Geological and palaeontological research in the Rhynie area, Aberdeenshire, Scotland, has progressed in several stages. Following early surveys in the nineteenth century, Dr William Mackie mapped the western margin of the basin in 1910–1913, and discovered the plant-bearing chert. Following trenching of the chert in 1913, Kidston & Lang described the plant fossils between 1917 and 1921 and Scourfield, Hirst and Maulik the arthropods in the 1920s. Following a ‘dark age’ of some 30 years, Geoffrey Lyon awakened interest in the late 1950s. Trenching in 1963–1971 provided Lyon and his co-workers with new material, and resulted in finds of new plants and reinterpretations of earlier work. The next phase was initiated by Winfried Remy's discovery of gametophytes in material given to him by Lyon. Since 1980, the Münster school has continued to make exciting discoveries. Aberdeen University involvement began in 1987 with geochemical work confirming a hot spring origin for the chert. Drill cores taken in 1988 and 1997, and further trenching have allowed structural, sedimentological and stratigraphic reappraisals, and resulted in the discovery of a new biota in the Windyfield chert. Long-term collaborative international research continues to advance interpretation of this unique Early Devonian hot spring system, and the remarkably diverse freshwater and terrestrial biota of the cherts.

A new geological map of the Early Devonian Rhynie Basin has been produced by traditional methods supplemented by trenching to bedrock and a ground magnetic survey. This shows that the basin margins are mostly fault-controlled and three trends are recognised: NE–SW, NNE–SSW and N–S. Three sets of open folds are distinguished with axial traces trending NE–SW, E–W and roughly NW–SE. The faults defining the basin margin and the folding may be related to basin formation within a regional strike-slip system of Early Devonian age.

The stratigraphic succession comprises three mappable units: a lower mixed unit of sandstones, shales, conglomerates and andesitic lava (>700 m), a middle unit of laminated grey shale and siltstone (>300 m); and an upper unit of laminated sandstones and shales (>300 m). These correlate with the Tillybrachty Sandstone and Quarry Hill Sandstone Formations (lower unit), and the Dryden Flags Formation (middle and upper units).

Small areas containing abundant chert float found outside the Rhynie SSSI may represent the surface expression of chert pods within the middle unit of laminated grey shale and siltstone, which also hosts the Rhynie cherts. The Windyfield cherts occur within the upper unit. No further centres of hydrothermal activity have been found in the northern half of the basin

Hydrothermal alteration at Rhynie, Aberdeenshire, Scotland, is concentrated along a fault zone, which juxtaposes surface deposits and the mineralised feeder zone to the Rhynie hotspring system. Mineralisation consists of breccias and veins filled with quartz, chert, calcite, K-feldspar and pyrite. Associated pervasive alteration comprises a high-temperature K-feldsparquartz-illite facies (formed at 250–350°C), a medium-temperature mixed layered illite/smectitequartz-K-feldspar-chlorite-calcite facies (formed at 150–200°C) and a low-temperature mixed layered illite/smectite-chlorite-calcite facies (formed at 100 to +150°C). The fluids responsible for mineralisation were mainly moderate- to high-temperature (Th =91–360°C), low-salinity (<0·2 to 2·9 wt.% NaCl eq.) H2O-NaCl-heated meteoric fluids comparable to modern and ancient hot-spring systems. The migration of these fluids was mainly restricted to a major fault zone bounding the Devonian basin. Fluids responsible for mineralisation, alteration and cementation elsewhere in the basin were low-temperature (Th 57 to 161°C), low- to high-salinity (<0·2 to 18 wt.% NaCl eq.) H2O-NaCl fluids, which resemble basinal brines.

The Windyfield chert site is located 700 m NE of the original Rhynie chert locality at Rhynie, Aberdeenshire, Scotland. Originally identified by concentrations of surface float material, a drilling and trenching programme of the area in 1997 revealed a chert ‘pod’ in situ interbedded with fluvial/lacustrine sandstones and hydrothermally altered shales. Chert morphologies identified from float blocks and trench material range from tabular beds to lenticular pods displaying massive, nodular, laminated and brecciated fabrics, and geyserite splash textures. A suite of floral and faunal associations, when combined with distinctive macro- and microscopic chert textures, has been used to interpret depositional conditions. Palaeoenvironments ranged from terrestrial laminated, brecciated and vegetated sinter sheets to low-temperature pools and marginal aquatic settings. The flora comprises six higher land plant species, nematophytes, charophytes, various fungi and probable cyanobacteria. Arthropods include branchiopod crustaceans, a euthycarcinoid, trigonotarbid arachnids, centipedes, eoarthropleurids and a possible hexapod. The biota of the Windyfield chert is closely comparable to that found in the Rhynie chert. Together, the Windyfield and Rhynie cherts contain the most diverse associated fossil arthropod fauna of terrestrial and freshwater origin from rocks of comparable age anywhere in the world.

