Genus ( / ˈ dʒ iː n ə s / ; pl. : genera / ˈ dʒ ɛ n ər ə / ) is a taxonomic rank above species and below family as used in the biological classification of living and fossil organisms as well as viruses . In binomial nomenclature , the genus name forms the first part of the binomial species name for each species within the genus.
194-721: Irritator is a genus of spinosaurid dinosaur that lived in what is now Brazil during the Albian stage of the Early Cretaceous Period , about 113 to 110 million years ago. It is known from a nearly complete skull found in the Romualdo Formation of the Araripe Basin . Fossil dealers had acquired this skull and sold it to the State Museum of Natural History Stuttgart . In 1996 ,
388-536: A display function when the animal was alive. The preserved part of Irritator ' s crest is deepest above the antorbital fenestra and lacks the vertical ridges seen in the crest of Spinosaurus . Like others in its family , Irritator possessed a long and bony structure on the roof of its mouth called a secondary palate , separating the oral from the nasal cavity. This is a feature observed in extant crocodilians , but absent in most theropod dinosaurs. Also like its relatives, Irritator had two additional openings on
582-551: A pterosaur , and later a maniraptoran dinosaur. In 1996, the animal was identified as a spinosaurid theropod . The holotype skull was thoroughly prepared before being redescribed in 2002, confirming this classification. Both Irritator and Angaturama belong to the Spinosaurinae subfamily . A generalist diet —like that of today's crocodilians —has been suggested; Irritator might have preyed mainly on fish and any other small prey animals it could catch. Fossil evidence
776-495: A chalk concretion containing the rear of a large skull with lower jaws near the town of Santana do Cariri in northeastern Brazil . This fossil was acquired by dealers who sold it to Rupert Wild of the State Museum of Natural History Stuttgart , Germany. At the time it was assumed to be the skull of a giant basal pterosaur , or flying reptile, since the Chapada do Araripe region is famous for its copious pterosaur finds, and
970-625: A common dental trait among spinosaurids. Both sides of Irritator ' s teeth were fluted, as in Spinosaurus , whereas Baryonyx exhibited them only on the lingual (inward facing) side of its teeth. Irritator ' s teeth were circular in cross section, as opposed to the laterally flattened condition of most theropod teeth. The enamel (first layer of the teeth) was thin, with a finely wrinkled texture also observed in Baryonyx. Both of Irritator ' s maxillae preserve nine teeth, although
1164-724: A consequence of the Atlantic Ocean , whose opening gradually separated the continents and contributed to differences between taxa. A similar scenario was suggested in 2014 by Brazilian paleontologist Manuel A. Medeiros and colleagues for the fauna of the Alcântara Formation, where Oxalaia has been found. But the paleobiogeography of spinosaurids remains highly theoretical and uncertain, with discoveries in Asia and Australia indicating that it may have been complex. The taphonomy (changes between death and fossilization) of
1358-487: A distinct ecological niche , therefore avoiding competition with more terrestrial theropods. Spinosaurines appear to have been more adapted for such lifestyles than baryonychines. A 2018 study by British paleontologist Thomas M. S. Arden and colleagues examined the morphology of spinosaurine skull bones for possible aquatic traits. They found that the frontal bones of Irritator , Spinosaurus , and Sigilmassasaurus were similar in being arched, concave on top, and narrowed at
1552-572: A fire broke out at the palace housing the Museu Nacional, largely destroying the fossil collections and possibly the exhibited Angaturama skeleton and fossil elements. The holotype of Oxalaia quilombensis , which was stored in the same building, may also have been destroyed. Even by maximal size estimates, Irritator was smaller than other known spinosaurids. Gregory S. Paul calculated its length at 7.5 meters (25 ft) and weight at 1 tonne (1.1 short tons). Thomas R. Holtz Jr. published
1746-407: A genus is determined by taxonomists . The standards for genus classification are not strictly codified, so different authorities often produce different classifications for genera. There are some general practices used, however, including the idea that a newly defined genus should fulfill these three criteria to be descriptively useful: Moreover, genera should be composed of phylogenetic units of
1940-541: A group of municipalities. These regions were created by the Brazilian Institute of Geography and Statistics for statistical purposes and do not, therefore, constitute an administrative area. Municipalities are analogous to counties in states of the United States; a city is the urban area of the municipality, and always has the same name as the municipality. A municipality may include cities other than
2134-484: A higher estimate of 8 meters (26 ft), with a weight between 0.9 to 3.6 tonnes (0.99 to 3.97 short tons). Estimates by Dougal Dixon were lower at 6 meters (20 ft) long and 2 meters (6.6 ft) high. When scaled by Aureliano and colleagues, the reconstructions from the study by Sales and Schultz provided a length of 6.5 meters (21 ft) for the Irritator challengeri holotype, and 8.3 meters (27 ft) for
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#17327799742602328-651: A later homonym of a validly published name is a nomen illegitimum or nom. illeg. ; for a full list refer to the International Code of Nomenclature for algae, fungi, and plants and the work cited above by Hawksworth, 2010. In place of the "valid taxon" in zoology, the nearest equivalent in botany is " correct name " or "current name" which can, again, differ or change with alternative taxonomic treatments or new information that results in previously accepted genera being combined or split. Prokaryote and virus codes of nomenclature also exist which serve as
2522-477: A left tibia from a particularly large individual. As is common with fossils from the Araripe Basin , the majority of spinosaurid material from the Romualdo Formation was collected under uncontrolled circumstances for use in the illegal fossil trade. As such, many specimens are partly damaged and without precise geological field data. Some of the Romualdo Formation postcrania were used as the basis for
2716-739: A long stretch of coast that was most accessible to Europe without, however, penetrating the interior. The colonists of the Dutch West India Company (WIC) in Brazil were under constant siege despite the presence in Recife of John Maurice of Nassau as governor. To help fight the Portuguese, the WIC sought the support of native peoples. By 1635, the majority of Tupi, mostly from Rio Grande do Norte and Paraíba , had given their support to
2910-628: A long time and redescribed as new by a range of subsequent workers, or if a range of genera previously considered separate taxa have subsequently been consolidated into one. For example, the World Register of Marine Species presently lists 8 genus-level synonyms for the sperm whale genus Physeter Linnaeus, 1758, and 13 for the bivalve genus Pecten O.F. Müller, 1776. Within the same kingdom, one generic name can apply to one genus only. However, many names have been assigned (usually unintentionally) to two or more different genera. For example,
3104-483: A major producer of cashew nuts , sugar cane , cocoa , cotton and tropical fruits in general (mainly coconut , papaya , melon , banana , mango , pineapple and guarana ). It also has relevant productions of soy , maize , bean , cassava and oranges . The region is subject to prolonged dry spells that are worse in El Niño years. This causes a periodic rural exodus . Government responses include dams and
3298-407: A member of the same genus, the latter would be the valid scientific name under rules of priority , since it was named almost a month earlier. The paleontologist Marco Schade and colleagues could not confirm the overlap in tooth positions in 2023 due to interpreting them differently, and could not provide information to resolve the issue. Besides the skull, the snout fragment, and some isolated teeth,
3492-473: A more developed obturator process , a blade-like structure on the bottom of the ischium. Martill and his team originally classified Irritator as a maniraptoran dinosaur in the clade Bullatosauria (a group no longer considered monophyletic ), as a close relative to the feathered ornithomimosaurs and troodontids . Given that its dental morphology , particularly long snout, and assumed fin-shaped crest were features unknown in "other" maniraptorans,
3686-446: A partial right tibia and fibula (shin and calf bones), most of the right femur (thigh bone), and part of an ulna (forearm bone). It also has the most complete hand known from a spinosaurid, including metacarpals , phalanges , one carpals , and one claw . Like in all spinosaurids, the claw of the first finger (the "thumb") was enlarged. The pelvic bones are well preserved, with
3880-719: A partly piscivorous (fish-eating) diet. Although much of the holotype's morphology turned out to be greatly different than they thought, later studies supported these observations. Spinosaurids had very narrow and elongated jaws with relatively homogeneous pointed teeth, an arrangement found particularly in animals like the Indian gharial —the most piscivorous extant crocodilian. The long conical teeth, which in spinosaurines did not possess serrated edges, were suitable for grabbing and holding prey. They differed from teeth of other theropods, which seemed geared towards tearing or cutting off seized body parts. Irritator shared with crocodilians
4074-478: A possible point of overlap, the third left maxillary tooth, and observed that the skull of Angaturama could have been larger than that of Irritator based on the proportions of the closely related genus Baryonyx . They therefore concluded that the two specimens do not belong to the same individual, Sales and Schultz noted that synonymy at the genus level would need to be verified by more extensively overlapping remains. If Angaturama and Irritator are regarded as
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#17327799742604268-760: A process of industrialization around Salvador, Fortaleza, and Recife, as well as a burgeoning agriculture industry around the São Francisco river . The regions of Brazil do not have their own governmental or administrative bodies, but they are well defined. Their boundaries and constituent states are part of the recognized geopolitical structure of the country. The Northeast Region is composed of nine states, with 1793 municipalities and two special municipalities, Saint Peter and Saint Paul Archipelago and Fernando de Noronha Archipelago; there are no unincorporated areas. Brazilian states are divided into Mesoregions, and Mesoregions into Microregions, each region representing
