediments of the Chotanagpur Plateau, eastern India, revealing that galling was the most common form of folivory in the latest Neogene monsoon-adapted tropical forests of eastern India. Diverse well-preserved galls are described on the basis of their size, shape and position on the host angiosperm leaf remains. Nine gall damage types (DT 32, DT 33, DT 34, DT 80, DT 83, DT 85, DT 110, DT 120, and DT 144) are identified. They are compared with extant galls and the probable gall inducers making morphologically similar galls on related host tropical plant species of Ficus, Mangifera, Albizia, Galactia, Ziziphus, Hylodesmum, Adina, and Psidium. The gall producers belong to the insect orders Orthoptera, Hemiptera, Hymenoptera, Coleoptera, Lepidoptera, and Diptera (Cecidomyiidae). Although the detailed morphology of the phytophagous insects associated with the recovered fossil leaves is unknown, our findings indicate that many modern plant-insect relationships were likely established by the Pliocene. The reconstructed warm, humid Pliocene climate with a weak monsoon seasonality was conducive to extensive galling activity during this time.     

In situ occurrence of a gall midge (Insecta,Diptera, Cecidomyiidae) on fossilized angiosperm leaf cuticle fragments from the Pliocene sediments of eastern India

In situ preservation of fossil insect damage in plant fossils is an excellent tool to study the coevolution of flora and fauna through geological time, but finding both damage and the insect causing that damage in the same specimen is a very rare phenomenon. Galling is a common form of angiosperm leaf damage, which can be regarded as a kind of extended phenotype of the causal insects, essentially the gall midges, but galls usually lack remains of the insects themselves. Here we report the in situ occurrence of a gall midge (Insecta, Diptera, Cecidomyiidae) as well as its pupal exuviae on the abaxial cuticular surface of fossilized leaf cuticle fragments of Fabaceae leaves (cf. Albizia) that also bear galls, recovered from the latest Neogene (Rajdanda Formation, Pliocene) sediments of the Chotonagpur Plateau, Jharkhand, northeastern India. This Pliocene gall midge features well-preserved legs, segmented antenna with distinct and enlarged scape, elongate curved setae, and longer than broad terminal plate of the ovipositor lamellae. The in situ presence of a gall midge on a host fabaceous leaf cuticle indicates the existence of a host-ectoparasite relationship in the ancient warm and humid tropical monsoon-influenced forests of eastern India during the Pliocene. This is the first authentic fossil record of an in situ phytophagous insect of Cecidomyiidae from India, as well as southeast Asia. Although the identification of the recovered phytophagous insect associated with the fossil leaf cuticle is only possible to family level, this find reveals that such plant-insect relationships existed in the Pliocene of eastern India.     

Evidence for diversification of <Emphasis Type="Italic">Calophyllum</Emphasis> L. (Calophyllaceae) in the Neogene Siwalik forests of eastern Himalaya

Here, we report fossil leaves, woods, and pollen grains comparable to Calophyllum L. (mainly to Calophyllum inophyllum L. and Calophyllum polyanthum Wall. ex Choisy) of Calophyllaceae from the upper (Kimin Formation, late Pliocene-early Pleistocene), middle (Subansiri Formation, Pliocene) Siwalik sediments of the Arunachal sub-Himalaya, and lower (Gish Clay Formation of Sevok Group; middle to late Miocene) Siwalik sediments of the Darjeeling foothills, eastern Himalaya. Their presence indicates a warm and humid tropical environment in the region during the period of Siwalik sedimentation. Considering all records of Calophyllum, it is suggested that Calophyllum was a frequent forest element throughout the period of Siwalik sedimentation during the Neogene (Miocene time). At present, C. polyanthum grows in the eastern Himalaya, but C. inophyllum is totally absent from north-eastern regions suggesting differential adaptability of these taxa to changing ecoclimatic conditions. Distinct climate change in the area, possibly related to the Himalayan Orogeny during Miocene–Pleistocene times, might have caused the disappearance of C. inophyllum from the entire eastern Himalaya and north-east Indian plains and a move to littoral/coastal and swampy forests of India and other adjoining south-east Asian regions, Polynesia, and the east coast of Africa. The past global distribution of Calophyllum is also discussed, and it is suggested that India may have been its primary centre of origin. This is the first time Cenozoic fossil leaves Calophyllum siwalikum Khan, R.A.Spicer & Bera, sp. nov. comparable to C. inophyllum are described using the both macro- and micromorphological characters.     