A harvestman (Arachnida: Opiliones) is described from the Early Devonian (Pragian) Rhynie cherts, Aberdeenshire, Scotland. Eophalangium sheari gen. et sp. nov. is the oldest known harvestman. The material includes both males and a female preserving, respectively, a cuticle-lined penis and ovipositor within the opisthosoma. Both these structures are of essentially modern appearance. The Rhynie fossils also show tracheae which are, again, very similar to those of living harvestmen. This is the oldest unequivocal record of arachnid tracheal respiration and indicates that E. sheari was terrestrial. An annulate, setose ovipositor in the female suggests that it can be excluded from the clades Dyspnoi and Laniatores, in which the ovipositor lacks such annulations. However, the penis shows evidence of two muscles, a feature of uncertain polarity seen in modern Troguloidea (Dyspnoi). The presence of median eyes and long legs excludes Cyphophthalmi, and thus, E. sheari is tentatively referred to the suborder Eupnoi. Therefore, this remarkable material is implicitly a crown-group harvestman and is one of the oldest known crown-group chelicerates. It also suggests an extraordinary degree of morphological stasis within the eupnoid line, with the Devonian forms differing little in gross morphology – and perhaps in reproductive behaviour – from their modern counterp

A new aquatic crustacean is described from the Early Devonian Rhynie chert. The small, broadly oviform carapace consists of a univalve symmetrical about its longitudinal axis, formed by a continuous cuticle with no fold or hinge. In transverse cross-section, the ventral surface exhibits a broadly ‘W’-shaped outline. Areas of fine, setose objects are preserved, positioned antero-ventrally on either side of the mid-line axis. In transverse sections, a ring-shaped internal structure is also visible. In longitudinal cross-section, this structure is resolved into an axially positioned, internally subdivided tube occupying the mid- and posterior part of the carapace. A small, anteriorly positioned rostrum has a deep pit on each side, perhaps indicative of the socket of an antennal appendage. The arthropods are commonly found clustered together around plant axes, comprising groups of up to 25 or more similarly sized individuals. The chert texture enclosing many individual specimens indicates a sub-aqueous preservational environment. The same organism has recently been discovered in preparations from the nearby and equivalently aged Windyfield hot spring deposit. The morphology of this new arthropod suggests affinity with Branchiopoda and it is tentatively placed within the Diplostraca (an order which contains both ‘conchostracans’ and ‘cladocerans’, or water fleas). The presence of a univalve test suggests a cladoceran affinity, but the lack of diagnostic appendages in the present material does not allow us to take the classification any further. This is potentially the earliest known occurrence of the group and would extend their fossil record back from the Early Cretaceous. The remarkable similarity between these arthropods and their present-day representatives strongly indicates morphological stasis within the group from early on in their evolutionary history

Fossils in the Pragian Rhynie cherts, Aberdeenshire, Scotland, are preserved with exquisite cellular detail, and provide much information on Early Devonian terrestrial and freshwater ecosystems. The fossils include abundant and diverse coprolites which demonstrate the existence of consumers differing in life-habit and diet. The coprolites are small (0.5–3 mm) and diverse in morphology and content, including groups of amorphous coprolites as well as coprolites with identifiable, particulate content. The present authors define three new ichnogenera to accommodate these coprolites:

• • Lancifaex encompasses elongate coprolites with particulate content, and includes three ichnospecies, distinguished on morphology, L. simplex, L. divisa and L. moniliforma.

• • Rotundafaex encompasses rotund coprolites with particulate content, and includes a single ichnospecies, R. aggregata.

• • Bacillafaex encompasses rod-shaped coprolites with amorphous content, and includes two ichnospecies, distinguished on size, B. constipatus and B. mina.

