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63-506: The formation underlies El Capitan in Guadalupe Mountains National Park , and the formation and its associated basin , shelf margin, and backreef formations have been described as "the largest, best-preserved, most accessible, and most intensively studied Paleozoic reef complex in the world." The Guadalupe Mountains were first described in the reports of 1849 and 1850 United States military expeditions to

126-596: A crustose stage; some later become frondose . As sessile encrusting organisms, the corallines are prone to overgrowth by other "fouling" algae. The group have many defences to such immuration, most of which depend on waves disturbing their thalli. However, the most relied-upon method involves waiting for herbivores to devour the potential encrusters. This places them in the unusual position of requiring herbivory, rather than benefiting from its avoidance. Many species periodically slough their surface epithallus – and anything attached to it. Some corallines slough off

189-537: A head in December 1877, when Charles Howard, who had attempted to claim the salt flats, and two colleagues were murdered by an angry mob. This quickly led to widespread violence against local Mexican-American families, leading many to flee south of the Rio Grande. By the time the dust had settled, the salt flats had been claimed and local residents were forced to pay for the salt that for centuries had been free. By

252-451: A living organism was probably Corallina in the 1st century AD. In 1837, Rodolfo Amando Philippi recognized coralline algae were not animals, and he proposed the two generic names Lithophyllum and Lithothamnion as Lithothamnium . For many years, they were included in the order Cryptonemiales as the family Corallinaceae until, in 1986, they were raised to the order Corallinales. Many corallines produce chemicals which promote

315-544: A millimetre to several centimetres high. Some are free-living as rhodoliths (rounded, free-living specimens). The morphological complexity of rhodoliths enhances species diversity, and can be used as a non-taxonomic descriptor for monitoring. Thalli can be divided into three layers: the hypothallus , perithallus and epithallus . The epithallus is periodically shed, either in sheets or piecemeal. Corallines live in varying depths of water, ranging from periodically exposed intertidal settings to 270 m water depth (around

378-543: A prosperous oilman and early conservationist whose Guadalupe Mountains Ranch eventually included 75,000 acres of land in the Guadalupes, including parts of McKittrick Canyon. An early advocate for establishing a park to protect the southern Guadalupes, Hunter campaigned for several years until his death in 1945. Inheriting his father’s desire to see the area protected, both for its scenic beauty and geological significance, Hunter's son, J.C. Hunter Jr., worked with officials of

441-421: A spike in coralline diversity, and the extinction of many delicately branched (and thus predation-prone) forms. The group's internal taxonomy is in a state of flux; molecular studies are proving more reliable than morphological methods in approximating relationships within the group. Recent advances in morphological classification based on skeletal ultrastructure, however, are promising. Crystal morphology within

504-520: A surface layer of epithallial cells, which in a few cases may be an antifouling mechanism which serves the same function as enhancing herbivore recruitment. This also affects the community, as many algae recruit on the surface of a sloughing coralline, and are then lost with the surface layer of cells. This can also generate patchiness within the community. The common Indo-Pacific corallines, Neogoniolithon fosliei and Sporolithon ptychoides , slough epithallial cells in continuous sheets which often lie on

567-681: A taxonomic grouping: Geniculate corallines are branching, tree-like organisms which are attached to the substratum by crustose or calcified, root-like holdfasts. The organisms are made flexible by having noncalcified sections (genicula) separating longer calcified sections (intergenicula). Nongeniculate corallines range from a few micrometres to several centimetres thick crusts. They are often very slow growing, and may occur on rock, coral skeletons, shells, other algae or seagrasses. Crusts may be thin and leafy to thick and strongly adherent. Some are parasitic or partly endophytic on other corallines. Many coralline crusts produce knobby protuberances ranging from

630-556: Is a giant fossil reef, extending at least from the Carlsbad area to the Glass Mountains of Texas. At its greatest development, the reef may have been built up to 200–300 feet (61–91 m) above the sea floor. Richardson (1904) found that the upper and lower beds of the formation were relatively unfossiliferous, but the middle section contained an abundant fossil assemblage unlike any other known at that time. King found that

