title: NXT TakeOver: In Your House (2021)

text: TakeOver was a series of professional wrestling shows that began in May 2014, as WWE's NXT brand held their second WWE Network-exclusive event, billed as TakeOver. In subsequent months, the "TakeOver" moniker became the brand used by WWE for all of their NXT live specials. While originally exclusive to the WWE Network, NXT TakeOver events also became available on traditional pay-per-view beginning with TakeOver 31 in October 2020, and then also on Peacock after the American version of the WWE Network merged under Peacock in March 2021. The 2021 TakeOver: In Your House was scheduled to be held on June 13, 2021, as the 35th NXT TakeOver event, as well as the second in the TakeOver: In Your House chronology, a subseries of TakeOvers that carry the In Your House name.


title: NXT TakeOver: In Your House (2021)

text: In Your House was a series of monthly PPV shows that was held by the then-World Wrestling Federation (WWF, renamed to WWE in 2002) from May 1995 to February 1999. They aired when the promotion was not holding one of its then-five major PPVs (WrestleMania, King of the Ring, SummerSlam, Survivor Series, and Royal Rumble), and were sold at a lower cost. The branding was retired following February 1999's event, as the company moved to install permanent names for each of its monthly PPVs. In Your House was as an NXT TakeOver event, thus the 2021 event was the 29th event to carry the In Your House name. It was also the first In Your House to air on traditional pay-per-view since the February 1999 event as well as the first In Your House to air on Peacock—the 2020 TakeOver: In Your House only aired on the WWE Network.


title: NXT TakeOver: In Your House (2021)

text: As a result of the COVID-19 pandemic that began affecting the industry in mid-March 2020, WWE had to present the majority of its programming from a behind closed doors set. NXT's programming was initially held at NXT's home base of Full Sail University in Winter Park, Florida. In October 2020, NXT's events were moved to the WWE Performance Center in Orlando, Florida, featuring the "Capitol Wrestling Center" setup, an homage to the Capitol Wrestling Corporation, the predecessor to WWE. Like the WWE ThunderDome utilized for Raw and SmackDown's programming, LED boards were placed around the Performance Center so that fans could attend virtually, while additionally, friends and family members of the wrestlers were in attendance along with a limited number of actual live fans divided from each other by plexiglass walls.


title: NXT TakeOver: In Your House (2021)

text: For in April 2021, the plexiglass walls were removed, and live audience capacity was increased. In Your House increased capacity further to nearly 300 spectators, being the largest attended NXT event at the Capitol Wrestling Center. Nearly all COVID-19 protocols were lifted including physical distancing requirements and the requirement to wear masks although anyone who had tested positive, within the preceding 14 days, were asked to stay home. The virtual audience was also removed with the increased live audience capacity.


title: NXT TakeOver: In Your House (2021)

text: The In Your House card included five matches. The storylines were produced on their weekly television program, "NXT".


title: NXT TakeOver: In Your House (2021)

text: A triple threat match between Kyle O'Reilly, Pete Dunne, and Johnny Gargano was scheduled on the June 1 episode of "NXT" to determine who would face Karrion Kross for the NXT Championship at TakeOver: In Your House. The match ended in a no-contest after Adam Cole interfered and attacked all three competitors. Kross would confront Cole and made a request to NXT General Manager William Regal to defend the title against Cole, O'Reilly, Dunne, and Gargano. Regal agreed to Kross' request and made the bout a fatal five-way match for the title at the event.


title: NXT TakeOver: In Your House (2021)

text: González viciously assaulted Blackheart while Moon was forced to watch the ordeal after being subdued by Kai, following the conclusion of the women's tag team match between the team of Shotzi Blackheart and Ember Moon and the team of Dakota Kai and NXT Women's Champion Raquel González on the May 25 episode of "NXT". The following week, Moon called out González and they both brawled in the ring. Later that same night, Moon announced that she would challenge González for the NXT Women's Championship at TakeOver: In Your House, which was made official.


title: NXT TakeOver: In Your House (2021)

text: On the May 25 episode of "NXT", Mercedes Martinez defeated Zayda Ramier. Afterwards that day, the lights went out and turned back on with a red lighting, and Mei Ying of Tian Sha appeared on the LED screen. As things went back to normal Martinez was given a black symbol on the back of her wrist. The following week, Xia Li was with Boa watching a video of her first round loss against Martinez in the inaugural Mae Young Classic (four years ago) saying it was a disgrace to her family and she would hurt her at TakeOver: In Your House. Later that night, Martinez said that she was aware of Li being different, after facing her in the Mae Young Classic, and that she was not afraid of Li and would defeat her at TakeOver.


title: NXT TakeOver: In Your House (2021)

text: On the February 10 episode of "NXT", Cameron Grimes made his return from injury and claimed to have become a GameStop investor during his time away (in reference to the GameStop stock rise), thus making him the "richest man in NXT". Grimes then began a feud with WWE Hall of Famer Ted DiBiase (The Million Dollar Man) after encountering him in a jewelry store over their watches. Over the next few weeks, DiBiase would continue to one-up Grimes, outbidding him in various purchases. On the May 25 episode of "NXT", LA Knight confronted Grimes during the Million Dollar Face-Off segment between Grimes and DiBiase, the latter of whom Knight defended and allied with. The following week, it was announced that a singles match between Grimes and Knight was scheduled for TakeOver: In Your House. DiBiase later announced that their match would instead be a ladder match for the revived Million Dollar Championship.


title: NXT TakeOver: In Your House (2021)

text: On the June 1 episode of "NXT", MSK (Nash Carter and Wes Lee) successfully defended their NXT Tag Team Championship against Legado Del Fantasma (Joaquin Wilde and Raúl Mendoza), after an assist from NXT North American Champion Bronson Reed, who took down Santos Escobar during the match. The following week, Escobar proposed a winner takes all match for the NXT North American Championship and NXT Tag Team Championships at TakeOver: In Your House, which Reed and MSK accepted.


title: NXT TakeOver: In Your House (2021)

text: The pay-per-view opened with NXT North American Champion Bronson Reed and NXT Tag Team Champions MSK (Wes Lee and Nash Carter) facing off against Legado del Fantasma (Santos Escobar, Joaquin Wilde, and Raul Mendoza) in a Winner Takes All Six-man Tag Team match. During the match, Wilde and Mendoza performed a High-Low on Lee for a nearfall, before Carter broke it up. Escobar performed a Michinoku Driver on Reed for a nearfall. In the climax, after Reed tackled Escobar against one of the plexiglass walls, MSK performed their elevated corkscrew blockbuster on Mendoza, followed by Reed performing the "Tsunami" on Mendoza to retain their respective championships.