First fossil evidence of Palaeocarya (Engelhardioideae: Juglandaceae) from India and its biogeographical implications

Even though presently indigenous to eastern Himalaya in India, no Engelhardioideae have been reported from the Cenozoic sediments of India till date. Here, we report the first Indian occurrence of a characteristic engelhardioid winged samaroid fruit having a tri-lobed wing (oblong-ovate median lobe and two lateral lobes) and a globose nut from the latest Neogene (Pliocene: Rajdanda Formation) sediments of Chotanagpur Plateau, eastern India. This is the first fossil evidence of relict family Juglandaceae from the Indian Cenozoic. We determine its taxonomic position on the basis of detailed macromorphological comparison with similar extant and fossil specimens and discuss its palaeoclimatic significance in terms of the present-day distribution of modern analogous species. We assign this Pliocene winged fruit specimen to the morphogenus Palaeocarya sect. Monocosta Manchester and describe it as a new species, namely Palaeocarya indica Hazra, Hazra M & Khan sp. nov. Palaeocarya sect. Monocosta has rich fossil records from the Cenozoic sediments of Europe, North America, and eastern Asia (China, Korea), but the modern analog, Engelhardia, is presently native only to India and neighboring Southeast Asia. We discuss the possible causes of disappearance of Engelhardia from the present-day vegetation of Chotanagpur Plateau. Its disappearance may be related to the gradual intensification of monsoonal rainfall seasonality since the Pliocene. Here, we also review in detail the biogeographic history of Palaeocarya sect. Monocosta and suggest its possible migration routes.     

Evidence of fungal decay in petrified legume wood from the Neogene of the Bengal Basin,India

Silicified fossil legume woods of Cynometroxylon Chowdhury & Ghosh collected from the Neogene (late Miocene) sediments of the Bengal Basin, eastern India, exhibit fungal decay seldom found in the fossil record. The wood possesses numerous perforate areas on the surface that seem to be the result of extensive fungal activity. In transverse section, the decayed areas (pockets) appear irregular to ellipsoidal in outline; in longitudinal section these areas of disrupted tissue are somewhat spindle-shaped. Individual pockets are randomly scattered throughout the secondary xylem or are restricted to a narrow zone. The aforesaid patterns of decay in fossil wood show similarities with that of white rot decay commonly produced by higher fungi, specifically basidiomycetes and ascomycetes. The host fossil wood harbors abundant ramifying and septate fungal hyphae with knob like swellings similar to pseudoclamps in basidiomycetes, and three-celled conidia-like reproductive structures. This record expands our current knowledge of wood decaying fungi-host plant interaction in the Neogene tropical forests of Peninsular India.     

A novel fossil-species of Meliolinites Selkirk (fossil Meliolaceae) and its life cycle stages associated with an angiosperm fossil leaf from the Siwalik (Mio-Pliocene) of Bhutan sub-Himalaya

Here, we report the in-situ occurrence of a new fossil-species of Meliolinites (fossil Meliolaceae), Meliolinites bhutanensis sp. nov. on the cuticle fragments of a compressed angiosperm dicot leaf recovered from the middle Siwalik (Formation II: latest Miocene to Pliocene) of Bhutan, eastern Himalaya. This unique foliicolous new fossil fungal species features well-preserved mycelia consisting of superficial, brown to dark brown, septate, thick-walled, branching hyphae with bi-cellular appressoria, unicellular phialides, and a characteristic long, slightly curved hyphal seta. The web-like, brown to dark brown fungal colonies also include globose to sub-globose, dark brown ascomata, and oblong to broadly cylindrical, 5-celled, 4-septate, brown to dark brown, mature ascospores. As almost all features of different stages in the life cycle (ascospores, mature germinating ascospores, superficial lateral hyphae, hyphal seta, hyphopodia, mycelial colony, and ascomata) of this new fossil-species are found, we have proposed the first time a possible life cycle of fossil-species of Meliolaceae. The in-situ evidence of M. bhutanensis on the host leaf cuticle indicates the possible existence of a host-ectoparasite relationship in Bhutan sub-Himalaya's ancient warm and humid tropical evergreen forest during the deposition. So, M. bhutanensis might have thrived generally under warm and humid climate conditions for its growth and development in the Mio-Pliocene time, which is in conformity with our recently published quantitative climatic data by CLAMP (Climate Leaf Analysis Multivariate Program) analysis.     