693-455: Is energetically costly, does not affect seaweed recruitment when herbivores are removed. The surface of these plants is usually kept clean by herbivores, particularly the pear limpet, Patella cochlear . Sloughing in this case is probably a means of eliminating old reproductive structures and grazer-damaged surface cells, and reducing the likelihood of surface penetration by burrowing organisms. The corallines have an excellent fossil record from

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756-740: Is overlain by the Artesia Group , while on the basin side, the Capitan rests on the Delaware Mountain Group and is overlain by the Castile Formation . The formation thus forms a narrow belt curving around the western side of the Delaware Basin that interfingers with backreef formations on the northwestern to southwestern side and with basin formations on the southeastern to northeastern side. The formation

819-545: Is part of the Guadalupe Mountains, an exposed portion of a Permian period reef uplifted and exposed by tectonic activity during the late Cretaceous period. The southern terminus of the Guadalupe Mountains, El Capitan looms over U.S. 62/180, where its imposing height and stark outline have made it one of the iconic images of the Trans-Pecos to generations of travelers. El Capitan is the southernmost peak of

882-443: Is probably an important factor affecting the distribution and grazing effects of herbivores within marine communities. Nothing is known about the microhabitat role of Indo-Pacific corallines. However, the most common species in the region, Hydrolithon onkodes , often forms an intimate relationship with the chiton Cryptoplax larvaeformis . The chiton lives in burrows it makes in H. onkodes plants, and comes out at night to graze on

945-590: The Solenoporaceae , a view that has been disputed. Their fossil record matches their molecular history, and is complete and continuous. The Sporolithaceae tend to be more diverse in periods of high ocean temperatures; the opposite is true for the Corallinaceae . The group's diversity has closely tracked the efficiency of grazing herbivores; for instance, the Eocene appearance of parrotfish marked

1008-477: The 18th century. This is particularly significant in Britain and France , where more than 300,000 tonnes of Phymatolithon calcareum ( Pallas , Adey & McKinnin) and Lithothamnion corallioides are dredged annually. The earliest use of corallines in medicine involved the preparation of a vermifuge from ground geniculate corallines of the genera Corallina and Jania . This use stopped towards

1071-600: The Castile Formation. As salt concentrations increased, laminated halite, anhydrite, sylvite, and polyhalite formed the Salado Formation, which eventually covered and grew beyond the lower Castile Formation. By the end of the Ochoan, these deposits had filled the roughly 1,800-ft-deep basin and covered the reef with dry land. Red silt and sand deposited by rivers crossing these new lands eventually formed

1134-546: The Early Cretaceous onwards, consistent with molecular clocks that show the divergence of the modern taxa beginning in this period. The fossil record of nonarticulated forms is better: the unmineralized genuiculae of articulated forms break down quickly, scattering the mineralized portions, which then decay more quickly. This said, non-mineralizing coralline algae are known from the Silurian of Gotland showing that

1197-657: The Guadalupe escarpment , an ancient limestone reef that forms the present-day Guadalupe Mountains . These mountains are an exposed portion of the Capitan Reef Barrier, a 350-mile long reef constructed primarily from calcareous sponges, encrusting algae, such as stromatolites, and lime-rich mud directly from the ocean. This reef surrounded much of the Delaware Sea , an inland ocean that covered parts of modern southern New Mexico and Trans-Pecos Texas in

1260-412: The Guadalupe Mountains area of the reef, the impact of this tectonic event can be clearly seen in the difference in height between the towering El Capitan and the adjacent salt flat graben, vertically driven 1,000 feet apart by the uplift. Once exposed, the natural forces of wind and rain slowly stripped away the softer sediments, further uncovering the ancient reef and revealing the sheer limestone walls of

1323-607: The Guadalupe Mountains. Deep in the Chihuahuan Desert, the area around El Capitan is composed of rugged mountains and wind-swept salt flats. As a result, most of the Native American presence in the vicinity was centered around 3 miles southeast of El Capitan in the area now called Pine Springs. An oasis of springs, seeps, and vegetation, this area exhibits evidence (mescal dumps, petroglyphs, artifacts, etc.) of human occupation for several centuries. Most recently,