title: NXT TakeOver: In Your House (2021)

text: Next, Xia Li (accompanied by Boa) faced Mercedes Martinez. In the end, Li performed a roundhouse kick to Martinez's head to win the match. Following the pinfall, Martinez attacked both Li and Boa with a steel chair sending them out of the ring. As the duo retreated up the entrance ramp, Mei Ying, who watched and walked on stage, took Martinez down. As Martinez attempted to attack Ying, Ying applied a choke hold to immobilize Martinez, throwing her off the ramp into a guardrail.


title: NXT TakeOver: In Your House (2021)

text: In a backstage segment, Johnny Gargano and Kyle O'Reilly, two of the five participants in the main event match, were shown brawling in the arena parking lot.


title: NXT TakeOver: In Your House (2021)

text: After that, LA Knight faced Cameron Grimes in a ladder match for the Million Dollar Championship. Ted DiBiase watched the match at ringside — this was a back-and-forth match. During the match, Grimes retrieved the gold-plated ladder that DiBiase had used to introduce the match (which was on the stage). As Grimes ascended the ladder, Knight prevented Grimes from retrieving the Championship by tipping the ladder over. In the closing moments, Grimes attempted to ascend the ladder again, only for Knight to send Grimes off the ladder, through another ladder which was positioned outside the ring. Knight then ascended and retrieved the championship to win the match. Following the match, DiBiase awarded the championship to Knight.


title: NXT TakeOver: In Your House (2021)

text: In the fourth match, Raquel Gonzalez (accompanied by Dakota Kai) defended the NXT Women's Championship against Ember Moon. During the match, Moon performed her "Eclipse" on Gonzalez, for only Kai to place Gonzalez's foot on the bottom rope to void the pin. This prompted Moon's tag team partner Shotzi Blackheart, who was not seen on NXT due to injury, to come out and brawl with Kai to the backstage area. Gonzalez then performed the "Chingona Bomb" on Moon to retain the championship. Later, Kai and Blackheart were shown still brawling in the backstage area.


title: NXT TakeOver: In Your House (2021)

text: In the main event, Karrion Kross (accompanied by Scarlett) defended the NXT Championship against Johnny Gargano, Pete Dunne, Kyle O'Reilly, and Adam Cole in a fatal five-way match. During the match, Dunne, Gargano, O'Reilly, and Cole attempted to take out Kross, however Kross dispatched all the challengers. Dunne and Gargano threw Kross into the house door of the In Your House set at the top of the stage, temporarily taking Kross out. Kross later returned, however O'Reilly and Cole performed a powerbomb on Kross onto the announcer's table, which did not break. Dunne performed the "Bitter End" on Kross for a nearfall, Gargano performed a "One Final Beat" on Dunne, and Cole performed a "Panama City Sunrise" on Gargano. In the end, Kross applied the "Kross Jacket" on O'Reilly, who passed out, to retain the title.


title: NXT TakeOver: In Your House (2021)

text: Following the match, NXT general manager William Regal, who looked sullen, told an interviewer that in view of the wild brawls, he was unable to keep order in NXT anymore and mused that it was time for a change.


title: NXT TakeOver: In Your House (2021)

text: TakeOver: In Your House generally received positive reviews from critics. Wrestling journalist Dave Meltzer awarded 4 stars to the six-man tag team match, 2.25 stars to the Xia Li/Mercedes Martinez match, 3.75 stars to the ladder match, 3 stars to the women's championship match, and 4.25 stars to the main event. The event as a whole received a 7.5 "Good" rating from 411mania's Kevin Pantoja. Fightful.com's Sean Ross Sapp gave each match generally positive ratings as well, with the main event receiving the highest score at 8.5/10 and the Li/Martinez match the lowest at 5.75/10.


title: NXT TakeOver: In Your House (2021)

text: William Regal opened the June 15 episode of "NXT" by talking about his history with the brand, tearfully recounting the many roles he had had with NXT since 2014. As he was about to reach a conclusion, he was interrupted by NXT Champion Karrion Kross, who gloated about his victory at TakeOver and mockingly inviting Regal to leave. Samoa Joe then made his return to NXT (and to WWE as a whole, having been released from his contract two months prior). Regal revealed that he planned to make Joe the new general manager of NXT, and then formally extended the offer. Joe declined, but made a counteroffer to become Regal's enforcer if Regal stayed in the role. Regal said he would accept on two conditions — that Joe would not be an active in-ring competitor and that he could not physically engage anyone unless provoked. Joe agreed, and began filling the role in segments throughout that night. At the end of the show, he and Regal shook hands on a successful night one of their partnership, with Joe stating it would be "night one of many."


title: NXT TakeOver: In Your House (2021)

text: During TakeOver, The Great American Bash was scheduled for July 6 as a special episode of "NXT". During the June 15 episode of "NXT", continuing on from their encounter at TakeOver, as well as their past feud, Kyle O'Reilly and Adam Cole were signed for a match at The Great American Bash.


title: NXT TakeOver: In Your House (2021)

text: Also during the following episode, LA Knight was formally presented with the Million Dollar Championship by Ted DiBiase. After an emotional promo where he thanked DiBiase for being a fixture of his childhood and called receiving the championship from him the honor of his career, Knight turned on and attacked DiBiase (who was saved by Cameron Grimes) in a video uploaded to WWE's YouTube channel, it was shown that Grimes had apologized to DiBiase following the match at TakeOver.


title: NXT TakeOver: In Your House (2021)

text: The Santos Escobar-Bronson Reed and Xia Li-Mercedes Martinez feuds both continued in promos, with Escobar promising to face Reed for the NXT North American Championship one-on-one in the future.


title: NXT TakeOver: In Your House (2021)

text: In September 2021, NXT was rebranded as NXT 2.0, returning the brand to its original function as WWE's developmental territory. The NXT TakeOver series was subsequently discontinued, but In Your House continued on as NXT's annual June event.


title: Namastey London

text: Namastey London () is a 2007 Indian romance film, directed and produced by Vipul Amrutlal Shah, starring


title: Namastey London

text: Akshay Kumar and Katrina Kaif in lead roles. It also stars Rishi Kapoor, Nina Wadia, Javed Sheikh, Upen Patel and Clive Standen in supporting roles. The film is originally based on a real life story of Kumar's friend. Riteish Deshmukh appears in a cameo role in the film.


