Diameter at breast height
Diameter at breast height, or DBH, is a standard method of expressing the diameter of the trunk or bole of a standing tree. DBH is one of the most common dendrometric measurements. Tree trunks are measured at the height of an adult's breast. However, this is defined differently in different situations; the convention is now 1.3 m above ground level. In many countries the diameter has been measured at 1.3 metres above ground. In some countries, such as Australia, New Zealand, the United States, India and South Africa, breast height diameter has been measured at a height of 1.4 metres at some times, but because of much active research into allometrics that are being applied to trees and forests, the convention of 1.3 m is now more appropriate. 4.5 ft was used. The height can make a substantial difference to the measured diameter. Ornamental trees are measured at 1.5 metres above ground. However, some authors maintain that the term DBH should be abolished because the heights at which the diameter is measured are so variable and because it may influence forestry calculations such as biomass.
Instead Dx was proposed whereby the x denotes the exact height above the floor at which the diameter is measured. For instance D130 denotes a diameter measured along the stem. On sloping ground, the "above ground" reference point is taken as the highest point on the ground touching the trunk, but some use the average between the highest and lowest points of ground. If the DBH point falls on a swelling in the trunk it is customary to measure the girth below the swelling at the point where the diameter is smallest. Other ambiguous settings for determining the exact place where to measure the diameter is given in Dahdouh-Guebas & Koedam. Treetec, a company based in Melbourne, has created a free online calculator to determine the DBH for multi-stemmed trees as per AS 4970-2009 Protection of Trees on Development Sites; the two most common instruments used to measure DBH are a girthing tape and calipers. A girthing tape measures the girth of the tree; the measure assumes the trunk has a circular cross-section and gives a directly converted reading of the diameter.
It is accurate for most plantation trees. Calipers consist of two parallel arms one of, fixed and the other able to slide along a scale. Calipers are held at right-angles to the trunk with the arms on either side of the trunk. Precision can be improved on non-circular stems by averaging two caliper measurements taken at right-angles. Electronic calipers are available enabling accurate measurements to be taken and stored for further analysis. DBH is used in estimating the amount of timber volume in a single tree or stand of trees utilising the allometric correlation between stem diameter, tree height and timber volume, it can be used in the estimation of the age of veteran trees, given that diameter increment is the only "constant non-reversible feature of tree growth". Biltmore stick Crown Forest inventory Site index Site tree Tree crown measurement Brokaw, N. and Thompson, J.. The H for DBH. Forest Ecology and Management 129: 89-91. Hamilton, G. J.. Forest Mensuration Handbook. Forestry Commission Booklet 39.
HMSO, London. Mackie, E. D. and Matthews, R. W.. Forest Mensuration, a handbook for practitioners. HMSO, Edinburgh. ISBN 0-85538-621-5 White, J.. Estimating the Age of Large and Veteran Trees in Britain. Forestry Commission Information Note 12. Surrey
The Carboniferous is a geologic period and system that spans 60 million years from the end of the Devonian Period 358.9 million years ago, to the beginning of the Permian Period, 298.9 Mya. The name Carboniferous means "coal-bearing" and derives from the Latin words carbō and ferō, was coined by geologists William Conybeare and William Phillips in 1822. Based on a study of the British rock succession, it was the first of the modern'system' names to be employed, reflects the fact that many coal beds were formed globally during that time; the Carboniferous is treated in North America as two geological periods, the earlier Mississippian and the Pennsylvanian. Terrestrial animal life was well established by the Carboniferous period. Amphibians were the dominant land vertebrates, of which one branch would evolve into amniotes, the first terrestrial vertebrates. Arthropods were very common, many were much larger than those of today. Vast swaths of forest covered the land, which would be laid down and become the coal beds characteristic of the Carboniferous stratigraphy evident today.
The atmospheric content of oxygen reached its highest levels in geological history during the period, 35% compared with 21% today, allowing terrestrial invertebrates to evolve to great size. The half of the period experienced glaciations, low sea level, mountain building as the continents collided to form Pangaea. A minor marine and terrestrial extinction event, the Carboniferous rainforest collapse, occurred at the end of the period, caused by climate change. In the United States the Carboniferous is broken into Mississippian and Pennsylvanian subperiods; the Mississippian is about twice as long as the Pennsylvanian, but due to the large thickness of coal-bearing deposits with Pennsylvanian ages in Europe and North America, the two subperiods were long thought to have been more or less equal in duration. In Europe the Lower Carboniferous sub-system is known as the Dinantian, comprising the Tournaisian and Visean Series, dated at 362.5-332.9 Ma, the Upper Carboniferous sub-system is known as the Silesian, comprising the Namurian and Stephanian Series, dated at 332.9-298.9 Ma.