4462-409: A reference for designating currently accepted genus names as opposed to others which may be either reduced to synonymy, or, in the case of prokaryotes, relegated to a status of "names without standing in prokaryotic nomenclature". An available (zoological) or validly published (botanical) name that has been historically applied to a genus but is not regarded as the accepted (current/valid) name for
4656-479: A sagittal crest on Angaturama ' s premaxillae may correspond with that of Irritator ' s nasal bones. Some objection has been raised to these assertions. Kellner and Campos in 2000 and Brazilian paleontologist Elaine B. Machado and Kellner in 2005 expressed the opinion that the fossils come from two different genera, and that the holotype of Angaturama limai was clearly more laterally flattened than that of Irritator challengeri . A review of both fossils by
4850-435: A septarian concretion . The tip of the upper jaw was also missing. Since there were no signs of erosion, it had most probably broken off during or after the fossil's collection. Evident corrosion on certain bones indicates acid preparation had been attempted. A vertical fracture was present across the middle of the skull, which had apparently been sealed with car body filler. In hopes of making it look more complete and valuable,
5044-551: A sideways-facing opening in the lower jaw, was oval and comparatively large. The dentary (tooth-bearing bone of the mandible) is unknown in Irritator , save for a possible remnant at the front of the surangular. Irritator challengeri ' s holotype is unique in that it is one of the few non-avian (or non-bird) dinosaur fossils found with a preserved stapes . Irritator had straight or only faintly recurved conical teeth, bearing sharply defined but unserrated edges. Flutes (lengthwise ridges) were present on its tooth crowns,
5238-477: A snout tip that was described a few weeks later also in 1996—as a potential junior synonym of Irritator . Both animals hail from the same stratigraphic units of the Araripe Basin. It was also previously proposed that Irritator and Angaturama ' s skull parts belonged to the same specimen. Although this has been cast into doubt, more overlapping fossil material is needed to confirm whether they are
5432-422: A so-called indústria da seca (drought industry). The term, coined by Antônio Callado , describes the phenomenon wherein local politicians use droughts as a tool to increase their electoral chances, effectively "trading drought relief for votes." This misappropriation of funds often benefits the wealthy—especially private landowners—to the detriment of the rural poor. Meio-Norte is a transition area between
5626-602: A spinosaurid and synonymized Irritatoridae with that family. Irritator was then assigned to the Baryonychidae along with Angaturama , Baryonyx , Suchomimus, and Spinosaurus by Oliver W.M. Rauhut in 2003. Thomas Holtz and colleagues in 2004 considered the Baryonychidae synonymous with Spinosauridae, and moved these genera to the latter family. Most later revisions have upheld these classifications. As spinosaurids, Irritator and Angaturama are placed within
5820-525: A stiff secondary palate and reduced antorbital fenestrae. In 2007, a finite-element analysis study by British paleontologist Emily J. Rayfield and colleagues found that these attributes, present in other spinosaurids as well, made the skull more resistant to torsion from prey item loads when feeding. The authors pointed out that in contrast, most theropods lacked secondary palates and had large antorbital fenestrae, exchanging strength for lighter skull builds. The nostrils of Irritator were shifted far back from
6014-418: A subadult, Irritator challengeri ' s holotype remains the most completely preserved spinosaurid skull yet found. The Angaturama snout tip expanded to the sides in a rosette -like shape, bearing long teeth and an unusually tall crest. One possible skeleton indicates it, like other spinosaurids, had enlarged first-finger claws and a sail running down its back. Irritator had been mistaken initially for
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6208-452: A subadult, so the mature animal may have been larger. The holotype skull of Irritator challengeri , although heavily damaged in certain places, is largely complete; missing only the tip of the snout and the front of the mandible . The preserved skull is 16.5 cm (6.5 in) tall and 10 cm (3.9 in) wide, its full length has been estimated at 60 cm (24 in), based on comparisons with Baryonyx . Irritator ' s skull
6402-427: A taxon; however, the names published in suppressed works are made unavailable via the relevant Opinion dealing with the work in question. In botany, similar concepts exist but with different labels. The botanical equivalent of zoology's "available name" is a validly published name . An invalidly published name is a nomen invalidum or nom. inval. ; a rejected name is a nomen rejiciendum or nom. rej. ;
6596-448: A theropod's buoyancy to a significant extent. In 2020, German paleontologist Marco Schade and colleagues analyzed the anatomy of the holotype skull braincase through CT scans, revealing numerous details about behavioral capabilities of Irritator . With the scans, they created a 3D model of the skull and braincase, discovering that Irritator had elongated olfactory tracts and a relatively large floccular recesses (area that pierces through
6790-406: A total of 17.6 million tons in 2018. Maranhão was the 7th largest producer in the country, with 681 thousand tons. Ceará was 9th, with 622 thousand tons. Bahia was 10th with 610 thousand tons. In total, the northeast produced 3,5 million tons. Regarding oranges , Bahia was the 4th largest producer in Brazil in 2018, with a total of 604 thousand tons. Sergipe was 6th, with 354 thousand tons. Alagoas
6984-455: A total of c. 520,000 published names (including synonyms) as at end 2019, increasing at some 2,500 published generic names per year. "Official" registers of taxon names at all ranks, including genera, exist for a few groups only such as viruses and prokaryotes, while for others there are compendia with no "official" standing such as Index Fungorum for fungi, Index Nominum Algarum and AlgaeBase for algae, Index Nominum Genericorum and
7178-764: A value of R$ 254 million. In vanadium , it produced 358 thousand tons, at a value of R$ 91 million. In the extraction of precious and semi-precious stones, Bahia has small or medium scale productions of amethyst, agate, diamond, emerald, garnet, opal, ruby, tourmaline and turquoise. There is also aquamarine production in Rio Grande do Norte, Ceará, Alagoas and Paraíba; grenades in Paraíba, Ceará and Rio Grande do Norte; opal in Piauí and Ceará; tourmaline in Ceará and Tourmaline Paraíba in Paraíba and Rio Grande do Norte. In 2017,
7372-423: A “polygon of drought.” Due to global warming , more frequent, more severe and longer droughts are estimated to hit the region over the next 90 years. Despite the increasing severity of the droughts, Brazil has become increasingly able to mitigate the negative social impacts associated with drought. While generally successful in helping those enduring drought, drought relief programs have historically resulted in
7566-473: Is carried out almost exclusively in the Northeast. The area occupied by cashew trees in Brazil in 2017 was estimated at 505,500 ha; of this total, 99.5% is located in the Northeast. The main producers in this region are Ceará (61.6% of the national area), Rio Grande do Norte and Piauí. However, Brazil, which in 2011 was the fifth largest world producer of cashew nuts, in 2016, fell to 14th position, with 1.5% of
7760-612: Is discouraged by both the International Code of Zoological Nomenclature and the International Code of Nomenclature for algae, fungi, and plants , there are some five thousand such names in use in more than one kingdom. For instance, A list of generic homonyms (with their authorities), including both available (validly published) and selected unavailable names, has been compiled by the Interim Register of Marine and Nonmarine Genera (IRMNG). The type genus forms
7954-500: Is further distinguished from Baryonyx , Suchomimus , and Cristatusaurus by having slightly over half as many teeth in the maxilla, and from Spinosaurus due to its comparatively larger and more forwardly positioned nostril openings, which, unlike in Spinosaurus , are also formed by the premaxilla. The narrow sagittal crest, which ends in a knob-like process above the frontals, is another autapomorphy separating Irritator from other spinosaurids. Although Angaturama limai ' s snout
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8148-781: Is generally narrower than in other spinosaurids, this might be due to damage sustained by the fossil; the holotype appears partly crushed and broken on its lower margin, with some of the preserved teeth having been sectioned off along their length. Therefore, Angaturama ' s only valid autapomorphy is its sagittal crest, which extends farther forwards on the rostrum and is more exaggerated than in other known spinosaurid skulls. Topology A: Benson and colleagues (2009) Baryonyx [REDACTED] Megalosauridae [REDACTED] Topology B: Sales and Schultz (2017) Baryonyx [REDACTED] In 1996, Martill and colleagues theorized that Irritator challengeri, with its elongated snout and unserrated conical teeth, likely had at least
8342-647: Is in 4th place among the 5 regions, behind the Midwest, South and Southeast. None of the states in the Northeast Region of Brazil has a percentage of GDP production above its percentage of inhabitants in the country, therefore, being in deficit – with this, the Federal Government of Brazil is obliged to redirect taxes extracted in the Southeast and South to the Northeast Region. The region is
8536-409: Is in second place in cocoa almond, atemoia, cupuaçu, lime and lemon, and third in banana, carambola, guava, papaya, watermelon, melon, cherry, pomegranate and table grapes. In all, 34 products from Bahia's fruit culture have an important participation in the national economy. Rio Grande do Norte is the largest producer of melon in the country. In 2017 it produced 354 thousand tons, distributed between
8730-403: Is interpreted as a coastal lagoon with irregular freshwater influence that contended with cycles of transgressing and regressing sea levels . The climate of the formation was tropical and largely corresponded to today's Brazilian climate. The regions surrounding the formation were arid to semi-arid , with most of the local flora being xerophytic (adapted to dry environments). Cycadales and
8924-403: Is known of an individual that ate a pterosaur, either from hunting or scavenging it. Irritator may have had semiaquatic habits, and inhabited the tropical environment of a coastal lagoon surrounded by dry regions. It coexisted with other carnivorous theropods as well as turtles, crocodyliforms , and a large number of pterosaur and fish species. The holotype of Irritator was excavated from