title: Namastey London

text: "Namastey London" released on 23 March 2007 and it received positive reviews from for its direction, screenplay, soundtrack and particularly Akshay Kumar's performance was highly acclaimed. It was also a box office success, earning 71.40 crore worldwide, thus becoming the ninth-highest grossing Hindi film of 2007. The success of the film ended a long dry spell of box office failures since the release of "" which released on January 2007.


title: Namastey London

text: Several critics noted similarities between the film and Manoj Kumar's "Purab Aur Paschim" (1970), though the director denied it being a remake.


title: Namastey London

text: Indian-born Manmohan Malhotra (Rishi Kapoor) re-located to London, England, established himself, returned to India, married Bebo (Nina Wadia), and after four years got a visa for her so that she could live with him. Shortly thereafter, she gave birth to Jasmeet (Katrina Kaif). Manmohan was always embarrassed by Bebo as she was overweight and could not speak English. As a result, he always left her out of important occasions, while he socialised. Bebo did not want Jasmeet to end up like her, so she got her educated in an English school, encouraged her to mingle with British friends, and Jasmeet was transformed into Jazz – a beautiful young woman, modern in looks, talk, habits, and heart.


title: Namastey London

text: Her father had set her to meet a young lad, Bobby Bedi (Ritesh Deshmukh), though the date turns out to be a disaster and Bobby rejects her. Manmohan thinks that it is impossible to get him an Indian son-in-law, although his British Pakistani friend, Parvez Khan (Javed Sheikh), is in a similar situation with his son, Imran (Upen Patel), who has an Indophobic girlfriend, Susan (Cathy Barry). Manmohan takes his family for a tour in India and forces Jasmeet to marry Arjun Singh (Akshay Kumar) who does not know how to speak English.


title: Namastey London

text: On their return to London, Jasmeet announces that she will marry an English man named Charlie Brown (Clive Standen), who is well educated and has good friends and connections, even with Jimmy Savile. She refuses to recognise her marriage to Arjun as there is no legal proof of the wedding. When Jasmeet is introduced to the friends of the Brown family, she is subjected to considerable abuse (Eve teasings, Indian rope tricks, Jai Shri Ram) as well as a quote from Winston Churchill: 'When we leave India, the country will be in the hands of goons'. Arjun then tells the gentleman about the achievements of India which includes about the derivation of English words from Sanskrit like 'Mother', 'Brother' , 'Trignometry' and 'Geometry'. During this period, Jazz becomes close with Arjun who still supports her. Charlie Brown then has a rugby match against the Indians/Pakistanis and Arjun, with the Desis emerging triumphant and Brown is verbally abusive towards them. Imran is then asked by Susan's parents to leave Islam, become a Christian, change his name to Emmanuel or Ian, as well as provide written proof that his family is not associated with terrorists. The film ends on a happy note when Imran decides not to be a Christian and Susan is accepted by Imran's family; Jazz realises that Arjun lied to her about not knowing English and runs away from her wedding to Charlie Brown, and goes with Arjun back to India where they are seen riding Arjun's motorcycle. While they are on the motorcycle Arjun reveals that all along he knew how to speak English, and that he had already dropped the clue when he got offended at Charlie's uncle's derogatory comments on India. He adds that he lied because he was still an Indian at heart.


title: Namastey London

text: The soundtrack album was released on 27 January 2007 and received good reviews. The songs were composed by Himesh Reshammiya. Salim–Sulaiman composed the film score. All songs were penned by Javed Akhtar.


title: Namastey London

text: "Namastey London" received positive reviews from critics upon release. Taran Adarsh of indiafm.com wrote, ""Namastey London" is addictive stuff. You watch it once, you wanna watch it again." Kumar received almost unanimous praise for his portrayal of Arjun Singh, with a critic saying "Kumar certainly leaves his mark throughout the flick. One of the coolest performances of the star." Adarsh said "One of his finest performances so far, "Namastey London" also marks the coming of age of an actor who was often dubbed as an action hero or a funster. Kumar not only wins the heart of Kaif on-screen, he's sure to win the hearts of millions of moviegoers with a terrific portrayal in this film." On Katrina Kaif's performance he wrote, "she handles the complex part with remarkable ease. The pretty lass is only getting better with every film." Despite the generally favorable reviews, Prachi Singh of "Moviewalah" panned the film, with a verdict of 2 out of 5 stars.


title: Namastey London

text: "Namastey London" opened to a good response and mainly picked up momentum after the second and third day of its release, a major reason being India's exit from the 2007 Cricket World Cup. The film did extremely well in the northern parts of India, especially in Punjab, Haryana, Rajasthan, and Uttar Pradesh. It also did well in Mumbai and in the first week of its release collected . The film was also received well in other parts of the country. The success of the film ended a long spell of box office failures since the release of "" which released on January 2007. The film went on to gross worldwide, making it one of the top hits of 2007.


title: Namastey London

text: The film was also successful overseas. It debuted at number 9 on the UK charts and similarly debuted within the top 20 in the United States and Australian charts. The film collected £238,841 in the first week of its release. As of 27 July 2007, the movie grossed an estimated $15,273,747 in five territories, which included the United States ($4,149,772), Australia ($197,148), India ($17,267,662), Malaysia ($15,285), and the United Kingdom ($9,021,900).


title: Namastey London

text: The film was unofficially remade in Bengali as "Paran Jai Jaliya Re" starring Dev and Subhashree Ganguly. The Kolkata High Court in December 2009 banned the screening of the film for plagiarism after a petition was filed by the film's producers.


title: Namastey London

text: In March 2011, Shah had announced a sequel, "Namaste England". Kumar and Sonakshi Sinha were initially signed to play the leads in the sequel. "Namaste England" was released in October 2018 with Parineeti Chopra and Arjun Kapoor starring.