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1386-513: The Mediterranean. Their ability to calcify in low light conditions makes them the some of deepest photosynthetic multicellular organisms in the ocean, having been found as deep as 268 metres (879 ft), and as such a critical base of mesophotic ecological systems. Since coralline algae contain calcium carbonate, they fossilize fairly well. They are particularly significant as stratigraphic markers in petroleum geology. Coralline rock

1449-640: The National Park Service to construct a deal that would enable the Park Service to purchase the land for use as a national park. Forced to run a tortuous gauntlet of state and federal legislators, park officials, and road engineers, as well as property owners and local ranchers, the Hunter donation, which included El Capitan and Guadalupe Peak, was finally deeded over to the Park Service in 1969. Three years later, Guadalupe Mountains National Park

1512-515: The Pecos, hoping to find a potential wagon route with enough timber and water to make consistent travel possible. One of the expeditions, led by Indian Agent Robert S. Neighbors, made its way to El Paso in 1849 through a difficult journey that required crossing and recrossing the Pecos and Devil's Rivers, and then surviving the long, dry stretch from the Davis Mountains to El Paso. Dismayed at

1575-469: The Permian period (about 290 million years ago). Near the end of the late Permian period, in the Ochoan epoch, the outlet that allowed sea water to enter the inland waters began to silt over, occasionally closing the inland sea from its source. Mineral-rich and cut off from replenishment, the inland sea began to evaporate into layers of alternating gray anhydrite/gypsum, brown calcite, and halite, which formed

1638-661: The Tessey, Gilliam, and Vidrio Limestones of the Glass Mountains of west Texas were correlative with the Capitan Formation, and redesigned them as members of the formation. However, by 1937, King had concluded that the Tessey Limestone was not part of the Capital Formation and removed it as a member. By 1942 he had restricted the definition of the Capitan Formation to reef limestone, consistent with

1701-656: The Vidrio Limestone Member had been highly dolomitized, destroying most of its fossil contents, but he recognized fossils of coralline algae , cup corals , crinoid stems, fusulinids, echinoid spines, and brachiopods. El Capitan (Texas) El Capitan ( Spanish : El Capitán ) is a peak in Culberson County , Texas , located within Guadalupe Mountains National Park . The 10th-highest peak in Texas at 8,085 ft (2,464 m), El Capitan

1764-480: The accepted framework for the stratigraphy of the area, and identified the Capitan Formation as late Permian in age. The Capitan Formation itself was first named by G.B. Richardson in 1904 for exposures in the Guadalupe Mountains . Richardson was impressed by the great mass of seemingly uniform limestone , forming vertical cliffs over 1,000 feet (300 m) tall, and noted that much of the limestone

1827-666: The area around El Capitan, permanent Anglo-European presence in the area dates from the late 1840s, when the region officially became a part of the United States after the American victory in the Mexican–American War . Eager to find a suitable trail from San Antonio to El Paso , and eventually the far-flung American territories on the Pacific coast, a number of expeditions pushed into the hot, dry, dangerous lands west of

1890-552: The area was a seasonal home to bands of the Mescalero Apache , who, steadily pushed south by the fiercely expansionist Comanche in the 18th century, used their local knowledge of springs and other scarce resources to maintain themselves in an otherwise harsh environment well into the 19th century, when Anglo expansion and settlement ended their delicately balanced way of life. While the occasional Spanish expedition, such as that of Antonio de Espejo in 1582, came close to

1953-424: The area. George Shumard was the first geologist to study the area, in 1855, and described an "upper white limestone" containing fossils. These included fusulinids and brachiopods , that were identified correctly by his brother, B.F. Shumard, as Permian in age. However, debate on whether the beds were Carboniferous or Permian in age continued until at least 1920. The work of Darton and Reeside in 1926 established

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2016-512: The calcified cell wall of coralline algae was found to have a high correspondence with molecular studies. These skeletal structures thus provide morphologic evidence for molecular relationships within the group. According to AlgaeBase : According to the World Register of Marine Species : According to ITIS : Fresh surfaces are generally colonized by thin crusts, which are replaced by thicker or branched forms during succession over