Thirteen specimens of type and/or figured Hexapoda from the Rhynie chert are housed at The Natural History Museum, London, UK. They are listed along with a complete bibliography of where they have been cited and figured. Two taxa are present: Rhyniella praecursor Hirst & Maulik 1926 (Collembola); and Rhyniognatha hirsti Tillyard 1928 (Insecta)

Brief descriptions and comments on relationships are given for the seven embryophytic sporophytes in the cherts at Rhynie, Aberdeenshire, Scotland. They are Rhynia gwynnevaughanii Kidston & Lang, Aglaophyton major D. S. Edwards, Horneophyton lignieri Barghoorn & Darrah, Asteroxylon mackiei Kidston & Lang, Nothia aphylla Lyon ex Høeg, Trichopherophyton teuchansii Lyon & Edwards and Ventarura lyonii Powell, Edwards & Trewin. The superb preservation of the silica permineralisations produced in the hot spring environment provides remarkable insights into the anatomy of early land plants which are not available from compression fossils and other modes of permineralisation. They include soft tissues, such as those surrounding stomata, rhizoids, apical and lateral meristems, and diversity in conducting cells, with inferences for palaeoecophysiology, including water use efficiency, transport and absorption, and for growth processes and patterns.

Several new gametophytes are described from the Early Devonian Rhynie chert. The new monotypic genus Remyophyton, being the gametophyte of Rhynia, is represented by a dense stand of in situ preserved gametophytes with antheridia- and archegonia-bearing axes. The gametophytes are remarkably small in comparison to those of other Rhynie chert plants. The generic diagnoses of Lyonophyton and Langiophyton are emended to include archegonia- and antheridiabearing axes. All essential stages of the reproductive cycle, i.e. sporophytes, and male and female gametophytes, can now be demonstrated for three of the six land plants from the Rhynie chert, i.e. Rhynia gwynne-vaughanii, Aglaophyton major and Horneophyton lignieri.

Detailed studies have been undertaken on in situ spores of the plant Horneophyton lignieri (Kidston & Lang) Barghoorn & Darrah, 1938 preserved in the Rhynie chert. It is concluded that mature spores are trilete, with proximal radial ribbing, and a distal ornament of coni. These spores are identical to Emphanisporites decoratus Allen, 1965, a widely reported and morphologically distinctive dispersed spore taxon. Emphanisporites decoratus is a common element of dispersed spore assemblages from throughout the stratigraphical sequence of the Rhynie outlier, and it is concluded that the dispersed spore taxon E. decoratus was produced by H. lignieri. The nature (reproductive biology, ecology and distribution) of H. lignieri is interpreted on the basis of evidence from the dispersed spore record from the Rhynie outlier and elsewhere. It is concluded that H. lignieri was a common and widespread plant of the Lower Devonian. It flourished in the unusual hot springs environment of the Rhynie chert because it was preadapted to do so, not because it was a highly specialized and adapted plant. Integrated evidence from the in situ and dispersed spore records permits a detailed consideration of the biology of the plant H. lignieri.

The discovery of new specimens of Palaeonitella cranii (Kidston & Lang) Pia in the Early Devonian Rhynie chert permits the classification of this charophyte alga. This relatively small charophyte is composed of multicellular furcating branchlets with antheridia attached to the parent plant by a pedicel. The non-calcified gyrogonites, which are not seen in life position, are composed of six sinistrally spiralling cells and have six coronula cells arranged in a single layer around an apical pore.

Because the oogonia of P. cranii have six sinistrally spiralling cells and an apical pore, Palaeonitella has been assigned to the Palaeocharaceae. The shape and form of the oogonia is reminiscent of the extant Chareae, but the morphology of the thalli is indicative of the Nitelleae. This suggests that P. cranii, although retaining some characteristics of the Nitelleae, has some of the characteristics of the Chareae, supporting existing molecular studies which place the Nitelleae at the base of the Characeae.

Palaeonitella cranii inhabited the freshwater alkaline pools and streams which formed on sinter aprons created by hot spring activity. Extant charophytes exist in a similar environment (pH 8·66–8·9, temperature 25–30°C in summer) in pools and channels fed by hot springs in Yellowstone National Park, Wyoming, USA.

The exquisite preservation of organisms in the Early Devonian Rhynie chert ecosystem has permitted the documentation of the morphology and life history biology of fungi belonging to several major taxonomic groups (e.g., Chytridiomycota, Ascomycota, Glomeromycota). The Rhynie chert also provides the first unequivocal evidence in the fossil record of fungal interactions that can in turn be compared with those in modern ecosystems. These interactions in the Rhynie chert involve both green algae and macroplants, with examples of saprophytism, parasitism, and mutualism, including the earliest mycorrhizal associations and lichen symbiosis known to date in the fossil record. Especially significant are several types of specific host responses to fungal infection that indicate that these plants had already evolved methods of defence similar and perhaps analogous to those of extant plants. This suggests that mechanisms underlying the establishment and sustenance of associations of fungi with land plants were well in place prior to the Early Devonian. In addition, a more complete understanding of the microbial organisms involved in this complex ecosystem can also provide calibration points for phylogenies based on molecular data analysis. The richness of the microbial community in the Rhynie chert holds tremendous potential for documenting additional fungal groups, which permits speculation about further interactions with abiotic and biotic components of the environment.