title: Onychophora

text: Onychophora (from , , "claws"; and , , "to carry"), commonly known as velvet worms (due to their velvety texture and somewhat wormlike appearance) or more ambiguously as peripatus (after the first described genus, "Peripatus"), is a phylum of elongate, soft-bodied, many-legged panarthropods. In appearance they have variously been compared to worms with legs, caterpillars, and slugs. They prey upon other invertebrates, which they catch by ejecting an adhesive slime. Approximately 200 species of velvet worms have been described, although the true number of species is likely greater. The two extant families of velvet worms are Peripatidae and Peripatopsidae. They show a peculiar distribution, with the peripatids being predominantly equatorial and tropical, while the peripatopsids are all found south of the equator. It is the only phylum within Animalia that is wholly endemic to terrestrial environments, at least among extant members. Velvet worms are generally considered close relatives of the Arthropoda and Tardigrada, with which they form the proposed taxon Panarthropoda. This makes them of palaeontological interest, as they can help reconstruct the ancestral arthropod. In modern zoology they are particularly renowned for their curious mating behaviours and the bearing of live young in some species.


title: Onychophora

text: Velvet worms are segmented animals with a flattened cylindrical body cross-section and rows of unstructured body appendages known as oncopods or lobopods (informally: stub feet). They grow to between 0.5 and 20 cm (.2 to 8 in), with the average being about 5 cm (2  in). The number of leg pairs ranges from as few as 13 (in "Ooperipatellus nanus") to as many as 43 (in "Plicatoperipatus jamaicensis"). Their skin consists of numerous, fine transverse rings and is often inconspicuously coloured orange, red or brown, but sometimes also bright green, blue, gold or white, and occasionally patterned with other colours. Segmentation is outwardly inconspicuous, and identifiable by the regular spacing of the pairs of legs and in the regular arrangement of skin pores, excretion organs and concentrations of nerve cells. The individual body sections are largely unspecialised; even the head develops only a little differently from the abdominal segments. Segmentation is apparently specified by the same gene as in other groups of animals, and is activated in each case, during embryonic development, at the rear border of each segment and in the growth zone of the stub feet. Although onychophorans fall within the protostome group, their early development has a deuterostome trajectory, (with the mouth and anus forming separately); this trajectory is concealed by the rather sophisticated processes which occur in early development.


title: Onychophora

text: The stub feet that characterise the velvet worms are conical, baggy appendages of the body, which are internally hollow and have no joints. Although the number of feet can vary considerably between species, their structure is basically very similar. Rigidity is provided by the hydrostatic pressure of their fluid contents, and movement is usually obtained passively by stretching and contraction of the animal's entire body. However, each leg can also be shortened and bent by internal muscles. Due to the lack of joints, this bending can take place at any point along the sides of the leg. In some species, two different organs are found within the feet:


title: Onychophora

text: On each foot is a pair of retractable, hardened (sclerotised) chitin claws, which give the taxon its scientific name: Onychophora is derived from the , , "claws"; and , , "to carry". At the base of the claws are three to six spiny "cushions" on which the leg sits in its resting position and on which the animal walks over smooth substrates. The claws are used mainly to gain a firm foothold on uneven terrain. Each claw is composed of three stacked elements, like Russian nesting dolls. The outermost is shed during ecdysis, which exposes the next element in — which is fully formed, so does not need time to harden before it is used.<ref name="doi10.1038/nature13576"></ref> (This distinctive construction identifies many early Cambrian fossils as early offshoots of the onychophoran lineage.) Apart from the pairs of legs, there are three further body appendages, which are at the head and comprise three segments:


title: Onychophora

text: The surface of the mandibles is smooth, with no ornamentation. The cuticle in the mandibles (and claws) is distinct from the rest of the body. It has an inner and outer component; the outer component has just two layers (whereas body cuticle has four), and these outer layers (in particular the inner epicuticle) are dehydrated and strongly tanned, affording toughness.


title: Onychophora

text: On the third head segment, to the left and right of the mouth, are two openings designated "oral papillae". Within these are a pair of large, heavily internally branched slime glands. These lie roughly in the centre of the body and secrete a sort of milky-white slime, which is used to ensnare prey and for defensive purposes. Sometimes the connecting "slime conductor" is broadened into a reservoir, which can buffer pre-produced slime. The slime glands themselves are probably modified crural glands. All three structures correspond to an evolutionary origin in the leg pairs of the other segments.


title: Onychophora

text: Unlike the arthropods, velvet worms do not possess a rigid exoskeleton. Instead, their fluid-filled body cavity acts as a hydrostatic skeleton, similarly to many distantly related soft-bodied animals that are cylindrically shaped, for example sea anemones and various worms. Pressure of their incompressible internal bodily fluid on the body wall provides rigidity, and muscles are able to act against it. The body wall consists of a non-cellular outer skin, the cuticula; a single layer of epidermis cells forming an internal skin; and beneath this, usually three layers of muscle, which are embedded in connective tissues. The cuticula is about a micrometer thick and covered with fine villi. In composition and structure, it resembles the cuticula of the arthropods, consisting of α-chitin and various proteins, although not containing collagen. It can be divided into an external epicuticula and an internal procuticula, which themselves consist of exo- and endo-cuticula. This multi-level structure is responsible for the high flexibility of the outer skin, which enables the velvet worm to squeeze itself into the narrowest crevices. Although outwardly water-repellent, the cuticula is not able to prevent water loss by respiration, and, as a result, velvet worms can live only in microclimates with high humidity to avoid desiccation. The surface of the cuticula is scattered with numerous fine papillae, the larger of which carry visible villi-like sensitive bristles. The papillae themselves are covered with tiny scales, lending the skin a velvety appearance (from which the common name is likely derived). It also feels like dry velvet to the touch, for which its water-repellent nature is responsible. Moulting of the skin (ecdysis) takes place regularly, around every 14 days, induced by the hormone ecdysone. The inner surface of the skin bears a hexagonal pattern. At each moult, the shed skin is replaced by the epidermis, which lies immediately beneath it; unlike the cuticula, this consists of living cells. Beneath this lies a thick layer of connective tissue, which is composed primarily of collagen fibres aligned either parallel or perpendicular to the body's longitudinal axis. The colouration of Onychophora is generated by a range of pigments. The solubility of these pigments is a useful diagnostic character: in all arthropods and tardigrades, the body pigment is soluble in ethanol. This is also true for the Peripatidae, but in the case of the Peripatopsidae, the body pigment is insoluble in ethanol.


title: Onychophora

text: Within the connective tissue lie three continuous layers of unspecialised smooth muscular tissue. The relatively thick outer layer is composed of annular muscles, and the similarly voluminous inner layer of longitudinal muscles. Between them lie thin diagonal muscles that wind backward and forward along the body axis in a spiral. Between the annular and diagonal muscles exist fine blood vessels, which lie below the superficially recognisable transverse rings of the skin and are responsible for the pseudo-segmented markings. Beneath the internal muscle layer lies the body cavity. In cross-section, this is divided into three regions by so-called dorso-ventral muscles, which run from the middle of the underbelly through to the edges of the upper side: a central midsection and on the left and right, two side regions that also include the legs.