2079-752: The community level; the presence of herbivores associated with corallines can generate patchiness in the survival of young stages of dominant seaweeds. This has been seen this in eastern Canada , and it is suspected the same phenomenon occurs on Indo-Pacific coral reefs , yet nothing is known about the herbivore enhancement role of Indo-Pacific corallines, or whether this phenomenon is important in coral reef communities. Some coralline algae develop into thick crusts which provide microhabitat for many invertebrates. For example, off eastern Canada , Morton found juvenile sea urchins , chitons , and limpets suffer nearly 100% mortality due to fish predation unless they are protected by knobby and undercut coralline algae. This

2142-503: The course of one (in the tropics) to ten (in the Arctic) years. However, the transition from crusts to branched form depends on environmental conditions. Crusts may also become detached and form calcareous nodules known as Rhodoliths . Their growth may be also disrupted by local environmental factors. While coralline algae are present in most hard substrate marine communities in photic depths, they are more common in higher latitudes and in

2205-588: The discovery in May 1923 of the Big Lake oil field in Texas and the drilling of the first commercial oil well in southeastern New Mexico in 1924. This culminated in the publication by E. Russell Lloyd in 1929 of his interpretation of the Capitan Limestone and associated formations as a gigantic fossil coral reef. Lloyd traced the reef nearly to Carlsbad and noted that the dissimilarity of the formations on

2268-747: The dolomitic Rustler Formation, and the Dewey Lake Formation, burying the reef even deeper. The Capitan Reef stayed buried for over 150 million years, until the late Cretaceous period of the Mesozoic era (80 million years ago), when tectonic activity associated with the Laramide Orogeny caused a significant uplift in the area and created a major fault line, the Border Fault, in the area west of the Delaware Basin. Exposing

2331-459: The early decades of the 20th century, an all-weather road had been constructed between El Paso and Carlsbad, New Mexico . Commissioned in 1928 as U.S. Highway 62 , the road brought a new generation of visitors to the area, particularly after the opening of nearby Carlsbad Caverns National Park in 1930. Decades before the creation of Guadalupe Mountains National Park , travelers by car and bus were already stopping at roadside rest stops to marvel at

2394-520: The end of the 18th century. Medical science now uses corallines in the preparation of dental bone implants. The cell fusions provide the matrix for the regeneration of bone tissue. Maërl is also used as a food additive for cattle and pigs , as well as in the filtration of acidic drinking water. As a colorful component of live rock sold in the marine aquarium trade, and an important part of reef health, coralline algae are desired in home aquariums for their aesthetic qualities, and ostensible benefit to

2457-573: The expedition of Diego de Vargas. Quickly becoming an important local resource, generations of Mexicans, and later, Mexican-Americans, braved the hot, dangerous, four-day trail from San Elizario on the Rio Grande to the Hueco Mountains and then east towards El Capitan to fill their wagons with the precious salt. This all changed in the late 1840s, when the region began to have a larger Anglo presence. While Mexican law and tradition had held

2520-697: The lineage has a much longer history than molecular clocks would indicate. The earliest known coralline deposits date from the Ordovician , although modern forms radiated in the Cretaceous . True corallines are found in rocks of Jurassic age onwards. Stem group corallines are reported from the Ediacaran Doushantuo formation ; later stem-group forms include Arenigiphyllum , Petrophyton , Graticula , and Archaeolithophyllum . The corallines were thought to have evolved from within

2583-519: The main reef structures that prevent oceanic waves from striking adjacent coastlines , helping to prevent coastal erosion . Because of their calcified structure, coralline algae have a number of economic uses. Some harvesting of maërl beds that span several thousand kilometres off the coast of Brazil takes place. These beds contain as-yet undetermined species belonging to the genera Lithothamnion and Lithophyllum . The collection of unattached corallines (maërl) for use as soil conditioners dates to