Phytogeographical Implication of Bridelia Will. (Phyllanthaceae) Fossil Leaf from the Late Oligocene of India

Background

The family Phyllanthaceae has a predominantly pantropical distribution. Of its several genera, Bridelia Willd. is of a special interest because it has disjunct equally distributed species in Africa and tropical Asia i.e. 18–20 species in Africa-Madagascar (all endemic) and 18 species in tropical Asia (some shared with Australia). On the basis of molecular phylogenetic study on Bridelia, it has been suggested that the genus evolved in Southeast Asia around 33±5 Ma, while speciation and migration to other parts of the world occurred at 10±2 Ma. Fossil records of Bridelia are equally important to support the molecular phylogenetic studies and plate tectonic models.

Results

We describe a new fossil leaf of Bridelia from the late Oligocene (Chattian, 28.4–23 Ma) sediments of Assam, India. The detailed venation pattern of the fossil suggests its affinities with the extant B. ovata, B. retusa and B. stipularis. Based on the present fossil evidence and the known fossil records of Bridelia from the Tertiary sediments of Nepal and India, we infer that the genus evolved in India during the late Oligocene (Chattian, 28.4–23 Ma) and speciation occurred during the Miocene. The stem lineage of the genus migrated to Africa via “Iranian route” and again speciosed in Africa-Madagascar during the late Neogene resulting in the emergence of African endemic clades. Similarly, the genus also migrated to Southeast Asia via Myanmar after the complete suturing of Indian and Eurasian plates. The emergence and speciation of the genus in Asia and Africa is the result of climate change during the Cenozoic.

Conclusions

On the basis of present and known fossil records of Bridelia, we have concluded that the genus evolved during the late Oligocene in northeast India. During the Neogene, the genus diversified and migrated to Southeast Asia via Myanmar and Africa via “Iranian Route”.     

A fossil fruit wing of Dipterocarpus from the middle Miocene of Fujian, China and its palaeoclimatic significance

Dipterocarpus zhengae sp. nov. is described from the middle Miocene Fotan Group of Zhangpu county, Fujian Province, Southeast China on the basis of a fruit wing. Three kinds of venation of the calyx longer lobes (enlarged sepals) occur in the winged fruits of extant Dipterocarpus. The fossil species is referred to the kind having three primary veins and within that category is most similar to extant Dipterocarpus gracilis Blume in the size of the longer lobe as well as its venation. The occurrence of the fruit wing of Dipterocarpus, together with palynological evidence, indicates unequivocally that a tropical climate and tropical rain forest occurred in Zhangpu during the middle Miocene, warmer and more humid than at the present day. The palaeobiogeography of Dipterocarpus is reviewed.     

Two new fossil woods from the early Miocene of Kutch,Gujarat, India and their significance

The Kutch region of western India (Gujarat State) is today arid to semiarid and characterised by mostly ephemeral streams which carry water during the monsoon. The uneven distribution of rainfall and disturbed topography are the result of climate change during the Cenozoic period. Two fossil woods, namely Bauhinium palaeomalabaricum Prakash and Prasad (Fabaceae) and Ebenoxylon indicum Ghosh and Kazmi (Ebenaceae), are described from Kutch in order to provide insights into the palaeovegetation and palaeoclimate. Because the modern representatives of the present and previously described taxa from the same horizon are thermophilic in nature and grow in evergreen to deciduous forests, a warm and humid climate is interpreted. Furthermore, the finding of some mangrove taxa in the assemblage indicates the lagoonal to intertidal environment at the time of deposition.     