The Silesian is contemporaneous with the late Mississippian Serpukhovian plus the Pennsylvanian. In Britain the Dinantian is traditionally known as the Carboniferous Limestone, the Namurian as the Millstone Grit, the Westphalian as the Coal Measures and Pennant Sandstone; the International Commission on Stratigraphy faunal stages from youngest to oldest, together with some of their regional subdivisions, are: A global drop in sea level at the end of the Devonian reversed early in the Carboniferous. There was a drop in south polar temperatures; these conditions had little effect in the deep tropics, where lush swamps to become coal, flourished to within 30 degrees of the northernmost glaciers. Mid-Carboniferous, a drop in sea level precipitated a major marine extinction, one that hit crinoids and ammonites hard; this sea level drop and the associated unconformity in North America separate the Mississippian subperiod from the Pennsylvanian subperiod. This happened about 323 million years ago, at the onset of the Permo-Carboniferous Glaciation.
The Carboniferous was a time of active mountain-building as the supercontinent Pangaea came together. The southern continents remained tied together in the supercontinent Gondwana, which collided with North America–Europe along the present line of eastern North America; this continental collision resulted in the Hercynian orogeny in Europe, the Alleghenian orogeny in North America. In the same time frame, much of present eastern Eurasian plate welded itself to Europe along the line of the Ural Mountains. Most of the Mesozoic supercontinent of Pangea was now assembled, although North China, South China continents were still separated from Laurasia; the Late Carboniferous Pangaea was shaped like an "O." There were two major oceans in the Carboniferous—Panthalassa and Paleo-Tethys, inside the "O" in the Carboniferous Pangaea. Other minor oceans were shrinking and closed - Rheic Ocean, the small, shallow Ural Ocean and Proto-Tethys Ocean. Average global temperatures in the Early Carboniferous Period were high: 20 °C.
However, cooling during the Middle Carboniferous reduced average global temperatures to about 12 °C. Lack of growth rings of fossilized trees suggest a lack of seasons of a tropical climate. Glaciations in Gondwana, triggered by Gondwana's southward movement, continued into the Permian and because of the lack of clear markers and breaks, the deposits of this glacial period are referred to as Permo-Carboniferous in age; the cooling and drying of the climate led to the Carboniferous Rainforest Collapse during the late Carboniferous. Tropical rainforests fragmented and were devastated by climate change. Carboniferous rocks in Europe and eastern North America consist of a repeated sequence of limestone, sandstone and coal beds. In North America, the early Carboniferous is marine
The Pinaceae are trees or shrubs, including many of the well-known conifers of commercial importance such as cedars, hemlocks, larches and spruces. The family is included in the order Pinales known as Coniferales. Pinaceae are supported as monophyletic by their protein-type sieve cell plastids, pattern of proembryogeny, lack of bioflavonoids, they are the largest extant conifer family in species diversity, with between 220 and 250 species in 11 genera, the second-largest in geographical range, found in most of the Northern Hemisphere, with the majority of the species in temperate climates, but ranging from subarctic to tropical. The family forms the dominant component of boreal and montane forests. One species, Pinus merkusii, grows just south of the equator in Southeast Asia. Major centres of diversity are found in the mountains of southwest China, central Japan, California. Members of the family Pinaceae are trees growing from 2 to 100 m tall evergreen, monoecious, with subopposite or whorled branches, spirally arranged, linear leaves.
The embryos of Pinaceae have three to 24 cotyledons. The female cones are large and woody, 2–60 cm long, with numerous spirally arranged scales, two winged seeds on each scale; the male cones are small, 0.5–6.0 cm long, fall soon after pollination. Seed dispersal is by wind, but some species have large seeds with reduced wings, are dispersed by birds. Analysis of Pinaceae cones reveals how selective pressure has shaped the evolution of variable cone size and function throughout the family. Variation in cone size in the family has resulted from the variation of seed dispersal mechanisms available in their environments over time. All Pinaceae with seeds weighing less than 90 mg are adapted for wind dispersal. Pines having seeds larger than 100 mg are more to have benefited from adaptations that promote animal dispersal by birds. Pinaceae that persist in areas where tree squirrels are abundant do not seem to have evolved adaptations for bird dispersal. Boreal conifers have many adaptions for winter.