title: Onychophora

text: The body cavity is known as a "pseudocoel", or haemocoel. Unlike a true coelom, a pseudocoel is not fully enclosed by a cell layer derived from the embryonic mesoderm. A coelom is, however, formed around the gonads and the waste-eliminating nephridia. As the name "haemocoel" suggests, the body cavity is filled with a blood-like liquid in which all the organs are embedded; in this way, they can be easily supplied with nutrients circulating in the blood. This liquid is colourless as it does not contain pigments; for this reason, it serves only a limited role in oxygen transport.


title: Onychophora

text: Two different types of blood cells (or haemocytes) circulate in the fluid: Amoebocytes and nephrocytes. The amoebocytes probably function in protection from bacteria and other foreign bodies; in some species, they also play a role in reproduction. Nephrocytes absorb toxins or convert them into a form suitable for elimination by the nephridia.


title: Onychophora

text: The haemocoel is divided by a horizontal partition, the diaphragm, into two parts: The pericardial sinus along the back and the perivisceral sinus along the belly. The former encloses the tube-like heart, and the latter, the other organs. The diaphragm is perforated in many places, enabling the exchange of fluids between the two cavities. The heart itself is a tube of annular muscles consisting of epithelial tissues, with two lateral openings (ostia) per segment. While it is not known whether the rear end is open or closed, from the front, it opens directly into the body cavity.


title: Onychophora

text: Since there are no blood vessels, apart from the fine vessels running between the muscle layers of the body wall and a pair of arteries that supply the antennae, this is referred to as an open circulation. The timing of the pumping procedure can be divided into two parts: Diastole and systole. During diastole, blood flows through the ostia from the pericardial sinus (the cavity containing the heart) into the heart. When the systole begins, the ostia close and the heart muscles contract inwards, reducing the volume of the heart. This pumps the blood from the front end of the heart into the perivisceral sinus containing the organs. In this way, the various organs are supplied with nutrients before the blood finally returns to the pericardial sinus via the perforations in the diaphragm. In addition to the pumping action of the heart, body movements also influence circulation.


title: Onychophora

text: Oxygen uptake occurs to an extent via simple diffusion through the entire body surface, with the coxal vesicles on the legs possibly being involved in some species. However, of most importance is gas exchange via fine unbranched tubes, the tracheae, which draw oxygen from the surface deep into the various organs, particularly the heart.


title: Onychophora

text: The walls of these structures, which are less than three micrometers thick in their entirety, consist only of an extremely thin membrane through which oxygen can easily diffuse. The tracheae originate at tiny openings, the spiracles, which themselves are clustered together in dent-like recesses of the outer skin, the atria. The number of "tracheae bundles" thus formed is on average around 75 bundles per body segment; they accumulate most densely on the back of the organism.


title: Onychophora

text: Unlike the arthropods, the velvet worms are unable to control the openings of their tracheae; the tracheae are always open, entailing considerable water loss in arid conditions. Water is lost twice as fast as in earthworms and forty times faster than in caterpillars. For this reason, velvet worms are dependent upon habitats with high air humidity.


title: Onychophora

text: The digestive tract begins slightly behind the head, the mouth lying on the underside a little way from the frontmost point of the body. Here, prey can be mechanically dismembered by the mandibles with their covering of fine toothlets. Two salivary glands discharge via a common conductor into the subsequent "throat", which makes up the first part of the front intestine. The saliva that they produce contains mucus and hydrolytic enzymes, which initiate digestion in and outside the mouth.


title: Onychophora

text: The throat itself is very muscular, serving to absorb the partially liquified food and to pump it, via the oesophagus, which forms the rear part of the front intestine, into the central intestine. Unlike the front intestine, this is not lined with a cuticula but instead consists only of a single layer of epithelial tissue, which does not exhibit conspicuous indentation as is found in other animals.


title: Onychophora

text: On entering the central intestine, food particles are coated with a mucus-based peritrophic membrane, which serves to protect the lining of the intestine from damage by sharp-edged particles. The intestinal epithelium secretes further digestive enzymes and absorbs the released nutrients, although the majority of digestion has already taken place externally or in the mouth. Indigestible remnants arrive in the rear intestine, or rectum, which is once again lined with a cuticula and which opens at the anus, located on the underside near to the rear end.


title: Onychophora

text: In almost every segment is a pair of excretory organs called nephridia, which are derived from coelom tissue. Each consists of a small pouch that is connected, via a flagellated conductor called a nephridioduct, to an opening at the base of the nearest leg known as a nephridiopore. The pouch is occupied by special cells called podocytes, which facilitate ultrafiltration of the blood through the partition between haemocoelom and nephridium.


title: Onychophora

text: The composition of the urinary solution is modified in the nephridioduct by selective recovery of nutrients and water and by isolation of poison and waste materials, before it is excreted to the outside world via the nephridiopore. The most important nitrogenous excretion product is the water-insoluble uric acid; this can be excreted in solid state, with very little water. This so-called uricotelic excretory mode represents an adjustment to life on land and the associated necessity of dealing economically with water.


title: Onychophora

text: A pair of former nephridia in the head were converted secondarily into the salivary glands, while another pair in the final segment of male specimens now serve as glands that apparently play a role in reproduction.


title: Onychophora

text: The entire body, including the stub feet, is littered with numerous papillae: warty protrusions responsive to touch that carry a mechanoreceptive bristle at the tip, each of which is also connected to further sensory nerve cells lying beneath. The mouth papillae, the exits of the slime glands, probably also have some function in sensory perception. Sensory cells known as "sensills" on the "lips" or labrum respond to chemical stimuli and are known as chemoreceptors. These are also found on the two antennae, which seem to be the velvet worm's most important sensory organs.


title: Onychophora

text: Except in a few (typically subterranean) species, one simply constructed eye (ocellus) lies behind each antenna, laterally, just underneath the head. This consists of a chitinous ball lens, a cornea and a retina and is connected to the centre of the brain via an optic nerve. The retina comprises numerous pigment cells and photoreceptors; the latter are easily modified flagellated cells, whose flagellum membranes carry a photosensitive pigment on their surface.