9118-554: Is no longer enough rainfall to support tropical rainforest, especially since the rainfall is extremely erratic from year to year. This transitional zone is known as the agreste and because it is located on the steep escarpment, was not generally used whilst flatter land was abundant. Today, with irrigation water available, however, the agreste , as its name suggest, is a major farming region. Despite containing no major city, it contains well developed medium large cities such as Caruaru , Campina Grande and Arapiraca . In Portuguese ,
9312-602: Is not known if it actively hunted these animals or simply scavenged the remains. In 2018, Aureliano and colleagues presented a possible scenario for the food web of the Romualdo Formation. The researchers proposed that spinosaurines from the formation may have also preyed on terrestrial and aquatic crocodyliforms , juveniles of their own species, turtles, and small to medium-sized dinosaurs. This would have made spinosaurines apex predators within this particular ecosystem. Examinations and digital reconstructions of Irritator published in 2023 by Schade, Rauhut et al., suggests that
9506-555: Is out of reach of the Intertropical Convergence Zone . The easterly trade winds blow across the region and bring abundant rainfall to the coast but producing clear, dry conditions inland where the escarpment blocks moisture flow. This gives rise to four distinct regions, the zona da mata on the coast, the agreste on the escarpment, sertão beyond and the Mid north. Before the arrival of Europeans, Nordeste
9700-502: Is practically three times larger than that of Pará, but Pará's productivity is practically three times greater. Some factors that explain this are: the crops in Bahia are more extractivist, and those in Pará have a more modern and commercial style, in addition to paraenses using more productive and resistant seeds, and their region providing resistance to Witch's broom . In 2018, the Northeast
9894-460: Is somewhat arbitrary. Although all species within a genus are supposed to be "similar", there are no objective criteria for grouping species into genera. There is much debate among zoologists about whether enormous, species-rich genera should be maintained, as it is extremely difficult to come up with identification keys or even character sets that distinguish all species. Hence, many taxonomists argue in favor of breaking down large genera. For instance,
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#173277997426010088-474: Is the type species , and the generic name is permanently associated with the type specimen of its type species. Should the specimen turn out to be assignable to another genus, the generic name linked to it becomes a junior synonym and the remaining taxa in the former genus need to be reassessed. In zoological usage, taxonomic names, including those of genera, are classified as "available" or "unavailable". Available names are those published in accordance with
10282-467: Is uncertain whether this specimen can be referred to Irritator or Angaturama , given that both are based only on skull material. In 2007, Machado and Kellner tentatively referred a rib fragment (MN 7021-V) to the Spinosauridae. However, the most complete spinosaur specimen retrieved from the Romualdo Formation is MN 4819-V, a partial skeleton lacking the skull. First reported in 1991, the specimen
10476-412: The alveoli (tooth sockets), the premaxilla had seven teeth altogether, the third tooth being the largest. The frontmost three teeth of the maxilla were also preserved. The premaxillary teeth increased in size from the first to third, shrank from the third to the sixth, and enlarged again from the sixth premaxillary to third maxillary positions. A 16 cm (6.3 in) diastema (gap in tooth row)
10670-569: The external nares (bony nostrils). As in all spinosaurids, the maxillae (main upper jaw bones) extended below and past the nostrils in a long, low branch that formed the lower border of this opening, consequently separating the premaxillae and nasal bones in that location. Irritator ' s maxillary sinuses (located in the body of the maxillae) bore a large oval opening, as in Allosaurus . The nostril openings were oval and, as in all spinosaurids, positioned farther back on
10864-417: The postacetabular ala (rear expansion). The preacetabular ala were somewhat enlarged at the front, in contrast to the more slender condition of the postacetabular ala. The brevis fossa (groove at the bottom of the postacetabular ala) was concave, as was the rear margin of the ischium. The pubis bore a relatively large and almost closed obturator notch , an indentation in the lower margin of
11058-464: The Angaturama limai holotype. Some of the skull bones of the former holotype had not yet fully co-ossified (fused), indicating that the specimen belonged to a subadult. The partial spinosaurine skeleton MN 4819-V represented a moderately sized individual, estimated by Machado at 5 to 6 meters (16 to 20 ft) in length. Many elements from this specimen were incorporated into the skeletal mount in
11252-675: The Atlantic seaboard in the northeast and southeast, northwest and west to the Amazon Basin and south through the Espinhaço highlands in southern Bahia . It encloses the São Francisco River and drainage basin , which were instrumental in the exploration, settlement and economic development of the region. The region lies entirely within the earth's tropical zone and encompasses Caatinga , Atlantic Forest and part of
11446-551: The Brazilian Institute of Geography and Statistics . Of Brazil 's twenty-six states, it comprises nine: Maranhão , Piauí , Ceará , Rio Grande do Norte , Paraíba , Pernambuco , Alagoas , Sergipe and Bahia , along with the Fernando de Noronha archipelago (formerly a separate territory, now part of Pernambuco). Chiefly known as Nordeste ("Northeast") in Brazil, this region was the first to be colonized by
11640-652: The Cerrado ecoregions . The climate is hot and semi-arid , varying from xeric in Caatinga, to mesic in Cerrado and hydric in the Atlantic Forest. The Northeast Region represents 18% of Brazilian territory, has a population of 57 million people, 28% of the total population of the country, and contributes 14.2% (2020) of Brazil's GDP. Nearly three quarters of the population live in urban areas clustered along
11834-621: The International Code of Zoological Nomenclature ; the earliest such name for any taxon (for example, a genus) should then be selected as the " valid " (i.e., current or accepted) name for the taxon in question. Consequently, there will be more available names than valid names at any point in time; which names are currently in use depending on the judgement of taxonomists in either combining taxa described under multiple names, or splitting taxa which may bring available names previously treated as synonyms back into use. "Unavailable" names in zoology comprise names that either were not published according to
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#173277997426012028-824: The International Plant Names Index for plants in general, and ferns through angiosperms, respectively, and Nomenclator Zoologicus and the Index to Organism Names for zoological names. Totals for both "all names" and estimates for "accepted names" as held in the Interim Register of Marine and Nonmarine Genera (IRMNG) are broken down further in the publication by Rees et al., 2020 cited above. The accepted names estimates are as follows, broken down by kingdom: The cited ranges of uncertainty arise because IRMNG lists "uncertain" names (not researched therein) in addition to known "accepted" names;
12222-414: The Irritator challengeri holotype specimen has been discussed by some researchers. The skull was found lying on its side. Preceding fossilization, several bones from the back of the braincase, as well as the dentary, splenial , coronoid, and right angular bones from the lower jaw, were lost. Other bones, mostly from the skull rear, had become disarticulated and displaced towards alternate regions of
12416-459: The Portuguese and other European peoples, playing a crucial role in the country's history . Nordeste' s dialects and rich culture, including its folklore, cuisines, music and literature, became the most easily distinguishable across the country. To this day, Nordeste is known for its history and culture, as well as for its natural environment and its hot weather. Nordeste stretches from
12610-562: The Romualdo Formation , whose rocks are dated to the Albian stage of the Early Cretaceous Period , about 110 million years ago. During this time, the Southern Atlantic Ocean was opening, forming the series of circum-Atlantic basins of southern Brazil and southwestern Africa, but the northeastern part of Brazil and West Africa were still connected. The Romualdo Formation is part of the Santana Group and, at
12804-686: The Sertão consists mainly of low uplands that form part of the Brazilian Highlands . Most parts of the sertão are between 200 and 500 meters above sea level, with higher elevations found on the eastern edge in the Planalto da Borborema , where it merges into a sub-humid region known as agreste , in the Serra da Ibiapaba in western Ceará and in the Serro do Periquito of central Pernambuco . In
12998-571: The automotive sector represents close to 22% of industrial GDP. Bahia had a Ford factory. It was created in Camaçari (2001). The Bahian automotive sector, led by Ford was in 2005 the third largest contributor (14.6%) to the Bahian GDP . Currently the state is trying to replace Ford, which stopped producing in Brazil. The state also has a petrochemical complex in Camaçari. In Pernambuco,
13192-446: The nasal cavity . Spinosaurus itself probably made heavier use of senses like vision or the mechanoreceptors on the tip of its snout, like those used by crocodilians to sense prey moving in the water. Another trait spinosaurs shared with gharials was an enlarged snout tip that wielded a rosette-shaped array of interlocking teeth, adept for impaling and catching fish. Although to a lesser degree than most known spinosaurs, this feature
13386-419: The nomenclature codes , which allow each species a single unique name that, for animals (including protists ), plants (also including algae and fungi ) and prokaryotes ( bacteria and archaea ), is Latin and binomial in form; this contrasts with common or vernacular names , which are non-standardized, can be non-unique, and typically also vary by country and language of usage. Except for viruses ,
13580-404: The platypus belongs to the genus Ornithorhynchus although George Shaw named it Platypus in 1799 (these two names are thus synonyms ) . However, the name Platypus had already been given to a group of ambrosia beetles by Johann Friedrich Wilhelm Herbst in 1793. A name that means two different things is a homonym . Since beetles and platypuses are both members of the kingdom Animalia,