The narrow conical shape of northern conifers, their downward-drooping limbs help them shed snow, many of them seasonally alter their biochemistry to make them more resistant to freezing, called "hardening". Classification of the subfamilies and genera of Pinaceae has been subject to debate in the past. Pinaceae ecology and history have all been used as the basis for methods of analyses of the family. An 1891 publication divided the family into two subfamilies, using the number and position of resin canals in the primary vascular region of the young taproot as the primary consideration. In a 1910 publication, the family was divided into two tribes based on the occurrence and type of long–short shoot dimorphism. A more recent classification divided the subfamilies and genera based on the consideration of features of ovulate cone anatomy among extant and fossil members of the family. Below is an example of; the 11 genera are grouped into four subfamilies, based on the microscopical anatomy and the morphology of the cones, wood and leaves: Subfamily Pinoideae: cones are biennial triennial, with each year's scale-growth distinct, forming an umbo on each scale, the cone scale base is broad, concealing the seeds from abaxial view, the seed is without resin vesicles, the seed wing holds the seed in a pair of claws, leaves have primary stomatal bands adaxial or on both surfaces.
Subfamily Piceoideae: cones are annual, without a distinct umbo, the cone scale base is broad, concealing the seeds from abaxial view, seed is without resin vesicles, the seed wing holds the seed loosely in a cup, leaves have primary stomatal bands adaxial or on both surfaces. Subfamily Laricoideae: cones are annual, without a distinct umbo, the cone scale base is broad, concealing the seeds from abaxial view, the seed is without resin vesicles, the seed wing holds the seed in a cup, leaves have primary stomatal bands abaxial only. Subfamily Abietoideae: cones are annual, without a distinct umbo, the cone scale base is narrow, with the seeds visible in abaxial view, the seed has resin vesicles, the seed wing holds the seed in a cup, leaves have primary stomatal bands abaxial only. External stresses on plants have the ability to change the structure and composition of forest ecosystems. Common external stress that Pinaceae experience are herbivore and pathogen attack which leads to tree death.
In order to combat these stresses, trees need to evolve defenses against these stresses. Pinaceae have evolved a myriad of mechanical and chemical defenses, or a combination of the two, in order to protect themselves against antagonists. Pinaceae have the ability to up-regulate a combination of constitutive mechanical and chemical strategies to further their defenses. Pinaceae defenses are prevalent in the bark of the trees; this part of the tree contributes a complex defensive boundary against external antagonists. Constitutive and induced defenses are both found in the bark. Constitutive defenses are the first line of defenses used against antagonists and can include sclerified cells, lignified periderm cells, secondary compounds such as phenolics and resins
Tāne Mahuta is a giant kauri tree in the Waipoua Forest of Northland Region, New Zealand. Its age is estimated to be between 1,250 and 2,500 years, it is the largest kauri known to stand today. It is named of birds; the tree is a remnant of the ancient subtropical rainforest that once grew on the North Auckland Peninsula. Other giant kauri are found nearby, notably Te Matua Ngahere. Tāne Mahuta is the most famous tree in New Zealand, along with Te Matua Ngahere, it was discovered and identified in early January 1924 when contractors surveyed the present State Highway 12 route through the forest. In 1928 Nicholas Yakas and other bushmen, who were building the road identified the tree. In April 2009, Tāne Mahuta was partnered with the tree Jōmon Sugi on Japan. During the New Zealand drought of 2013, 10,000 litres of water from a nearby stream was diverted to Tāne Mahuta, showing signs of dehydration. In 2018, the tree was considered threatened by kauri dieback, a fatal disease which has infected nearby kauri trees.