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2646-479: The maximum penetration of light). Some species can tolerate brackish or hypersaline waters, and only one strictly freshwater coralline species exists. (Some species of the morphologically similar, but non-calcifying, Hildenbrandia , however, can survive in freshwater.) A wide range of turbidities and nutrient concentrations can be tolerated. Corallines, especially encrusting forms, are slow growers, and expand by 0.1–80 mm annually. All corallines begin with

2709-507: The mountains, and an easy path through Guadalupe Pass to the Pecos. This road, which became known as the "upper" road, to distinguish itself from the original "lower" road through Fort Davis and up the Rio Grande to El Paso, soon became popular among traders and emigrants, their long trains of wagons and mules passing near the brooding face of El Capitan. This traffic increased in 1858 as the Butterfield Stagecoach began to use

2772-467: The order Corallinales . They are characterized by a thallus that is hard because of calcareous deposits contained within the cell walls. The colors of these algae are most typically pink, or some other shade of red, but some species can be purple, yellow, blue, white, or gray-green. Coralline algae play an important role in the ecology of coral reefs . Sea urchins , parrot fish , and limpets and chitons (both mollusks) feed on coralline algae. In

2835-626: The prospect of returning by the same route, the party instead struck off east from El Paso towards the Hueco Tanks and Guadalupe Pass, which they hoped to follow to an established crossing of the Pecos River near the modern Texas-New Mexico border long used by Mexicans and Native Americans. Successful in reaching the Pecos, Neighbors quickly descended the river and made his way back to San Antonio, where he reported to his superiors that this route offered good water at Hueco Tanks, good timber near

2898-622: The protection of Forts Stockton and Davis, the Pinery station continued to be used by soldiers, freighters, and emigrants long after. During this period, El Capitan also bore witness to the El Paso Salt War, a violent struggle between Mexican-American residents and Anglo businessmen over access to the salt flats extending west from the base of the mountain. Long known to the Apache, the salt flats were first identified by Europeans in 1692 by

2961-535: The reef materials into a sturdy structure. Corallines are particularly important in constructing the algal ridge's reef framework for surf-pounded reefs in both the Atlantic and Indo-Pacific regions. Algal ridges are carbonate frameworks constructed mainly by nongeniculate coralline algae (after Adey, 1978). They require high and persistent wave action to form, so develop best on windward reefs with little or no seasonal change in wind direction. Algal ridges are one of

3024-443: The rock surfaces. These patches of pink "paint" are actually living crustose coralline red algae. The red algae belong to the division Rhodophyta , within which the coralline algae form the order Corallinales . There are over 1600 described species of nongeniculate coralline algae. The corallines are presently grouped into two families on the basis of their reproductive structures. Coralline algae are widespread in all of

3087-406: The salt flats as communal property, American tradition considered them unclaimed lands, which could be claimed by any citizen and purchased as private property. By the 1870s, attempts by local businessmen to claim the salt flats were being met with violent opposition by local residents, for whom the free salt was an important adjunct to the regional economy of farming and ranching. The conflict came to

3150-458: The settlement of the larvae of certain herbivorous invertebrates , particularly abalone . Larval settlement is adaptive for the corallines because the herbivores remove epiphytes which might otherwise smother the crusts and preempt available light. Settlement is also important for abalone aquaculture ; corallines appear to enhance larval metamorphosis and the survival of larvae through the critical settlement period. It also has significance at

3213-503: The sheer nature of the peak. While no trail to the summit has been developed, hikers can climb up to the top of El Capitan by first climbing to near the summit of Guadalupe Peak on the developed 4.5-mile Guadalupe Peak Trail, scrambling down to the south onto the Guadalupe Peak-El Capitan saddle, and then hiking up the back of El Capitan to the summit. Coralline algae Coralline algae are red algae in

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3276-418: The sheer sides of El Capitan, take pictures, and write quick messages on the backs of postcards emblazoned with the mountain’s imposing face. During this period, the land that included El Capitan was owned by James Adolphus Williams, whose Williams Ranch house is still extant on park land west of the mountain. After several economic setbacks, Williams sold the land in 1941 to Judge J.C. Hunter of Van Horn, Texas,