13774-608: The superfamily Megalosauroidea , with Spinosauridae being a possible sister taxon to the Megalosauridae . In 1998, Sereno and colleagues defined two subfamilies within the Spinosauridae based on craniodental (skull and tooth) characteristics. They were Spinosaurinae, where they placed Spinosaurus and Irritator ; and Baryonychinae, to which they assigned Baryonyx, Suchomimus, and Cristatusaurus . Spinosaurines were distinguished by their unserrated, straighter, and more widely spaced teeth, as well as smaller first teeth of
13968-501: The transfer of the São Francisco River . The worst recent droughts were in 1993, 1998 and 1999. The latter was the worst in fifty years. In 2017, the Northeast Region was the largest producer of coconut in the country, with 74.0% of national production. Bahia produced 351 million fruits, Sergipe, 234 million, and Ceará 187 million. However, the sector has been suffering strong competition and losing market to Indonesia,
14162-484: The Atlantic coast and about 15 million people live in the hinterland. It is an impoverished region: 43.5% of the population lives in poverty, defined as less than $ 2/day. The capital of each state including the states of Salvador , Recife , Fortaleza and São Luís is its largest city; those four capitals are coastal cities with a population of more than one million. Nordeste has nine international airports, and
14356-471: The Brazilian paleontologists Marcos A. F. Sales and Cesar L. Schultz in 2017 noted that the specimens also differ in other aspects of their preservation: the Irritator specimen is brighter in color and is affected by a vertical crack, while the Angaturama specimen bears many cavities; the damage to the teeth of the Irritator challengeri holotype is also much less severe. Sales and Schultz also identified
14550-583: The Dutch, as they viewed the Portuguese as more brutal and believed that they would be better off if the Dutch remained in Brazil. The military aid provided by the Tupi population proved to be useful in 1645, when Portuguese colonists who had remained in Dutch-controlled territory began to revolt. Tupi mediators such as Poty and Paraupaba were instrumental in maintaining strong Dutch-Tupi relations during
14744-482: The German museum often bought such pieces. As it promised to be a unique discovery of singular importance, German and British pterosaur experts were contacted to study the exemplar. A paper describing it as a pterosaur had already been submitted for publication when the authors, German paleontologist Eberhard Frey and British paleontologist David Martill, were disabused of this notion by the peer reviewers , who suggested
14938-592: The Itapecuru Group, part of the São Luís Basin. This larger species, known only from an isolated snout tip and upper jaw fragment, lived during the Cenomanian stage, around six to nine million years after Irritator and Angaturama . Oxalaia quilombensis is differentiated from Angaturama limai by its broader, more rounded snout and lack of a sagittal crest on the premaxillae. In September 2018,
15132-465: The National Museum of Rio de Janeiro, which had a length of 6 meters (20 ft) and a height of 2 meters (6.6 ft). However, spinosaurids from the Romualdo Formation possibly attained greater sizes. Although LPP-PV-0042 is represented only by a tibia fragment, Aureliano and colleagues estimated its length at roughly 10 meters (33 ft). Bone histology indicates that this individual was
15326-612: The Northeast Region had close to 13% of the country's industrial GDP. Bahia has 4.4% of the national industrial GDP, Pernambuco 2.7%, Ceará 1.9%, Maranhão 1.1%, Rio Grande do Norte 0.9%, Paraíba 0.7%, Sergipe 0.6%, Alagoas 0.5% and Piauí 0.4%,. It is the least industrialized region of the country, in proportion per inhabitant. The states that have some relevant industrial level are Bahia, Pernambuco and Ceará. Bahian industry have automobile and tyre industries, footwear and textiles, furniture, food and beverages, cosmetics and perfumes, information technology and naval sectors. In Brazil,
15520-409: The Northeast produced close to 10.7 million tons, or 9% of the Brazilian total. The largest producers in the Northeast were Bahia (5.3 million tons), Maranhão (3 million tons) and Piauí (2.4 million tons). In the production of maize , in 2018 Brazil was the 3rd largest producer in the world, with 82 million tons. The Northeast produced about 8.4% of the country's total. Bahia was the largest producer in
15714-537: The Northeast, with 2.2 million tons. Piauí was the 2nd largest producer in the Northeast, with 1.5 million tons, and Maranhão was the 3rd largest, with 1.3 million tons. In 2018, the South Region was the main producer of beans with 26.4% of the total, followed by the Midwest (25.4%), Southeast Region (25.1%), Northeast (20.6%) and North (2.5%). The largest producers in the Northeast were Ceará, Bahia, Piauí and Pernambuco. In cassava production, Brazil produced
15908-478: The Philippines and India, the world's largest producers, who even export coconut water to Brazil. In addition to climatic problems, the low productivity of coconut palms in the Northeast Region is the result of factors related to the variety of coconut harvested and the technological level used in coastal regions. In these areas, the semi-extractive cultivation system still prevails, with low fertility and without
16102-464: The Portuguese colonials for several decades, it was finally taken and destroyed and its leader Zumbi dos Palmares was captured and beheaded. His head was then displayed in a public plaza in Recife. Although the king of Portugal had abolished the enslavement of native peoples in 1570, it continued until the Marquis of Pombal declared them to be 'free men' in 1755. However, by that time the practice
16296-407: The Romualdo Formation has also yielded postcrania l remains that may belong to spinosaurids, many of which are hitherto undescribed, and all of them pertaining to the Spinosaurinae subfamily . In 2004, parts of a spinal column (MN 4743-V) were unearthed at the formation. Brazilian paleontologist Jonathas Bittencourt and Kellner assigned these, due to their structure, to the Spinosauridae. It
16490-441: The Romualdo Formation is indeed exposed there, and the matrix encasing the holotype has the same color and texture as those rocks, this locality can be regarded as the probable site of the discovery of the fossil. Irritator challengeri was the first dinosaur described from the Romualdo Formation, and its holotype specimen represents the most completely preserved spinosaurid skull known. Angaturama limai , another spinosaurid from
16684-540: The Romualdo Formation tibia fragment. CT scanning of the specimen in the University of São Carlos revealed the presence of osteosclerosis (high bone density). This condition had previously only been observed in Spinosaurus aegyptiacus, as a possible way of facilitating submersion in water by making its bones heavier. The presence of this condition on the Brazilian leg fragment showed that compact bones had already evolved in spinosaurines at least 10 million years before
16878-568: The Romualdo Member of the then Santana Formation ). This assignment was confirmed by microfossils of the ostracod Pattersoncypris , and fish scales from the ichthyodectid Cladocyclus , both found in the Romualdo Formation. Questioning of local fossil dealers hinted at a locality near the village of Buxexé close to Santana Do Cariri at the flank of the Chapada do Araripe, at a height of approximately 650 meters (2,130 ft). Since
17072-615: The Treaty of Tordesillas, plundered Nordeste' s coast, sacked Bahia in 1604, and even temporarily captured Salvador , which had been Brazil's first capital and general seat of government since 1549. However, the Portuguese soon regained control of Salvador, and the Dutch were unable to recapture it, despite repeated attempts. In 1630, the Dutch captured Pernambuco and made Recife ( Dutch : Mauritsstad ) their capital. By 1640, they had set up more permanently in Nordeste and controlled
17266-512: The aboriginal Tupi Indian language of Brazil. The specific name honors the late Brazilian paleontologist Murilo R. de Lima, who informed Kellner of the specimen in 1991. In 1997, British paleontologists Alan J. Charig and Angela C. Milner considered Angaturama a likely junior synonym of Irritator , noting that both genera had retracted nostrils, long jaws, and characteristic spinosaurid dentition. Paul Sereno and colleagues in 1998 agreed with this possibility, and additionally observed that
17460-410: The adoption of cultural management practices. The three states that have the largest production, Bahia, Sergipe and Ceará, present a yield three times lower than that of Pernambuco, which is in 5th place in the national production. This is because most of the coconut trees in these three states are located in coastal areas and cultivated in semi-extractivist systems. The production of cashew in Brazil
17654-534: The appearance of Spinosaurus in Morocco. According to phylogenetic bracketing —a method used to infer unknown traits in organisms by comparison with their relatives—osteosclerosis therefore might have been the norm in the Spinosaurinae. The significance of these traits was questioned in a later 2018 publication, where Canadian paleobiologist Donald Henderson argued that osteosclerosis would not have changed
17848-423: The authors to suggest a possible junior synonymy of Irritator with Spinosaurus . Sues and colleagues noted that more overlapping skull material was needed for further diagnosis. As more of Spinosaurus ' s skull became known, later research maintained separation of the two taxa. Although the site of discovery is uncertain, the specimen most probably stems from the Romualdo Formation (previously designated
18042-442: The base for higher taxonomic ranks, such as the family name Canidae ("Canids") based on Canis . However, this does not typically ascend more than one or two levels: the order to which dogs and wolves belong is Carnivora ("Carnivores"). The numbers of either accepted, or all published genus names is not known precisely; Rees et al., 2020 estimate that approximately 310,000 accepted names (valid taxa) may exist, out of
18236-413: The cities of Mossoró, Tibau and Apodi. The Northeast region accounted for 95.8% of the country's production in 2007. In addition to Rio Grande do Norte, which in 2005 produced 45.4% of the country's total, the other 3 largest in the country were Ceará, Bahia and Pernambuco. In the production of papaya , in 2018 Bahia was the 2nd largest producer state in Brazil, almost equaling with Espírito Santo. Ceará
18430-406: The cities of the northeastern countryside. These fairs gave rise to cities like Campina Grande, Feira de Santana and Caruaru. The Northeast region housed 93.2% of the goat herd (8,944,461 heads) and 64.2% of the sheep herd (11,544,939 heads) in Brazil in 2017. Bahia concentrated 30.9% of the goat herd and 20.9% of the national sheep herd. Casa Nova (BA) took first place in the municipal ranking with