New Zealand's Department of Conservation initiated a plan to protect and save the tree from kauri dieback. The measurements above were taken in 2002 by Dr. Robert Van Pelt, a forest ecology researcher and affiliate assistant professor at the University of Washington. Former measurements taken in 1971 by the New Zealand Forestry Service may be found on The New Zealand Tree Register
The Cambrian Period was the first geological period of the Paleozoic Era, of the Phanerozoic Eon. The Cambrian lasted 55.6 million years from the end of the preceding Ediacaran Period 541 million years ago to the beginning of the Ordovician Period 485.4 mya. Its subdivisions, its base, are somewhat in flux; the period was established by Adam Sedgwick, who named it after Cambria, the Latin name of Wales, where Britain's Cambrian rocks are best exposed. The Cambrian is unique in its unusually high proportion of lagerstätte sedimentary deposits, sites of exceptional preservation where "soft" parts of organisms are preserved as well as their more resistant shells; as a result, our understanding of the Cambrian biology surpasses that of some periods. The Cambrian marked a profound change in life on Earth. Complex, multicellular organisms became more common in the millions of years preceding the Cambrian, but it was not until this period that mineralized—hence fossilized—organisms became common; the rapid diversification of life forms in the Cambrian, known as the Cambrian explosion, produced the first representatives of all modern animal phyla.
Phylogenetic analysis has supported the view that during the Cambrian radiation, metazoa evolved monophyletically from a single common ancestor: flagellated colonial protists similar to modern choanoflagellates. Although diverse life forms prospered in the oceans, the land is thought to have been comparatively barren—with nothing more complex than a microbial soil crust and a few molluscs that emerged to browse on the microbial biofilm. Most of the continents were dry and rocky due to a lack of vegetation. Shallow seas flanked the margins of several continents created during the breakup of the supercontinent Pannotia; the seas were warm, polar ice was absent for much of the period. Despite the long recognition of its distinction from younger Ordovician rocks and older Precambrian rocks, it was not until 1994 that the Cambrian system/period was internationally ratified; the base of the Cambrian lies atop a complex assemblage of trace fossils known as the Treptichnus pedum assemblage. The use of Treptichnus pedum, a reference ichnofossil to mark the lower boundary of the Cambrian, is difficult since the occurrence of similar trace fossils belonging to the Treptichnids group are found well below the T. pedum in Namibia and Newfoundland, in the western USA.
The stratigraphic range of T. pedum overlaps the range of the Ediacaran fossils in Namibia, in Spain. The Cambrian Period was followed by the Ordovician Period; the Cambrian is divided into ten ages. Only three series and six stages are named and have a GSSP; because the international stratigraphic subdivision is not yet complete, many local subdivisions are still used. In some of these subdivisions the Cambrian is divided into three series with locally differing names – the Early Cambrian, Middle Cambrian and Furongian. Rocks of these epochs are referred to as belonging to Upper Cambrian. Trilobite zones allow biostratigraphic correlation in the Cambrian; each of the local series is divided into several stages. The Cambrian is divided into several regional faunal stages of which the Russian-Kazakhian system is most used in international parlance: *Most Russian paleontologists define the lower boundary of the Cambrian at the base of the Tommotian Stage, characterized by diversification and global distribution of organisms with mineral skeletons and the appearance of the first Archaeocyath bioherms.
The International Commission on Stratigraphy list the Cambrian period as beginning at 541 million years ago and ending at 485.4 million years ago. The lower boundary of the Cambrian was held to represent the first appearance of complex life, represented by trilobites; the recognition of small shelly fossils before the first trilobites, Ediacara biota earlier, led to calls for a more defined base to the Cambrian period. After decades of careful consideration, a continuous sedimentary sequence at Fortune Head, Newfoundland was settled upon as a formal base of the Cambrian period, to be correlated worldwide by the earliest appearance of Treptichnus pedum. Discovery of this fossil a few metres below the GSSP led to the refinement of this statement, it is the T. pedum ichnofossil assemblage, now formally used to correlate the base of the Cambrian. This formal designation allowed radiometric dates to be obtained from samples across the globe that corresponded to the base of the Cambrian. Early dates of 570 million years ago gained favour, though the methods used to obtain this number are now considered to be unsuitable and inaccurate.
A more precise date using modern radiometric dating yield a date of 541 ± 0.3 million years ago. The ash horizon in Oman from which this date was recovered corresponds to a marked fall in the abundance of carbon-13 that correlates to equivalent excursions elsewhere in the world, to the disappearance of distinctive Ediacaran fossils. There are arguments that the dated horizon in Oman does not correspond to the Ediacaran-Cambrian boundary, but represents a facies change from marine to evaporite-dominated strata — which w
A leaf is an organ of a vascular plant and is the principal lateral appendage of the stem. The leaves and stem together form the shoot. Leaves are collectively referred to as foliage, as in "autumn foliage". A leaf is a thin, dorsiventrally flattened organ borne above ground and specialized for photosynthesis. In most leaves, the primary photosynthetic tissue, the palisade mesophyll, is located on the upper side of the blade or lamina of the leaf but in some species, including the mature foliage of Eucalyptus, palisade mesophyll is present on both sides and the leaves are said to be isobilateral. Most leaves have distinct upper surface and lower surface that differ in colour, the number of stomata, the amount and structure of epicuticular wax and other features. Leaves can have many different shapes and textures; the broad, flat leaves with complex venation of flowering plants are known as megaphylls and the species that bear them, the majority, as broad-leaved or megaphyllous plants. In the clubmosses, with different evolutionary origins, the leaves are simple and are known as microphylls.