title: Onychophora

text: The rhabdomeric eyes of the Onychophora are thought to be homologous with the median ocelli of arthropods; this would suggest that the last common ancestor of arthropods may have only had median ocelli.


title: Onychophora

text: However, the innervation shows that the homology is limited: The eyes of Onychophora form behind the antenna, whereas the opposite is true in arthropods.


title: Onychophora

text: Both sexes possess pairs of gonads, opening via a channel called a gonoduct into a common genital opening, the gonopore, which is located on the rear ventral side. Both the gonads and the gonoduct are derived from true coelom tissue.


title: Onychophora

text: In females, the two ovaries are joined in the middle and to the horizontal diaphragm. The gonoduct appears differently depending on whether the species is live-bearing or egg-laying. In live-bearing species, each exit channel divides into a slender oviduct and a roomy "womb", the uterus, in which the embryos develop. The single vagina, to which both uteri are connected, runs outward to the gonopore. In egg-laying species, whose gonoduct is uniformly constructed, the genital opening lies at the tip of a long egg-laying apparatus, the ovipositor. The females of many species also possess a sperm repository called the "receptacle seminis", in which sperm cells from males can be stored temporarily or for longer periods.


title: Onychophora

text: Males possess two separate testes, along with the corresponding sperm vesicle (the vesicula seminalis) and exit channel (the vasa efferentia). The two vasa efferentia unite to a common sperm duct, the vas deferens, which in turn widens through the ejaculatory channel to open at the gonopore. Directly beside or behind this lie two pairs of special glands, which probably serve some auxiliary reproductive function; the rearmost glands are also known as anal glands. A penis-like structure has so far been found only in males of the genus "Paraperipatus" but has not yet been observed in action.


title: Onychophora

text: There are different mating procedures: In some species males deposit their spermatophore directly into the female's genitals opening, while others deposit it on the female's body, where the cuticle will collapse and allowing the sperm cells to migrate into the female. There are also Australian species where the male place their spermatophore on top of their head, which is then pressed against the female's genitals. In these species the head have elaborate structures like spikes, spines, hollow stylets, pits, and depressions, whose purpose is to either hold the sperm and / or assist in the sperm transfer to the female. The males of most species also secrete a pheromone from glands on the underside of the legs to attract females.


title: Onychophora

text: Velvet worms live in all tropical habitats and in the temperate zone of the Southern Hemisphere, showing a circumtropical and circumaustral distribution. Individual species are found in Central and South America; the Caribbean islands; equatorial West Africa and Southern Africa; northeastern India; Thailand; Indonesia and parts of Malaysia; New Guinea; Australia; and New Zealand.


title: Onychophora

text: Fossils have been found in Baltic amber, indicating that they were formerly more widespread in the Northern Hemisphere when conditions were more suitable.


title: Onychophora

text: Velvet worms always sparsely occupy the habitats where they are found: They are rare among the fauna which they are a part of.


title: Onychophora

text: All extant velvet worms are terrestrial (land-living) and prefer dark environments with high air humidity. They are found particularly in the rainforests of the tropics and temperate zones, where they live among moss cushions and leaf litter, under tree trunks and stones, in rotting wood or in termite tunnels. They also occur in unforested grassland, if there exist sufficient crevices in the soil into which they can withdraw during the day, and in caves. Two species live in caves, a habitat to which their ability to squeeze themselves into the smallest cracks makes them exceptionally well-adapted and in which constant living conditions are guaranteed. Since the essential requirements for cave life were probably already present prior to the settlement of these habitats, this may be described as exaptation.


title: Onychophora

text: Some species of velvet worms are able to occupy human-modified land-uses, such as cocoa and banana plantations in South America and the Caribbean, but for others, conversion of rainforests is likely one of the most important threats to their survival (see Conservation).


title: Onychophora

text: Velvet worms are photophobic: They are repelled by bright light sources. Because the danger of desiccation is greatest during the day and in dry weather, it is not surprising that velvet worms are usually most active at night and during rainy weather. Under cold or dry conditions, they actively seek out crevices in which they shift their body into a resting state.


title: Onychophora

text: The Onychophora forcefully squirt glue-like slime from their oral papillae; they do so either in defense against predators or to capture prey. The openings of the glands that produce the slime are in the papillae, a pair of highly modified limbs on the sides of the head below the antennae. Inside, they have a syringe-like system that, by a geometric amplifier, allows for fast squirt using slow muscular contraction. High speed films show the animal expelling two streams of adhesive liquid through a small opening (50 to 200 microns) at a speed of . The interplay between the elasticity of oral papillae and the fast unsteady flow produces a passive oscillatory motion (30–60 Hz) of the oral papillae. The oscillation causes the streams to cross in mid air, weaving a disordered net; the velvet worms can control only the general direction where the net is thrown.


title: Onychophora

text: The slime glands themselves are deep inside the body cavity, each at the end of a tube more than half the length of the body. The tube both conducts the fluid and stores it until it is required. The distance that the animal can propel the slime varies; usually it squirts it about a centimetre, but the maximal range has variously been reported to be ten centimetres, or even nearly a foot, although accuracy drops with range. It is not clear to what extent the range varies with the species and other factors. One squirt usually suffices to snare a prey item, although larger prey may be further immobilised by smaller squirts targeted at the limbs; additionally, the fangs of spiders are sometimes targeted. Upon ejection, it forms a net of threads about twenty microns in diameter, with evenly spaced droplets of viscous adhesive fluid along their length. It subsequently dries, shrinking, losing its stickiness, and becoming brittle. Onychophora eat their dried slime when they can, which is appropriate, because it takes an onychophoran about 24 days to replenish an exhausted slime repository.


title: Onychophora

text: The slime can account for up to 11% of the organism's dry weight and is 90% water; its dry residue consists mainly of proteins — primarily a collagen-type protein. 1.3% of the slime's dry weight consists of sugars, mainly galactosamine. The slime also contains lipids and the surfactant nonylphenol. Onychophora are the only organisms known to produce this latter substance. It tastes "slightly bitter and at the same time somewhat astringent". The proteinaceous composition accounts for the slime's high tensile strength and stretchiness. The lipid and nonylphenol constituents may serve one of two purposes. They may line the ejection channel, stopping the slime from sticking to the organism when it is secreted; or they may slow the drying process long enough for the slime to reach its target.