18624-538: The country, namely the extraction and export of pau Brasil, or brazilwood . Indigenous peoples helped Europeans with the extraction of brazilwood in exchange for spices. They also engaged in an exchange of goods ( Portuguese : escambo ), trading things like animal skins for knives and other valuables. Brazilwood was highly valued in Europe where it was used to make violin bows (especially the Pau de Pernambuco variety) and for
18818-671: The creation of a replica Angaturama skeleton, later mounted at the Federal University-owned Museu Nacional do Rio de Janeiro ( National Museum of Rio de Janeiro ). The skeleton depicted the animal carrying an anhanguerid pterosaur in its jaws. It was the centerpiece of the Dinossauros no Sertão (Dinosaurs of Sertão) exhibit, which opened to the public in March 2009, becoming the first large carnivorous Brazilian dinosaur to be put on display. Some of
19012-729: The decline of the indigenous population, which intensified as colonization, commercial interest, and disease escalated in the region. After resistance from indigenous peoples and opposition to their enslavement from the Jesuits , the Portuguese colonials began importing black African slaves in 1530, largely to Bahia. In 1552, the seat of the first Catholic bishop of Brazil was established in Nordeste. French colonists not only tried to settle in present-day Rio de Janeiro , from 1555 to 1567 (the so-called France Antarctique episode), but also in present-day São Luís , from 1612 to 1614 (the so-called France Equinoxiale ). The Dutch , also opposed to
19206-507: The examinations reaffirmed classification of spinosaurids as megalosauroids and that megalosauroids and allosauroidea were part of a monophyletic Carnosauria, with Monolophosaurus standing as a sister taxon to spinosauridae. Many spinosaurs likely had semiaquatic habits, as has been shown in studies using techniques like isotope analysis and bone histology . It has been found that they probably took advantage of aquatic prey and environments (usually marginal and coastal habitats) to occupy
19400-435: The extinct conifer Brachyphyllum were the most widespread plants. This environment was dominated by pterosaurs, including: Anhanguera , Araripedactylus , Araripesaurus , Brasileodactylus , Cearadactylus , Coloborhynchus , Santanadactylus , Tapejara , Thalassodromeus , Tupuxuara , Barbosania , Maaradactylus , Tropeognathus , and Unwindia . The known dinosaur fauna besides Irritator
19594-525: The fact that the soil in Bahia is more favorable, in addition to the absence of diseases in the region. The most famous users of the product, however, acquire 90% to 100% of their guarana from the Amazon region, such as AMBEV and Coca-Cola . Bahian guarana prices are well below those of other states, but Sudam's tax exemptions lead the beverage industry to prefer to purchase seeds in the North, which helps maintain
19788-564: The feeling the authors felt (understated here) when discovering that the snout had been artificially elongated." The type species , Irritator challengeri, was named after Professor Challenger , a character in Arthur Conan Doyle 's novels, specifically The Lost World . Two years earlier, Frey and Martill had named a new pterosaur species from the Crato Formation Arthurdactylus conandoylei , after
19982-446: The form "author, year" in zoology, and "standard abbreviated author name" in botany. Thus in the examples above, the genus Canis would be cited in full as " Canis Linnaeus, 1758" (zoological usage), while Hibiscus , also first established by Linnaeus but in 1753, is simply " Hibiscus L." (botanical usage). Each genus should have a designated type , although in practice there is a backlog of older names without one. In zoology, this
20176-435: The fossil belonged to a theropod dinosaur. The skull was flattened sideways somewhat and, as is common with fossils, was partly crushed. The right side was well preserved, while the left was extensively damaged during collection. Some of the skull's hindmost upper surface had eroded, and the lower jaw lacked its front end, both owing to breakage during fossilization. Parts of the specimen were also cracked due to being part of
20370-492: The fossil skeleton of one Baryonyx . Naish and colleagues in 2004 supported the theory that Irritator hunted both aquatic and terrestrial animals as a generalist within the coastal area and in addition probably searched for carrion . A tooth belonging to Irritator was discovered still inserted into the fossil neck vertebral column of an ornithocheirid pterosaur, likely with a wingspan of 3.3 m (11 ft). This indicates that Irritator ate pterosaurs as well, although it
20564-497: The fossil traders had severely obscured the skull beneath plaster; a widespread practice among local collectors in the Chapada do Araripe, especially on fish fossils. The buyers were unaware of the modifications to the specimen until it was sent to universities in the United Kingdom for CT scan imaging. This revealed the collectors had tried to reconstruct the skull by grafting parts of the maxilla (main upper jaw bone) onto
20758-490: The front of the rostrum (snout). The skull (designated SMNS 58022) became the holotype specimen of the new genus and species Irritator challengeri in February 1996 , when it was first scientifically described by paleontologists David M. Martill, Arthur R.I. Cruickshank, Eberhard Frey, Philip G. Small and Maria Clarke. In this paper, Martill and his team wrote that the generic name Irritator came "from irritation,
20952-477: The front; features that would have resulted in the eyes being positioned further up on the head than in other theropods. In particular, the broad lower jaw and narrowed frontals of Irritator resulted in the orbits facing at a steep incline towards the midline of the skull, whereas most theropods had laterally facing orbits. These traits would have permitted the animal to see above the waterline when submerged. In 2018, Aureliano and colleagues conducted an analysis on
21146-737: The generic name (or its abbreviated form) still forms the leading portion of the scientific name, for example, Canis lupus lupus for the Eurasian wolf subspecies, or as a botanical example, Hibiscus arnottianus ssp. immaculatus . Also, as visible in the above examples, the Latinised portions of the scientific names of genera and their included species (and infraspecies, where applicable) are, by convention, written in italics . The scientific names of virus species are descriptive, not binomial in form, and may or may not incorporate an indication of their containing genus; for example,
21340-542: The giant cashew trees, which are the majority in the Region, are exploited in an almost extractive manner, with low use of technology. In the production of cocoa , for a long time, Bahia led the Brazilian production. Today, it is disputing the leadership of national production with the state of Pará. In 2017 Pará obtained the leadership for the first time. In 2019, people from Pará harvested 135 thousand tons of cocoa, and Bahians harvested 130 thousand tons. Bahia's cocoa area
21534-416: The head before burial. Naish and colleagues in 2004 asserted that the Romualdo Formation dinosaur fauna is represented by animals that died near shorelines or rivers before being carried out to sea, where their floating remains were eventually fossilized. In 2018, Aureliano and colleagues argued against this scenario, stating that the Irritator challengeri holotype's mandible was preserved in articulation with
21728-456: The head body. In addition, the vestibular part of the endosseous labyrinth has a large anterior semicircular canal with a lateral oriented semicircular canal. Both floccular recesses and semicircular canal suggest that Irritator could coordinate fast head movements and had a downward inclined snout posture, enabling an unobstructed, stereoscopic forward vision, which is important for distance perception and therefore precise snatching movements of
21922-697: The head rapidly. In 2015, German paleontologist Serjoscha W. Evers and colleagues found evidence for similar adaptations in the African spinosaur Sigilmassasaurus . The neck vertebrae of this genus have a heavily furrowed undersurface. This is consistent with the attachment of powerful neck muscles for use in fishing or rapidly snatching small prey, a trait also observed in extant crocodilians and birds. Sales and Schultz in 2017 found that Irritator and baryonychines might have relied more on their sense of smell for hunting than Spinosaurus did, since they had larger, less retracted nostrils and more room in their skulls for
22116-568: The high rainfalls region of Amazon rainforest and the semi arid region of Sertão (hot and drought) covering the state of Maranhão and half of Piauí . The Northeast region comprises the drainage basins of the São Francisco , Canindé , and Parnaíba Rivers . Geographically, Nordeste consists chiefly of an eroded continental craton with many low hills and small ranges. The highest peaks are around 1,850 metres (6,070 ft) in Bahia, while further north there are no peaks above 1,123 metres (3,684 ft). On its northern and western side,
22310-401: The highest added value of Amazonian guarana. The pharmaceutical industries and importers, on the other hand, buy more guarana from Bahia, due to the price. In 2017, the Northeast had 12.9% of the Brazilian cattle herd. In the sertão, producers often suffer losses due to constant droughts. There are also goat breeds, which are more resistant, pigs, sheep and birds. Livestock fairs are common in
22504-552: The holotype of Angaturama seems to complete that of Irritator (meaning that they could belong to the same specimen). Authors including Éric Buffetaut and Mohamed Ouaja in 2002, Cristiano Dal Sasso and colleagues in 2005, Tor G. Bertin in 2010, Darren Naish in 2013, and Madani Benyoucef and colleagues in 2015 supported this conclusion. In their redescription of Irritator , Sues and colleagues pointed out that both holotypes are equally as narrow, and share transversely round teeth with defined yet unserrated edges. They also noted that
22698-435: The installed capacity for processing cashew nuts in the Northeast, however, the Region only managed to produce around a quarter of that quantity. Among the main world producers, Brazil has the lowest productivity. Several factors are pointed out as the cause of the low productivity and the fall in the Brazilian production of cashew nuts. One reason is that most orchards are in a phase of natural decline in production. In addition,
22892-645: The lagoon. Similarities between the fauna of the Romualdo and Crato Formations to that of Middle Cretaceous Africa suggest that the Araripe Basin was connected to the Tethys Sea , though this link was likely sporadic, because the lack of marine invertebrates indicates the basin had a non-marine depositional setting. Spinosaurids had already achieved a cosmopolitan distribution during the Early Cretaceous. Sereno and colleagues in 1998 suggested that with