3339-594: The stratigraphic conventions in the Guadalupe Mountains, and removed the Gilliam and most of the Vidrio Limestone from the formation. The Capitan Formation consists of compact, massive, light grey to white limestone with minor dolomite . Its total thickness is 1,000–2,000 feet (300–610 m). On the backreef side of the formation, the Capitan rests on the Goat Seep Dolomite and grades into and

3402-402: The structure of the reef, help cement the reef together, and are important sources of primary production. Coralline algae are especially important in reef construction, as they lay down calcium carbonate as calcite. Although they contribute considerable bulk to the calcium carbonate structure of coral reefs, their more important role in most areas of the reef, is in acting as the cement which binds

3465-433: The surface of the coralline. This combination of grazing and burrowing results in a peculiar growth form (called "castles") in H. onkodes , in which the coralline produces nearly vertical, irregularly curved lamellae. Coralline algae are part of the diet of shingle urchins ( Colobocentrotus atratus ). Nongeniculate corallines are of particular significance in the ecology of coral reefs, where they add calcareous material to

3528-496: The surface of the plants. Not all sloughing serves an antifouling function. Epithallial shedding in most corallines is probably simply a means of getting rid of damaged cells whose metabolic function has become impaired. Morton and his students studied sloughing in the South African intertidal coralline alga, Spongites yendoi , a species which sloughs up to 50% of its thickness twice a year. This deep-layer sloughing, which

3591-588: The temperate Mediterranean Sea , coralline algae are the main builders of a typical algal reef, the Coralligène ("coralligenous"). Many are typically encrusting and rock-like, found in marine waters all over the world. Only one species lives in freshwater. Unattached specimens ( maerl , rhodoliths ) may form relatively smooth compact balls to warty or fruticose thalli. A close look at almost any intertidal rocky shore or coral reef will reveal an abundance of pink to pinkish-grey patches, distributed throughout

3654-617: The two sides of the reef, now known as the basin and backreef shelf facies. Two months later, a “Symposium on Pennsylvanian and Permian stratigraphy of southwestern United States” appeared in the August, 1929 issue of the Bulletin of the American Association of Petroleum Geologists , which provided a flood of new details on the Capitan reef. As part of that symposium, Philip B. King and R.E. King presented their conclusion that

3717-474: The upper road between El Paso and the ford on the Pecos called Pope’s Crossing , after John Pope , the Army officer who had surveyed the crossing two years before. Taking advantage of the water and timber available, the stagecoach line constructed a station at the Pinery, later called Pine Springs, 3 miles southeast of El Capitan. Though this station only lasted a year before the route was moved south to come under

3780-522: The world's oceans, where they often cover close to 100% of rocky substrata . Only one species, Pneophyllum cetinaensis , is found in freshwater. Its ancestor lived in brackish water, and was already adapted to osmotic stress and rapid changes in water salinity and temperature. Many are epiphytic (grow on other algae or marine angiosperms), or epizoic (grow on animals), and some are even parasitic on other corallines. Corallines have been divided into two groups, although this division does not constitute

3843-406: Was dolomitized . He was also impressed with the abundant fossils found in the middle beds of the formation, forming a fossil assemblage unlike anything else known at that time. Richardson interpreted the Guadalupe Mountains as an east-dipping monocline with a fault on the steep western boundary, and believed El Capitan itself was a product of erosion . Interest in the formation was rekindled by

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3906-411: Was officially opened, allowing public access to the area. As previously noted, El Capitan is located within the boundaries of Guadalupe Mountains National Park, a fee-area located on U.S. Highway 62/180 between Carlsbad and El Paso. The southern terminus of the Guadalupe range, El Capitan is guarded by cliffs on three sides, and those faces are rarely climbed due to the unstable condition of the rock and

3969-531: Was used as building stone since the ancient Greek culture. The calcite crystals composing the cell wall are elongated perpendicular to the cell wall. The calcite normally contains magnesium (Mg) , with the magnesium content varying as a function of species and water temperature. If the proportion of magnesium is high, the deposited mineral is more soluble in ocean water, particularly in colder waters, making some coralline algae deposits more vulnerable to ocean acidification . The first coralline alga recognized as

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