title: Onychophora

text: Velvet worms/Onychophora move in a slow and gradual motion that makes them difficult for prey to notice. Their trunk is raised relatively high above the ground, and they walk with non-overlapping steps.


title: Onychophora

text: To move from place to place, the velvet worm crawls forward using its legs; unlike in arthropods, both legs of a pair are moved simultaneously. The claws of the feet are used only on hard, rough terrain where a firm grip is needed; on soft substrates, such as moss, the velvet worm walks on the foot cushions at the base of the claws.


title: Onychophora

text: Actual locomotion is achieved less by the exertion of the leg muscles than by local changes of body length. This can be controlled using the annular and longitudinal muscles. If the annular muscles are contracted, the body cross-section is reduced, and the corresponding segment lengthens; this is the usual mode of operation of the hydrostatic skeleton as also employed by other worms. Due to the stretching, the legs of the segment concerned are lifted and swung forward. Local contraction of the longitudinal muscles then shortens the appropriate segment, and the legs, which are now in contact with the ground, are moved to the rear. This part of the locomotive cycle is the actual leg stroke that is responsible for forward movement. The individual stretches and contractions of the segments are coordinated by the nervous system such that contraction waves run the length of the body, each pair of legs swinging forward and then down and rearward in succession. "Macroperipatus" can reach speeds of up to four centimetres per second, although speeds of around 6 body-lengths per minute are more typical.<ref name="doi10.1111/j.1096-3642.1950.tb01699.x"></ref> The body gets longer and narrower as the animal picks up speed; the length of each leg also varies during each stride.


title: Onychophora

text: The brains of Onychophora, though small, are very complex; consequently, the organisms are capable of rather sophisticated social interactions. Behaviour may vary from genus to genus, so this article reflects the most-studied genus, "Euperipatoides".


title: Onychophora

text: The "Euperipatoides" form social groups of up to fifteen individuals, usually closely related, which will typically live and hunt together. Groups usually live together; in drier regions an example of a shared home would be the moist interior of a rotting log. Group members are extremely aggressive towards individuals from other logs. Dominance is achieved through aggression and maintained through submissive behaviour. After a kill, the dominant female always feeds first, followed in turn by the other females, then males, then the young.


title: Onychophora

text: When assessing other individuals, individuals often measure one another up by running their antennae down the length of the other individual. Once hierarchy has been established, pairs of individuals will often cluster together to form an "aggregate"; this is fastest in male-female pairings, followed by pairs of females, then pairs of males.


title: Onychophora

text: Social hierarchy is established by a number of interactions: Higher-ranking individuals will chase and bite their subordinates while the latter are trying to crawl on top of them. Juveniles never engage in aggressive behaviour, but climb on top of adults, which tolerate their presence on their backs.


title: Onychophora

text: Hierarchy is quickly established among individuals from a single group, but not among organisms from different groups; these are substantially more aggressive and very rarely climb one another or form aggregates. Individuals within an individual log are usually closely related; especially so with males. This may be related to the intense aggression between unrelated females.


title: Onychophora

text: Velvet worms are ambush predators, hunting only by night, and are able to capture animals at least their own size, although capturing a large prey item may take almost all of their mucus-secreting capacity. They feed on almost any small invertebrates, including woodlice (Isopoda), termites (Isoptera), crickets (Gryllidae), book/bark lice (Psocoptera), cockroaches (Blattidae), millipedes and centipedes (Myriapoda), spiders (Araneae), various worms, and even large snails (Gastropoda). Depending on their size, they eat on average every one to four weeks. They are considered to be ecologically equivalent to centipedes (Chilopoda). The most energetically favourable prey are two-fifths the size of the hunting onychophoran. Ninety percent of the time involved in eating prey is spent ingesting it; re-ingestion of the slime used to trap the insect is performed while the onychophoran locates a suitable place to puncture the prey, and this phase accounts for around 8% of the feeding time, with the remaining time evenly split between examining, squirting, and injecting the prey. In some cases, chunks of the prey item are bitten off and swallowed; undigestable components take around 18 hours to pass through the digestive tract. Onychophora probably do not primarily use vision to detect their prey; although their tiny eyes do have a good image-forming capacity, their forward vision is obscured by their antennae; their nocturnal habit also limits the utility of eyesight. Air currents, formed by prey motion, are thought to be the primary mode of locating prey; the role of scent, if any, is unclear. Because it takes so long to ingest a prey item, hunting mainly happens around dusk; the onychophorans will abandon their prey at sunrise. This predatory way of life is probably a consequence of the velvet worm's need to remain moist. Due to the continual risk of desiccation, often only a few hours per day are available for finding food. This leads to a strong selection for a low cost-benefit ratio, which cannot be achieved with a herbivorous diet.


title: Onychophora

text: Velvet worms literally creep up on their prey, with their smooth, gradual and fluid movement escaping detection. Once they reach their prey, they touch it very softly with their antennae to assess its size and nutritional value. After each poke, the antenna is hastily retracted to avoid alerting the prey. This investigation may last anywhere upwards of ten seconds, until the velvet worm makes a decision as to whether to attack it, or until it disturbs the prey and the prey flees. Hungry Onychophora spend less time investigating their prey and are quicker to apply their slime. Once slime has been squirted, Onychophora are determined to pursue and devour their prey, in order to recoup the energy investment. They have been observed to spend up to ten minutes searching for removed prey, after which they return to their slime to eat it. In the case of smaller prey, they may opt not to use slime at all. Subsequently, a soft part of the prey item (usually a joint membrane in arthropod prey) is identified, punctured with a bite from the jaws, and injected with saliva. This kills the prey very quickly and begins a slower process of digestion. While the onychophoran waits for the prey to digest, it salivates on its slime and begins to eat it (and anything attached to it). It subsequently tugs and slices at the earlier perforation to allow access to the now-liquefied interior of its prey. The jaws operate by moving backwards and forwards along the axis of the body (not in a side-to-side clipping motion as in arthropods), conceivably using a pairing of musculature and hydrostatic pressure. The pharynx is specially adapted for sucking, to extract the liquefied tissue; the arrangement of the jaws about the tongue and lip papillae ensures a tight seal and the establishment of suction. In social groups, the dominant female is the first to feed, not permitting competitors access to the prey item for the first hour of feeding. Subsequently, subordinate individuals begin to feed. The number of males reaches a peak after females start to leave the prey item. After feeding, individuals clean their antennae and mouth parts before re-joining the rest of their group.