23086-633: The largest component, with 23,236 ± 5,379 accepted genus names, of which 20,845 ± 4,494 are angiosperms (superclass Angiospermae). By comparison, the 2018 annual edition of the Catalogue of Life (estimated >90% complete, for extant species in the main) contains currently 175,363 "accepted" genus names for 1,744,204 living and 59,284 extinct species, also including genus names only (no species) for some groups. The number of species in genera varies considerably among taxonomic groups. For instance, among (non-avian) reptiles , which have about 1180 genera,
23280-404: The largest numbers of both species. Regarding pork, Brazil had almost 42 million hogs in 2017. The Northeast had 13% of the total (5.4 million). In poultry farming, Brazil had a total of 1.4 billion chickens in 2017. The Northeast had 11.6% of the total (164 million). In milk production, Brazil produced 33.5 billion liters in 2017. The Northeast produced 11.6% of the total (3.9 billion liters). In
23474-404: The left maxilla's tooth crowns are more intact, and there are traces of a tenth tooth in the rock matrix. The teeth were deeply inserted into the jaw and widely spaced towards the front of the maxilla. The first and second preserved maxillary teeth were the largest, at 32 mm (1.3 in) and 40 mm (1.6 in) in crown length. The seven remaining teeth became progressively smaller towards
23668-402: The lizard genus Anolis has been suggested to be broken down into 8 or so different genera which would bring its ~400 species to smaller, more manageable subsets. Northeastern Brazil The Northeast Region of Brazil ( Portuguese : Região Nordeste do Brasil [ʁeʒiˈɐ̃w nɔʁˈdɛstʃi du bɾaˈziw] ) is one of the five official and political regions of the country according to
23862-422: The lower jaws of this spinosaurid could rotate and open laterally, with a gape similar to that of modern pelicans; this would have allowed the theropod to swallow very large prey items. It was also discovered to have a very weak but rapid bite as a result of this analysis. The study also reaffirms that Irritator likely possessed binocular vision and would have held its snout at an inclined orientation. Additionally,
24056-509: The lush aquatic life as a primary food source. They also hypothesized that following storm events, pterosaur and fish carcasses might have washed up on the shoreline, providing theropods with plenty of carrion. Multiple piscivorous animals were present in the formation, which might in theory have led to high competition. Aureliano and colleagues stated there must have, therefore, been some degree of niche partitioning , where different animals would have fed on prey of varied sizes and locations within
24250-401: The more smoothly sloping snouts of other spinosaurs. In the premaxilla, a broken-off tooth with partial tooth crown was recovered. The strongly extended and straight teeth with unserrated conical crowns, which measured 6 to 40 mm (0.24 to 1.57 in) in length, were singly embedded. This indicates continuous tooth replacement where new teeth were pushed up between the old ones. Judging by
24444-403: The most (>300) have only 1 species, ~360 have between 2 and 4 species, 260 have 5–10 species, ~200 have 11–50 species, and only 27 genera have more than 50 species. However, some insect genera such as the bee genera Lasioglossum and Andrena have over 1000 species each. The largest flowering plant genus, Astragalus , contains over 3,000 species. Which species are assigned to a genus
24638-428: The name could not be used for both. Johann Friedrich Blumenbach published the replacement name Ornithorhynchus in 1800. However, a genus in one kingdom is allowed to bear a scientific name that is in use as a generic name (or the name of a taxon in another rank) in a kingdom that is governed by a different nomenclature code. Names with the same form but applying to different taxa are called "homonyms". Although this
24832-434: The north, the Sertão extends to the northern coastal plains of Rio Grande do Norte state, whilst in the south it fades out in the northern fringe of Minas Gerais . Because the Sertão lies close to the equator, temperatures remain nearly uniform throughout the year and are typically tropical, often extremely hot in the west. However, the sertão is distinctive in its low rainfall compared to other areas of Brazil. Because of
25026-486: The novelist himself. When Martill and colleagues first described Irritator challengeri , the holotype was still extensively encased in calcareous rock matrix . Under the supervision of American paleontologist Hans-Dieter Sues, technician Diane M. Scott from the University of Toronto at Mississauga assumed the task of fully extracting the skull bones from the rock, allowing for a detailed redescription in 2002. Published by Sues, Frey, Martill, and Scott, this inspection of
25220-403: The now fully prepared specimen negated many of Martill and colleagues' original observations, which were based on misinterpretations of the damaged and largely concealed skull. The estimated length of the complete skull was 24 cm (9.4 in) shorter than previously proposed. What was originally thought to be a prominent head crest proved to be an unattached, indeterminate bone fragment. As in
25414-547: The number of 12 teeth in MSNM V4047, an upper jaw fossil referred to Spinosaurus . The hindmost tooth of the Irritator specimen's left maxilla was not yet fully erupted, and only the tip of it was visible. The holotype of Angaturama limai consists only of the front part of the upper jaw, comprising the tips of the paired premaxillae and the frontmost ends of the maxillae. The specimen measures 19.2 cm (7.6 in) in height and 11 cm (4.3 in) in length, with
25608-444: The one autapomorphy (distinguishing feature) diagnosed in maniraptorans at the time, which was having its jugal (cheek) bone forming part of the antorbital fenestra. He also pointed out that since Irritator challengeri ' s holotype lacked the tip of its snout, it would not be possible to know if Spinosaurus ' s dentary could complement it or not. Based on comparisons with Spinosaurus , Kellner resolved Irritator as
25802-592: The one which gives it its name. The largest state of the Northeast region in terms of area, population and economic output is Bahia; its capital Salvador is the largest city of Nordeste . In 2016, the Northeast was in 3rd place among the 5 regions of Brazil, in terms of total wealth. The Southeast comes in 1st with 53.1% of GDP, the South in 2nd with 17%, Northeast in 3rd with 14.4% and the Center-West in 4th with 10.2%. In GDP per capita (that is, GDP per inhabitant), it
25996-643: The opening of the Tethys Sea, spinosaurines would have evolved in the south (Africa, in Gondwana ) and baryonychines in the north (Europe, in Laurasia ). In following, Machado and Kellner theorized in 2005 that spinosaurines would have then spread to South America from Africa. Sereno and colleagues postulated that divergent evolution between spinosaurines in South America and Africa likely occurred as
26190-497: The original postcranial elements (including the fossil pelvis and sacral vertebrae) were presented alongside the mount. In press releases of the exhibit's opening, Kellner informally implied MN 4819-V as belonging to Angaturama . This is also reflected in the specimen's inclusion in the skeletal mount. In 2011, a third Brazilian spinosaur, Oxalaia quilombensis , was named and described from the Alcântara Formation of
26384-594: The plateaus fall steadily to the coast and into the basin of the Tocantins River in Maranhão, but on the eastern side it falls off quite sharply to the coast except in the valley of the São Francisco river. The steep slopes and long cliffs of the eastern coastline are known as " The Great Escarpment ". The escarpment serves an extremely important climatic function because for most of the year, Nordeste
26578-512: The population in 1775 to 27.5% in 1855. Between 1877 and 1879, a large part of the sertão was hit by the most severe drought in Brazilian history, known as the Great Drought , with Ceará being the hardest hit region. As a result, thousands of Nordestinos migrated to other parts of Brazil, especially to the Amazon , and about 200,000 people died. Before the rise of Sudeste , Nordeste
26772-418: The premaxilla. In 2005, Dal Sasso and colleagues assumed Irritator ' s nostrils as being located above the middle of the maxillary tooth row; more posteriorly than in baryonychines, but less so than in Spinosaurus . Sales and Schultz in 2017 found that Irritator ' s nostrils were in fact positioned closer to the front of the jaw, as in Baryonyx and Suchomimus ; this more forward nostril placement
26966-455: The premaxillae would have complemented a convex and enlarged mandible tip. The premaxillae connected with each other on the bottom to form Angaturama ' s secondary palate, which was also partially contributed to by two processes extending from the maxillae. The snout was strongly compressed laterally, and the premaxillae gently tapered towards the top to form a tall sagittal crest 1 to 2 mm (0.039 to 0.079 in) in thickness. This crest
27160-454: The presence of: hybodont sharks , guitarfish , gars , amiids , ophiopsids , oshuniids , pycnodontids , aspidorhynchids , cladocyclids , bonefishes , chanids , mawsoniids and some uncertain forms. According to Naish and colleagues, the lack of herbivorous dinosaurs could mean that the local vegetation was scant and thus incapable of sustaining a large population of them. The abundant carnivorous theropods would have then likely turned to
27354-435: The previous study, Sues and colleagues regarded the African and possibly South American genus Spinosaurus as the most similar taxon to Irritator , because they shared many dental features, including mostly straight conical tooth crowns , thin enamel , well-defined edges with no serrations , and lengthwise fluting. Since little was known of Spinosaurus ' s skull at the time, these similarities were enough for
27548-423: The production of eggs, Brazil produced 4.2 billion dozens in 2017. The Northeast produced 16.1% (683 million dozens). The Northeast was the 2nd largest honey producer in the country in 2017, losing to the South region. The total produced in the country was 41.6 thousand tons. The Northeast produced 30.7% (12.7 thousand tons). In 2017, the Northeast was the largest shrimp producer in the country. National production
27742-541: The provisions of the ICZN Code, e.g., incorrect original or subsequent spellings, names published only in a thesis, and generic names published after 1930 with no type species indicated. According to "Glossary" section of the zoological Code, suppressed names (per published "Opinions" of the International Commission of Zoological Nomenclature) remain available but cannot be used as the valid name for
27936-413: The rear part of the bone that allowed for the passage of the obturator nerve . The upwards projecting neural spines of the sacrum were elongated, as is typical in spinosaurs. In life, these would have been covered in skin, forming a " sail " down the animal's back. MN 4819-V is distinguished from Suchomimus due to its longer and shallower ilium with a less curved upper margin, and from Baryonyx by having