Liverwort fossils from the Late Triassic of Baiyin City,Gansu Province,and their geological significance

Although liverworts are widely distributed around the world with a large number of extant species, reliable fossil records are relatively rare. Here, we report a new species, Ricciopsis baojishanensis Han and Yan, n. sp. (Ricciaceae) and an unnamed species, Hepaticites sp. from the Late Triassic Nanying’er Formation in Baojishan Basin, Baiyin City, Gansu Province, Northwest China. The generic designation is based on detailed comparison of the gross morphology with related fossil and extant species. The new species is characterized by its rosette-forming thallus, dichotomous branching, ribbon-like segments and entire margins. The current fossils represent the first record of liverwort from the Late Triassic in Baojishan Basin, Gansu Province. Based on the different fossil records of the Ricciaceae, we suggest that these taxa were widely distributed during Late Triassic to Oligocene worldwide, mainly in warm temperate and tropical environments, similar with their current distribution. The discovery of the present fossils indicates that the climate of Baojishan Basin in Late Triassic is warmer and more humid than that of today.     

A unique record of Cercis from the late early Miocene of interior Asia and its significance for paleoenvironments and paleophytogeography

The climatic impacts of the Tibetan Plateau since the Neogene and the phytogeographic pattern changes of formerly widely-distributed forest communities on the plateau remain poorly constrained. Today, Cercis L. (Fabaceae) is a well-known arborescent genus typically distributed in subtropical to warm temperate zones of the Northern Hemisphere, and Paleogene fossil occurrences from Eurasia and North America show a long history of the genus in mid-low latitudes of the Northern Hemisphere. Here, we describe a fossil species, Cercis zekuensis sp. nov. based on well-preserved fruits from the early Miocene of the northeastern Tibetan Plateau. Detailed morphological comparison (e.g., ventral margin with a veinless wing) of extant and fossil members of Cercis and other genera confirmed validity of the present taxonomic identity. Based on the comparison with extant relatives and their climate preferences, this unexpected occurrence of thermophilic Cercis in northeastern Tibetan Plateau indicates this area had higher temperature and precipitation in the Miocene than today. Integrated with inferred (paleo-)temperature lapse rates, this indicates a low paleoelevation of less than 2.4 km. In contrast with the present-day alpine climate here (~3.7 km), such a low elevation facilitated a more favorable habitat with comparatively high biodiversity and warm temperate forests at that time, as were evidenced by co-occurring megafossils. Moreover, the present existence of Cercis implies the genus was widespread in interior Asia during the early Neogene and shows its modern disjunction or diversification between eastern and central Asia was possibly shaped by the late Cenozoic regional tectonic uplift and consequential environmental deterioration.     

Paleoequatorial rain forest of western India during the EECO: evidence from Uvaria L. fossil and its geological distribution pattern

During the early Eocene, Rajasthan was positioned near the equator and had a warm and humid tropical climate dominated by tropical rainforests like the present-day equatorial forests of South India. Many of the plants retrieved as fossils from Rajasthan are growing there as refugee. This study further strengthens this view as it reports a new species of Uvaria L. from the early Eocene sediments of Bikaner (Rajasthan) showing its best resemblance with the extant U. zeylanica Deless. ex DC., which is presently growing in the evergreen forests of South India and Sri Lanka. The genus is thought to have originated in Africa, and the present finding gives an idea about its geologic distribution in Asia and Australasia via India relying on ‘stepping stone’ hypothesis during the Early Eocene Climatic Optimum (EECO) when climatic conditions were favourable for the luxuriant growth of tropical vegetation. A general cooling trend after EECO and change in the configuration of land and sea affected the climate on the regional scale causing total devastation of tropical evergreen forests that existed in western India during the depositional time; this change is ultimately responsible for creating dry and desertic conditions prevailing in the area at present.     