Some leaves, such as bulb scales, are not above ground. In many aquatic species the leaves are submerged in water. Succulent plants have thick juicy leaves, but some leaves are without major photosynthetic function and may be dead at maturity, as in some cataphylls and spines. Furthermore, several kinds of leaf-like structures found in vascular plants are not homologous with them. Examples include flattened plant stems called phylloclades and cladodes, flattened leaf stems called phyllodes which differ from leaves both in their structure and origin; some structures of non-vascular plants function much like leaves. Examples include the phyllids of liverworts. Leaves are the most important organs of most vascular plants. Green plants are autotrophic, meaning that they do not obtain food from other living things but instead create their own food by photosynthesis, they capture the energy in sunlight and use it to make simple sugars, such as glucose and sucrose, from carbon dioxide and water. The sugars are stored as starch, further processed by chemical synthesis into more complex organic molecules such as proteins or cellulose, the basic structural material in plant cell walls, or metabolised by cellular respiration to provide chemical energy to run cellular processes.
The leaves draw water from the ground in the transpiration stream through a vascular conducting system known as xylem and obtain carbon dioxide from the atmosphere by diffusion through openings called stomata in the outer covering layer of the leaf, while leaves are orientated to maximise their exposure to sunlight. Once sugar has been synthesized, it needs to be transported to areas of active growth such as the plant shoots and roots. Vascular plants transport sucrose in a special tissue called the phloem; the phloem and xylem are parallel to each other but the transport of materials is in opposite directions. Within the leaf these vascular systems branch to form veins which supply as much of the leaf as possible, ensuring that cells carrying out photosynthesis are close to the transportation system. Leaves are broad and thin, thereby maximising the surface area directly exposed to light and enabling the light to penetrate the tissues and reach the chloroplasts, thus promoting photosynthesis.
They are arranged on the plant so as to expose their surfaces to light as efficiently as possible without shading each other, but there are many exceptions and complications. For instance plants adapted to windy conditions may have pendent leaves, such as in many willows and eucalyptss; the flat, or laminar, shape maximises thermal contact with the surrounding air, promoting cooling. Functionally, in addition to carrying out photosynthesis, the leaf is the principal site of transpiration, providing the energy required to draw the transpiration stream up from the roots, guttation. Many gymnosperms have thin needle-like or scale-like leaves that can be advantageous in cold climates with frequent snow and frost; these are interpreted as reduced from megaphyllous leaves of their Devonian ancestors. Some leaf forms are adapted to modulate the amount of light they absorb to avoid or mitigate excessive heat, ultraviolet damage, or desiccation, or to sacrifice light-absorption efficiency in favour of protection from herbivory.
For xerophytes the major constraint drought. Some window plants such as Fenestraria species and some Haworthia species such as Haworthia tesselata and Haworthia truncata are examples of xerophytes. and Bulbine mesembryanthemoides. Leaves function to store chemical energy and water and may become specialised organs serving other functions, such as tendrils of peas and other legumes, the protective spines of cacti and the insect traps in carnivorous plants such as Nepenthes and Sarracenia. Leaves are the fundamental structural units from which cones are constructed in gymnosperms and from which flowers are constructed in flowering plants; the internal organisation of most kinds of leaves has evolved to maximise exposure of the photosynthetic organelles, the chloroplasts, to light and to increase the absorption of carbon dioxide while at the same time controlling water loss. Their surfaces are waterproofed by the plant cuticle and gas exchange between the mesophyll cells and the atmosphere is controlled by minute openings called stomata which open or close to regulate the rate exchange of carbon dioxide and water vapour into
The Devonian is a geologic period and system of the Paleozoic, spanning 60 million years from the end of the Silurian, 419.2 million years ago, to the beginning of the Carboniferous, 358.9 Mya. It is named after Devon, where rocks from this period were first studied; the first significant adaptive radiation of life on dry land occurred during the Devonian. Free-sporing vascular plants began to spread across dry land, forming extensive forests which covered the continents. By the middle of the Devonian, several groups of plants had evolved leaves and true roots, by the end of the period the first seed-bearing plants appeared. Various terrestrial arthropods became well-established. Fish reached substantial diversity during this time, leading the Devonian to be dubbed the "Age of Fishes." The first ray-finned and lobe-finned bony fish appeared, while the placoderms began dominating every known aquatic environment. The ancestors of all four-limbed vertebrates began adapting to walking on land, as their strong pectoral and pelvic fins evolved into legs.