28130-435: The rear, one of the last ones measuring 6 mm (0.24 in) in estimated crown length. CT scans performed on the specimen revealed replacement teeth on both sides of the upper jaw. Their roots ran deep into the maxillae and converged close to the midline, nearly reaching the top of the skull. Based on comparisons with Irritator ' s relatives, the maxillae were probably lined with a total of 11 teeth each, similar to
28324-757: The red dye it produced. Countries like France , which disagreed with the Treaty of Tordesillas (a papal bull decreed by the Spanish-born Pope Alexander VI in 1493 which sought to divide South America between the Spanish and the Portuguese), launched many attacks on the coast to steal the wood. Soon after their arrival, Portuguese settlers began to displace native peoples and enslave them as field laborers, leading to conflicts in which many natives died. These conflicts were one contributor to
28518-503: The region has the second largest number of passengers (roughly 20%) in Brazil. The Zona da Mata comprises the rainforest zones of Nordeste (part of the Atlantic Forest or Mata Atlântica ) in the humid eastern coast, where the region's largest capital cities are also located. The forest area was much larger before suffering from centuries of deforestation and exploration. For many years, sugar cane cultivation in this region
28712-517: The relatively cool temperatures in the South Atlantic Ocean , the Intertropical Convergence Zone remains north of the region for most of the year. Consequently, conditions are very dry for most of the year. Although annual rainfall averages between 500 and 800 millimeters over most of the sertão and 1300 millimeters on the northern coast of Fortaleza , it is confined to a short rainy season. This season extends from January to April in
28906-541: The researchers erected the new family Irritatoridae within the clade. They recognized Irritator ' s affinities with Spinosaurus , in that they both had similarly shaped and unserrated teeth, but noted that the latter's mandible would not conform with the front of Irritator ' s upper jaw, and that other non-avian dinosaurs like Compsognathus and Ornitholestes also bore no serrations on some or all of their teeth. Some of these claims were questioned in 1996 by Kellner who found that Irritator ' s skull lacked
29100-617: The rest of the skull, whereas it would have likely detached had the carcass been floating at sea. They also noted that the corpse would have quickly sunk due to the osteosclerosis of the skeleton. The researchers, therefore, concluded that fossils from the Santana Group represent organisms that were buried in their natural habitat, instead of having been deposited allochthonously (other than at their present position). [REDACTED] [REDACTED] [REDACTED] [REDACTED] [REDACTED] Genus The composition of
29294-421: The right side better articulated than the left. The fused sacral vertebrae are still attached to the pelvis, which lacks the distal ends of both of its pubic bones and ischia (lower and rearmost hip bones). The ilium (main hip bone) is 55.3 cm (21.8 in) long. The preacetabular ala (front expansion) of the ilium was curved on the bottom and was somewhat shorter and deeper than
29488-484: The same animal or not. Other spinosaurid skeletal material, some of which could belong to Irritator or Angaturama , was retrieved from the Romualdo Formation, allowing for a replica skeleton to be made and mounted for display at the National Museum of Rio de Janeiro in 2009. Estimated at between 6 and 8 meters (20 and 26 ft) in length, Irritator weighed around 1 tonne (1.1 short tons), making it one of
29682-470: The same kind as other (analogous) genera. The term "genus" comes from Latin genus , a noun form cognate with gignere ('to bear; to give birth to'). The Swedish taxonomist Carl Linnaeus popularized its use in his 1753 Species Plantarum , but the French botanist Joseph Pitton de Tournefort (1656–1708) is considered "the founder of the modern concept of genera". The scientific name (or
29876-488: The same time and place as Irritator challengeri , was described by the Brazilian paleontologists Alexander W. A. Kellner and Diogenes de Almeida Campos in February 1996. Kept today under specimen number USP GP/2T-5 at the University of São Paulo , the holotype specimen consists of an isolated snout tip from the Romualdo Formation. It was extracted from a calcareous nodule using a technique originally developed for pterosaur fossils. The generic name Angaturama means "noble" in
30070-408: The scientific epithet) of a genus is also called the generic name ; in modern style guides and science, it is always capitalised. It plays a fundamental role in binomial nomenclature , the system of naming organisms , where it is combined with the scientific name of a species : see Botanical name and Specific name (zoology) . The rules for the scientific names of organisms are laid down in
30264-428: The semicircular canals and connects the brain with the inner ear). The flocculus , itself, is an important element in the coordination and control of head and ocular movements during gaze stabilization (visual ability during head movement), by being involved in the coordination of the vestibulo-ocular (VOR) and vestibulo-collic (VCR) reflexes. The flocculus appears to be enlarged in taxa that rely on quick movements of
30458-410: The skull roof (called the postnasal fenestrae) as well as long and only partially diverging basipterygoid processes (bony extensions connecting the braincase with the palate ). The back of the lower jaw was deep, its rear upper surface consisting mainly of the large surangular bone, which articulated with the shallower angular bone below it. The mandibular fenestra ,
30652-463: The skull than in typical theropods. Irritator ' s nostrils were both proportionately and absolutely smaller than in Suchomimus and Baryonyx , but larger than those of Spinosaurus . The opening behind the orbit (eye socket), the lateral temporal fenestra , was very large, while the antorbital fenestra , in front of the eyes, was long and elliptical. The orbit itself was deep and wider at
30846-409: The smallest spinosaurids known. Its long, shallow and slender snout was lined with straight and unserrated conical teeth. Lengthwise atop the head ran a thin sagittal crest , to which powerful neck muscles were likely anchored. The nostrils were positioned far back from the tip of the snout, and a rigid secondary palate on the roof of the mouth would have strengthened the jaw when feeding. Belonging to
31040-401: The snout tip; this, along with the secondary palate, which separated the animal's nasal passages from the inside of its mouth, made respiration possible even if most of the jaw was underwater or held prey. In particular, the sagittal crest of Irritator is an indication for pronounced neck musculature, which would have been necessary to close the jaws quickly against water resistance and withdraw
31234-496: The snout. These inferences seem to be an agreement with a piscivore life-style. They also noted that the relatively well-developed cochlear duct may have enable an average hearing frequency of 1,950 Hz with a frequency band width of 3,196 Hz. However, they considered these ranges as rough estimates and established an overall frequency range of 350–3,550 Hz, making Irritator to be placed under bird hearing but above crocodiles. Irritator and Angaturama are known from
31428-497: The specific name particular to the wolf. A botanical example would be Hibiscus arnottianus , a particular species of the genus Hibiscus native to Hawaii. The specific name is written in lower-case and may be followed by subspecies names in zoology or a variety of infraspecific names in botany . When the generic name is already known from context, it may be shortened to its initial letter, for example, C. lupus in place of Canis lupus . Where species are further subdivided,
31622-432: The specimen became the holotype of the type species Irritator challengeri . The genus name comes from the word "irritation", reflecting the feelings of paleontologists who found the skull had been heavily damaged and altered by the collectors. The species name is a homage to the fictional character Professor Challenger from Arthur Conan Doyle 's novels. Some paleontologists regard Angaturama limai —known from
31816-412: The standard format for a species name comprises the generic name, indicating the genus to which the species belongs, followed by the specific epithet, which (within that genus) is unique to the species. For example, the gray wolf 's scientific name is Canis lupus , with Canis ( Latin for 'dog') being the generic name shared by the wolf's close relatives and lupus (Latin for 'wolf') being
32010-444: The struggle against the Portuguese. At the end of 1653, the Portuguese succeeded in capturing Recife, effectively ending Dutch Brazil and culminating in their surrender in 1654. Slave resistance began during the colonial era, in the 17th century, and eventually led to the formation of quilombos , or settlements of runaway and free-born African slaves. The Quilombo dos Palmares , the largest and most well-known of these settlements,
32204-403: The taxon is termed a synonym ; some authors also include unavailable names in lists of synonyms as well as available names, such as misspellings, names previously published without fulfilling all of the requirements of the relevant nomenclatural code, and rejected or suppressed names. A particular genus name may have zero to many synonyms, the latter case generally if the genus has been known for
32398-471: The thinner frontal bones, situated behind them, were smooth and concave on top; both of these bones formed the upper rim of the orbit. A thin sagittal crest , constructed from the elongate nasal bones, extended along the skull midline before stopping just above the eye in a slightly flattened bulge. Although the complete shape and height of this structure is unknown in Irritator , these head crests were commonplace in spinosaurids, having possibly served
32592-723: The time Irritator was described, was thought to be a member of what was then considered the Santana Formation. The Romualdo Formation is a Lagerstätte (a sedimentary deposit that preserves fossils in excellent condition) consisting of limestone concretions embedded in shales , and overlies the Crato Formation. It is well known for preserving fossils three-dimensionally in limestone concretions, including many pterosaur fossils. In addition to muscle fibres of pterosaurs and dinosaurs, fish, preserving gills, digestive tracts, and hearts, have been found there. The formation
32786-427: The top (where the eyeball was placed) than the bottom. The lacrimal bone separated the orbit from the antorbital fenestra, forming the upper and lower rear margins of the latter with two processes that enclosed a 40-degree angle; similar to Baryonyx , where it enclosed 35 degrees. Unlike in Baryonyx , Irritator ' s lacrimal did not form a bony horn core. The prefrontal bones were large and sturdy, while