Diversity of Menispermaceae from the Paleocene and Eocene of South China

We present here the earliest known Asian fossil records of the Menispermaceae based on fossil fruits from Paleocene and Eocene localities in South China. A new genus and species, Paleoorbicarpum parvum sp. nov., and two new species of Stephania Loureiro, S. ornamenta sp. nov. and S. geniculata sp. nov., are recognized from Paleocene deposits of the Sanshui Basin, Guangdong, and a new occurrence of the widespread Eocene species Stephania auriformis (Hollick) Han & Manchester is recognized from the Maoming Basin, Guangdong. The Paleocene Stephania specimens described here represent the earliest fossil endocarp record of the Menispermaceae in eastern Asia. This discovery shows that the moonseed family had arrived in tropical and humid South China by at least the middle Paleocene, which provides important evidence for the origin and phytogeographic history of the family.     

Warm and humid Trans-Himalaya during the late Miocene: plant fossil evidence

Neogene fossil records from the Indus Basin sedimentary rocks (IBSR), deposited in the Indus Tsangpo Suture Zone (ITSZ), are very rare, but are important to understand the history of plant diversity and paleoclimate in the Himalaya. We report fossil wood ascribed to Ebenoxylon siwalicus Prakash from late Miocene sediments of the Karit Formation belonging to ITSZ. The anatomical details of the fossil wood, such as small to medium-sized vessels occluded with tyloses, scanty paratracheal to diffuse-in-aggregate axial parenchyma, 1–3 seriate homo to heterocellular rays, bordered intervessel pits with lenticular apertures and simple perforations, suggest its close affinity with Diospyros Linnaeus of the family Ebenaceae. Further anatomical details suggest a close resemblance with extant D. ehretioides Don and D. macrophylla Blume. The present fossil, along with previously known fossil records of Lagerstroemia (Lythraceae) and palms, indicate that the Trans-Himalaya was warm and humid during the late Miocene, quite different from the modern cool and dry climate in the study area.     

Santonian Angiosperm-dominated leaf-assemblage from Piolenc (Vaucluse,Sud-Est de la France)

Lower Santonian sandy to coaly-clayey beds, collected near the village of Piolenc (Vaucluse) contain amber bearing insects and extremely well-preserved leaf cuticle compressions of Corystospermales, Cycadales, Bennettitales, Gymnosperms and mostly Angiosperms. The sedimentological and taphonomic studies indicate that the clays bearing the fossil cuticles were deposited along the Alpine Sea shore into supratidal ponds or marshes around which the plants studied herein grew. Moreover, the ‘nearest living relative’ and ‘foliar physiognomy’ methods, and the sedimentological and palaeontological studies agree with the occurrence of a warm and humid, tropical climate, including a dry season. To cite this article: B. Gomez et al., C. R. Palevol 2 (2003).     

Two Eocene fossil fruits from the Changchang Basin of Hainan Island,China

Two types of fossil fruit, one belonging to Palaeocarya sp. (Juglandaceae) and the other to Acer cf. A. miofranchetii Hu et Chaney (Aceraceae), are found in the Eocene coal-bearing series from the Changchang Basin of Hainan Island, China. This is the first fossil record of Palaeocarya and Acer in a tropical area of China. These fossils provide evidence for an investigation of the phytogeographic history of these two genera. Since their extant relative genera are distributed mostly in northern temperate or tropical–subtropical mountainous regions, I propose that the Changchang Basin of Hainan Island was close to a mountainous region in the Eocene; the plants bearing these fruits were growing at a mid-high altitude with a relatively cool climate, and the fruits were not preserved in situ but transported to the fossil site. The characters of other associated fossil plants and palynological data also support this hypothesis.     

The diversity and paleoenvironmental significance of Calophyllum (Clusiaceae) from the Miocene of southeastern China

Abstract

Three species within the genus Calophyllum collected from middle Miocene Fotan Group sediments in Zhangpu County, Fujian, southeastern China are described in this paper. These fossils include Calophyllum zhangpuensis sp. nov., Calophyllum striatum, and Calophyllum suraikholaensis. The new fossil species C. zhangpuensis sp. nov. is oval, possesses entire leaves with closely spaced parallel secondary veins and has a round, or slightly retuse, apex. These specimens represent the first known fossil records of this relative wide leaf-type form of Calophyllum from China and have a length:width (L:W) ratio less than 3:1. In combination with the known modern geographic distribution and habitats of this wide leaf-type Calophyllum and other plants, data suggest that the middle Miocene Fotan flora is indicative of a warm climate. Thus, based on available fossil data, we speculate that this genus probably originated in India during the Paleocene before spreading from India to Bangladesh and into China, Sumatra, Malaysia, Indonesia, and Java during the Neogene, leading to its modern distribution. At least, the 3 fossil species in this region can explain floristic exchange between India, Fujian, and South China, which is consistent with previous studies; the occurrence of these 3 species indicates that Calophyllum began to diversity in China no later than the Miocene.     