In the oceans, primitive sharks became more numerous than in the Late Ordovician. The first ammonites, species of molluscs, appeared. Trilobites, the mollusc-like brachiopods and the great coral reefs, were still common; the Late Devonian extinction which started about 375 million years ago affected marine life, killing off all placodermi, all trilobites, save for a few species of the order Proetida. The palaeogeography was dominated by the supercontinent of Gondwana to the south, the continent of Siberia to the north, the early formation of the small continent of Euramerica in between; the period is named after Devon, a county in southwestern England, where a controversial argument in the 1830s over the age and structure of the rocks found distributed throughout the county was resolved by the definition of the Devonian period in the geological timescale. The Great Devonian Controversy was a long period of vigorous argument and counter-argument between the main protagonists of Roderick Murchison with Adam Sedgwick against Henry De la Beche supported by George Bellas Greenough.
Murchison and Sedgwick named the period they proposed as the Devonian System. While the rock beds that define the start and end of the Devonian period are well identified, the exact dates are uncertain. According to the International Commission on Stratigraphy, the Devonian extends from the end of the Silurian 419.2 Mya, to the beginning of the Carboniferous 358.9 Mya. In nineteenth-century texts the Devonian has been called the "Old Red Age", after the red and brown terrestrial deposits known in the United Kingdom as the Old Red Sandstone in which early fossil discoveries were found. Another common term is "Age of the Fishes", referring to the evolution of several major groups of fish that took place during the period. Older literature on the Anglo-Welsh basin divides it into the Downtonian, Dittonian and Farlovian stages, the latter three of which are placed in the Devonian; the Devonian has erroneously been characterised as a "greenhouse age", due to sampling bias: most of the early Devonian-age discoveries came from the strata of western Europe and eastern North America, which at the time straddled the Equator as part of the supercontinent of Euramerica where fossil signatures of widespread reefs indicate tropical climates that were warm and moderately humid but in fact the climate in the Devonian differed during its epochs and between geographic regions.
For example, during the Early Devonian, arid conditions were prevalent through much of the world including Siberia, North America, China, but Africa and South America had a warm temperate climate. In the Late Devonian, by contrast, arid conditions were less prevalent across the world and temperate climates were more common; the Devonian Period is formally broken into Early and Late subdivisions. The rocks corresponding to those epochs are referred to as belonging to the Lower and Upper parts of the Devonian System. Early DevonianThe Early Devonian lasted from 419.2 ± 2.8 to 393.3 ± 2.5 and began with the Lochkovian stage, which lasted until the Pragian. It spanned from 410.8 ± 2.8 to 407.6 ± 2.5, was followed by the Emsian, which lasted until the Middle Devonian began, 393.3± 2.7 million years ago. During this time, the first ammonoids appeared. Ammonoids during this time period differed little from their nautiloid counterparts; these ammonoids belong to the order Agoniatitida, which in epochs evolved to new ammonoid orders, for example Goniatitida and Clymeniida.
This class of cephalopod molluscs would dominate the marine fauna until the beginning of the Mesozoic era. Middle DevonianThe Middle Devonian comprised two subdivisions: first the Eifelian, which gave way to the Givetian 387.7± 2.7 million years ago. During this time the jawless agnathan fishes began to decline in diversity in freshwater and marine environments due to drastic environmental changes and due to the increasing competition and diversity of jawed fishes; the shallow, oxygen-depleted waters of Devonian inland lakes, surrounded by primitive plants, provided the environment necessary for certain early fish to develop such essential characteristics as well developed lungs, the ability to crawl out of the water and onto the land for short periods of time. Late DevonianFinally, the Late Devonian started with the Frasnian, 382.7 ± 2.8 to 372.2 ± 2.5, during which the first forests took shape on land. The first tetrapods appeared in the fossil record in the ensuing Famennian subdivisi