32980-414: The total in the region (10.3 million tons). Paraíba has 11.9% of northeastern production (5.5 million tons) and Bahia, 10.24% of production (4.7 million tons). Bahia is the 2nd largest producer of cotton in Brazil, losing only to Mato Grosso. In 2019, it harvested 1.5 million tonnes of the product. In soy , Brazil produced close to 120 million tons in 2019, being the largest world producer. In 2019,
33174-399: The total volume of nuts produced in the world. Vietnam, Nigeria, India and Côte d'Ivoire were the world's largest cashew nut producers in 2016, with 70.6% of global production. In recent years, there has been increased competition with some African countries, where government programs have driven the expansion of culture and processing capacity. It is estimated that at 295 thousand tons per year
33368-576: The values quoted are the mean of "accepted" names alone (all "uncertain" names treated as unaccepted) and "accepted + uncertain" names (all "uncertain" names treated as accepted), with the associated range of uncertainty indicating these two extremes. Within Animalia, the largest phylum is Arthropoda , with 151,697 ± 33,160 accepted genus names, of which 114,387 ± 27,654 are insects (class Insecta). Within Plantae, Tracheophyta (vascular plants) make up
33562-429: The virus species " Salmonid herpesvirus 1 ", " Salmonid herpesvirus 2 " and " Salmonid herpesvirus 3 " are all within the genus Salmonivirus ; however, the genus to which the species with the formal names " Everglades virus " and " Ross River virus " are assigned is Alphavirus . As with scientific names at other ranks, in all groups other than viruses, names of genera may be cited with their authorities, typically in
33756-402: The west, but in the eastern Sertão it generally occurs from March to June. However, rainfall is extremely erratic and in some years the rains are minimal, leading to catastrophic drought . The duration of droughts has increased over the last 36 years, with the 2012–2016 drought being the longest drought in the region's history. Because of this vulnerability to the climate, Sertão is also known as
33950-445: The width of the palatal region being 4 to 5 mm (0.16 to 0.20 in). The suture between the maxilla and premaxilla was jagged at the front and straightened out towards the rear. The lower margin of the premaxillae was concave, with the concavity reaching its apex at the sixth premaxillary tooth. The front of the snout was expanded, forming the spoon-shaped terminal rosette characteristic of spinosaurids. This concave underside of
34144-413: The word sertão ( Portuguese pronunciation: [seʁˈtɐ̃w] , meaning " backcountry " or " outback ") first referred to the vast hinterlands of Asia and South America that Lusitanian explorers encountered. In Brazil, the geographical term referred to backlands away from the Atlantic coastal regions where the Portuguese first settled in South America in the early sixteenth century. Geographically,
34338-471: Was 41 thousand tons. Rio Grande do Norte (37.7%) and Ceará (28.9%) were the largest producers. Aracati-CE was the municipality with the highest participation . In Northeast Region, Bahia stands out, with 1.68% of the national mineral participation (4th place in the country). In 2017, at gold , it produced 6.2 tons, at a value of R$ 730 million. At copper , it produced 56 thousand tons, at a value of R$ 404 million. At chrome , it produced 520 thousand tons, at
34532-436: Was 7th with 166 thousand tons. Bahia is the second largest fruit producer in the country, with more than 3.3 million tons a year, behind São Paulo. The north of Bahia is one of the main fruit suppliers in the country. The State is one of the main national producers of ten types of fruit. In 2017, Bahia led the production of cajarana, coconut, count fruit or pinecone, soursop, umbu, jackfruit, licuri, mango and passion fruit, and
34726-646: Was already rare. Salvador remained the colonial capital until 1763 when it was succeeded by Rio de Janeiro , the new economic power center of that era. In 1850, the Eusébio de Queirós Law was passed, abolishing the international slave trade in Brazil. Following the ban, some slaves from Nordeste were sold to the Southeast region of Brazil ( Portuguese : Sudeste ), primarily to the state of Rio de Janeiro . The percentage of slaves in Salvador dropped from 41.6% of
34920-407: Was also present in the Angaturama limai holotype. In 2002, Sues and colleagues pointed out, however, that there would be no reason to assume that the Spinosauridae specialized completely in fishing. They stressed rather that this head morphology indicates a generalistic feeding , particularly on small prey animals. In fact, portions of a young Iguanodon , a terrestrial herbivore, were found inside
35114-734: Was destroyed in the War of Canudos (1896–1897), and the Cangaceiros in the 1930s. During the administration of Juscelino Kubitschek (1956–61), the Superintendency for the Development of the Northeast , or Sudene , was created with the goal of promoting the development of the Northeast region. In the following decades, under the Populist Republic and the military dictatorship , the investments made by Sudene catalyzed
35308-416: Was founded around 1600 in the Serra da Barriga hills, in the present state of Alagoas . Palmares, at the height of its power, was an independent, self-sustaining republic, hosting a population of over 30,000 free African men, women and children. There were over 200 buildings in the community, a church, four smithies, and a council house. Although Palmares managed to defend itself from the Dutch military and
35502-401: Was in 3rd place among the regions that most produce sugar cane in the country. Brazil is the world's largest producer, with 672.8 million tons harvested this year. The Northeast harvested 45.7 million tons, 6.8% of national production. Alagoas is the largest producer, with 33.3% of Northeastern production (15.2 million tons). Pernambuco is the 2nd largest producer in the Northeast, with 22.7% of
35696-493: Was in 3rd place and Rio Grande do Norte in 4th place. Bahia was the largest producer of mango in the country in 2019, with production of around 281 thousand tons per year. Juazeiro (130 thousand tons per year) and Casa Nova (54 thousand tons per year) are at the top of the list of Brazilian cities that lead the cultivation of fruit. In the production of banana , in 2018 Bahia was the 2nd largest national producer. Pernambuco came in 5th place. Regarding pineapple , in 2018 Paraíba
35890-476: Was inhabited by indigenous peoples, mostly speaking languages of the Tupi–Guarani family. In the Sertão region, Tapuia tribes could also be found. It was the first area of Brazil to be colonized when roughly 1,500 Portuguese arrived on April 22, 1500, under the command of Pedro Álvares Cabral at Porto Seguro , in what is now the state of Bahia . The coast of Nordeste saw the first economic activity of
36084-451: Was larger and extended farther forwards on the snout than in other known spinosaurids. The frontmost upper border of the premaxilla had a small bulge that overhung the base of the crest. This bulge was apparently damaged on its upper surface, indicating that the top of the crest may have extended even farther over and forwards from that point. The front of Angaturama ' s snout hence had a vertically straight or concave margin, atypical from
36278-399: Was long, narrow, and somewhat triangular in cross section. The braincase was inclined backwards, and deeper than it was long. Extending from it was an elongate and low snout, with both sides relatively flat and slightly angled towards the skull midline. Only the rear ends of the paired premaxillae (frontmost snout bones) remain intact, forming the front upper and lower borders of
36472-488: Was present between the last premaxillary and first maxillary tooth. Though no skeletal remains were discovered with the original Angaturama snout tip, one partial skeleton (MN 4819-V) from a different location may belong to the genus. But since there is no overlapping material between the two specimens, direct comparisons cannot be made. MN 4819-V comprises a largely intact pelvis , some dorsal (back) and caudal (tail) vertebrae , five sacral (hip) vertebrae,
36666-410: Was referred by Kellner to the Spinosauridae in 2001 because of its tall sacral neural spines and the enlarged condition of the hand claw . The skeleton was fully described in 2010 in an as-of-yet unpublished master's thesis by Machado. An incomplete hind limb (MPSC R-2089) mentioned in 2013 might also pertain to the Spinosauridae. In 2018 , Tito Aureliano and his team described LPP-PV-0042, part of
36860-499: Was represented by other theropods like the tyrannosauroid Santanaraptor , the compsognathid Mirischia , an indeterminate unenlagiine dromaeosaurid , and a megaraptoran . The crocodyliforms Araripesuchus and Caririsuchus , as well as the turtles Brasilemys , Cearachelys , Araripemys , Euraxemys , and Santanachelys , are known from the deposits. There were also clam shrimps , sea urchins , ostracods, and molluscs . Various well-preserved fish fossils record
37054-475: Was the 2nd largest producer state in Brazil. Bahia is the largest Brazilian producer of guaraná . In 2017, Brazilian production was close to 3.3 million tons. Bahia harvested 2.3 million (mainly in the city of Taperoá), Amazonas 0.7 million (mainly in the city of Maués) and the rest of the country, 0.3 million. Despite the fact that the fruit originated in the Amazon, since 1989 Bahia has beaten Amazonas in terms of production volume and guarana productivity, due to
37248-480: Was the center of the African slave trade in Brazil, the center of the sugar industry in Brazil, and Brazil's main seaport. During the Old Republic, the interior of the Northeast was under the dominion of local oligarchs called coronéis , who enforced their power through violence and fear. Two notable examples of responses to this oppression were the messianic community of Canudos, in the interior of Bahia, which
37442-419: Was the mainstay of Brazil's economy , being superseded only when coffee production developed in the late 19th century. Sugar cane is cultivated on large estates whose owners maintain tremendous political influence. Since the escarpment does not generate any further rainfall on its slopes from the lifting of the trade winds , annual rainfall decreases steadily inland. After a relatively short distance, there
37636-446: Was typically considered characteristic of baryonychines. Nevertheless, Irritator also bore unserrated teeth, a trait associated with spinosaurines. Sales and Schultz thus noted that the Araripe Basin spinosaurids Irritator and Angaturama might represent intermediate forms between the earlier baryonychines and later spinosaurines, and that further research may eventually render the former a paraphyletic (unnatural) group. Irritator
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