Coryphoid Palm Leaf Fossils from the Maastrichtian–Danian of Central India with Remarks on Phytogeography of the Coryphoideae (Arecaceae)

Premise of research

A large number of fossil coryphoid palm wood and fruits have been reported from the Deccan Intertrappean beds of India. We document the oldest well-preserved and very rare costapalmate palm leaves and inflorescence like structures from the same horizon.

Methodology

A number of specimens were collected from Maastrichtian–Danian sediments of the Deccan Intertrappean beds, Ghughua, near Umaria, Dindori District, Madhya Pradesh, India. The specimens are compared with modern and fossil taxa of the family Arecaceae.

Pivotal results

Sabalites dindoriensis sp. nov. is described based on fossil leaf specimens including basal to apical parts. These are the oldest coryphoid fossil palm leaves from India as well as, at the time of deposition, from the Gondwana- derived continents.

Conclusions

The fossil record of coryphoid palm leaves presented here and reported from the Eurasian localities suggests that this is the oldest record of coryphoid palm leaves from India and also from the Gondwana- derived continents suggesting that the coryphoid palms were well established and wide spread on both northern and southern hemispheres by the Maastrichtian–Danian. The coryphoid palms probably dispersed into India from Europe via Africa during the latest Cretaceous long before the Indian Plate collided with the Eurasian Plate.     

Occurrence of Phoma Sacc. in the phyllosphere of Neogene Siwalik forest of Arunachal sub-Himalaya and its palaeoecological implications

The present study reports in situ occurrence of two new epiphyllous fungal species of Phomites (comparable to modern genus Phoma Sacc.) on angiospermic leaf remains recovered from the Siwalik sediments (middle Miocene to early Pleistocene) of Arunachal Pradesh, eastern Himalaya. We describe two new species i.e. Phomites siwalicus Vishnu, Khan et Bera S, sp. nov. and Phomites neogenicus Vishnu, Khan et Bera S, sp. nov. on the basis of structural details of pycnidia. The pycnidium is a globose or slightly lens-shaped, ostiolate with a collar layer consisting of thick walled cells, sunken in leaf cuticle, with one-celled conidiospores and short-ampulliform conidiogenous cells. Host leaves resemble to those of extant Dipterocarpus C. F. Gaertn., Shorea Roxb. ex C. F. Gaertn. (Dipterocarpaceae), Dysoxylum Blume (Meliaceae), and Poaceae Barnhart. In situ occurrence of two Phomites morphotypes on the said leaf remains suggests a possible host–parasite interaction in the moist evergreen forest of Arunachal sub-Himalaya during Mio-Pleistocene period. The occurrence of Phomites in appreciable numbers indicates a humid climate favored by high rate of precipitation during Siwalik sedimentation, which is also consistent with our previously published climatic data obtained from the study of the macroscopic plant remains.     

Gleditisioxylon riojana nov. sp. en el Neógeno de Argentina: sus relaciones paleoambientales y biogeográficas

A new fossil species of Leguminosae is described from Neogene of northwest of Argentina. The good preservation of tissues and the diagnostic characters present in this fossil wood let assigning it to a new species of Gleditisioxylon Müller-Stoll and Madel. Gleditisioxylon riojana nov. sp. (Caesalpinoideae) has diagnostic features such as: growth rings distinct, semi-annular porosity, vestured pits, helical thickenings, simple plates, paratracheal parenchyma and rays 1-6 seriate. The possible climatic conditions of Toro Negro Formation were inferred by the use of xylological characters presents in this fossil wood. The presence of Gleditsioxylon added to other data, suggest a new hypothesis to explain the disjunction of Gleditsia L. genus and the occurrence of a single extant species in tropical and subtropical South America.