Modern, silica-precipitating hot springs, like those found in the Taupo Volcanic Zone (TVZ) on the North Island of New Zealand, are natural laboratories for assessing microbial silicification. Many of the silicified microbes found in the siliceous sinters of these spring systems seem to be life-like replicas of the original microbes. Such preservation reflects the fact that many of the microbes are replaced and encrusted by opal-A before they are destroyed by desiccation and decay. The taxonomic fidelity of these silicified microbes depends on the preservation potential of those features which are needed to identify them. For example, identification of extant cyanobacteria, relies on as many as 37 different features, most of which are not preserved by silicification.

In the hot-spring systems of the TVZ, characterisation of cyanobacteria which have been replaced and encrusted by opal-A is typically restricted to colony morphology, the length, diameter and morphology of the filament, and the presence/absence of septa, branching or a sheath. In many cases, description is limited to a subset of these parameters. Such a limited set of morphological characteristics severely impedes identifications in terms of extant taxa. The physical changes which accompany the stepwise diagenetic progression from opal-A to opal-CT ± opal-C to microcrystalline quartz may lead to further degradation of the silicified microbes and the loss of more taxonomically important features. Clearly, considerable care must be taken when trying to name silicified microorganisms and make palaeoenvironmental inferences.

The Tahunaatara sinter, Taupo Volcanic Zone, New Zealand, is a ∼17–20-kyr-old hot-spring deposit of opal-A mineralogy. It is interbedded with fluvial, lacustrine and volcaniclastic sediments, some silicified by infusing thermal waters. The exposed sinter (∼4 m thick, 90 m long) was truncated at its southern margin by a landslide, which deposited a conglomerate (up to 2 m thick, 56 m long) of sinter blocks and associated strata nearby. Kaolinite-rich cobbles at the base of the conglomerate indicate a change in the thermal regime and its probable trigger: acid steamcondensate produced alteration. Clasts in the landslide are oriented SW, the same direction as flattened plant reeds entombed in sinter, and as intercalated fluvial beds. Thus, thermal waters, stream flow and the landslide all likely followed the same palaeo-valley, which is similar in terrain and stratigraphy to the Devonian Rhynie hydrothermal system. The plant-rich, layered, in situ sinter contains fossilised microbes and rare stromatolites, and was deposited on mid- to distal slopes adjacent to marshes. Ash falls, fluvial activity and ponding occurred during and after the thermal activity. Unsilicified tephric Ohakea loess (∼26–17 kyr BP) and Taupo Tephra (1·86 kyr BP) blanket both sinter and landslide. Today, the deposits form resistant remnants in a topographically inverted landscape.

During experiments conducted within the vent pool of Medusa Geyser, Norris Geyser Basin, Yellowstone National Park, USA, amorphous opaline silica (opal-A) was deposited on/within plant tissues within 30 days of immersion. Initially, deposition created inter/intra-cellular films which lined cell walls plus intercellular colloid suspensions (sols) of opal-A nano/microspheres. By 330 days, opal-A deposition created a robust external and internal matrix that stabilised tissues against collapse and replicated plant structure. Opal-A films increased to micron-order thicknesses and intracellular sols were created. Systematic variation of opal-A fabric between tissues comprising living/dead cells at the time of deposition indicate that cell function, architecture and shape influence fabric development. Heterogeneity of opal-A fabric within adjacent cells of similar structure/function indicates spatially/temporally fluctuating physicochemical conditions and the presence of intraorganic microenvironments. Early deposition of opal-A films suggests a period of low silica supersaturation and slow opal-A deposition. In contrast, intracellular sols suggest high levels of supersaturation, and rapid opal-A deposition. Shell-like microsphere growth suggests cyclic variation of silica supersaturation, and alternations between rapid and slower opal-A deposition. Microsphere growth to the upper limit of colloidal stability and colloidal crystal structures indicate prolonged sol stability, whilst floc-like microsphere networks indicate localised sol instabi