title: Onychophora

text: Almost all species of velvet worm reproduce sexually. The sole exception is "Epiperipatus imthurni", of which no males have been observed; reproduction instead occurs by parthenogenesis. All species are in principle sexually distinct and bear, in many cases, a marked sexual dimorphism: the females are usually larger than the males and have, in species where the number of legs is variable, more legs. The females of many species are fertilized only once during their lives, which leads to copulation sometimes taking place before the reproductive organs of the females are fully developed. In such cases, for example at the age of three months in "Macroperipatus torquatus", the transferred sperm cells are kept in a special reservoir, where they can remain viable for longer periods. Fertilization takes place internally, although the mode of sperm transmission varies widely. In most species, for example in the genus "Peripatus", a package of sperm cells called the spermatophore is placed into the genital opening of the female. The detailed process by which this is achieved is in most cases still unknown, a true penis having been observed only in species of the genus "Paraperipatus". In many Australian species, there exist dimples or special dagger- or axe-shaped structures on the head; the male of "Florelliceps stutchburyae" presses a long spine against the female's genital opening and probably positions its spermatophore there in this way. During the process, the female supports the male by keeping him clasped with the claws of her last pair of legs. The mating behavior of two species of the genus "Peripatopsis" is particularly curious. Here, the male places two-millimetre spermatophores on the back or sides of the female. Amoebocytes from the female's blood collect on the inside of the deposition site, and both the spermatophore's casing and the body wall on which it rests are decomposed via the secretion of enzymes. This releases the sperm cells, which then move freely through the haemocoel, penetrate the external wall of the ovaries and finally fertilize the ova. Why this self-inflicted skin injury does not lead to bacterial infections is not yet understood (though likely related to the enzymes used to deteriorate the skin or facilitate the transfer of viable genetic material from male to female). Velvet worms are found in egg-laying (oviparous), egg-live-bearing (ovoviviparous) and live-bearing (viviparous) forms.


title: Onychophora

text: A female can have between 1 and 23 offspring per year; development from fertilized ovum to adult takes between 6 and 17 months and does not have a larval stage. This is probably also the original mode of development. Velvet worms have been known to live for up to six years.


title: Onychophora

text: The velvet worm's important predators are primarily various spiders and centipedes, along with rodents and birds, such as, in Central America, the clay-coloured thrush ("Turdus grayi"). In South America, Hemprichi's coral snake ("Micrurus hemprichii") feeds almost exclusively on velvet worms. For defence, some species roll themselves reflexively into a spiral, while they can also fight off smaller opponents by ejecting slime. Various mites (Acari) are known to be ectoparasites infesting the skin of the velvet worm. Skin injuries are usually accompanied by bacterial infections, which are almost always fatal.


title: Onychophora

text: The South African species "Peripatopsis capensis" has been inadvertently introduced to Santa Cruz Island in the Galapagos Islands, where it co-occurs with native velvet worms.


title: Onychophora

text: The global conservation status of velvet worm species is difficult to estimate; many species are only known to exist at their type locality (the location at which they were first observed and described). The collection of reliable data is also hindered by low population densities, their typically nocturnal behaviour and possibly also as-yet undocumented seasonal influences and sexual dimorphism. To date, the only onychophorans evaluated by the IUCN are:


title: Onychophora

text: The primary threat comes from destruction and fragmentation of velvet worm habitat due to industrialisation, draining of wetlands, and slash-and-burn agriculture. Many species also have naturally low population densities and closely restricted geographic ranges; as a result, relatively small localised disturbances of important ecosystems can lead to the extinction of entire populations or species. Collection of specimens for universities or research institutes also plays a role on a local scale. There is a very pronounced difference in the protection afforded to velvet worms between regions: in some countries, such as South Africa, there are restrictions on both collecting and exporting, while in others, such as Australia, only export restrictions exist. Many countries offer no specific safeguards at all. Tasmania has a protection programme that is unique worldwide: one region of forest has its own velvet worm conservation plan, which is tailored to a particular velvet worm species.


title: Onychophora

text: In their present forms, the velvet worms are probably very closely related to the arthropods, a very extensive taxon that incorporates, for instance, the crustaceans, insects, and arachnids. They share, among other things, an exoskeleton consisting of α-chitin and non-collagenous proteins; gonads and waste-elimination organs enclosed in true coelom tissue; an open blood system with a tubular heart situated at the rear; an abdominal cavity divided into pericardial and perivisceral cavities; respiration via tracheae; and similar embryonic development. Segmentation, with two body appendages per segment, is also a shared feature.


title: Onychophora

text: However, the antennae, mandibles, and oral papillae of velvet worms are probably "not" homologous to the corresponding features in arthropods; i.e., they probably developed independently.


title: Onychophora

text: Another closely related group are the comparatively obscure water bears (Tardigrada); however, due to their very small size, water bears have no need for – and hence lack – blood circulation and separate respiratory structures: shared characteristics that support common ancestry of velvet worms and arthropods.


title: Onychophora

text: Together, the velvet worms, arthropods, and water bears form a monophyletic taxon, the Panarthropoda, i.e., the three groups collectively cover all descendants of their last common ancestor. Due to certain similarities of form, the velvet worms were usually grouped with the water bears to form the taxon Protoarthropoda. This designation would imply that both velvet worms and water bears are not yet as highly developed as the arthropods. Modern systematic theories reject such conceptions of "primitive" and "highly developed" organisms and instead consider exclusively the historical relationships among the taxa. These relationships are not as yet fully understood, but it is considered probable that the velvet worms' sister groups form a taxon designated Tactopoda, thus:


title: Onychophora

text: For a long time, velvet worms were also considered related to the annelids. They share, among other things, a worm-like body; a thin and flexible outer skin; a layered musculature; paired waste-elimination organs; as well as a simply constructed brain and simple eyes. Decisive, however, was the existence of segmentation in both groups, with the segments showing only minor specialisation. The parapodia appendages found in annelids therefore correspond to the stump feet of the velvet worms. Within the Articulata hypothesis developed by Georges Cuvier, the velvet worms therefore formed an evolutionary link between the annelids and the arthropods: worm-like precursors first developed parapodia, which then developed further into stub feet as an intermediate link in the ultimate development of the arthropods' appendages. Due to their structural conservatism, the velvet worms were thus considered "living fossils". This perspective was expressed paradigmatically in the statement by the French zoologist A. Vandel:


title: Onychophora

text: Modern taxonomy does not study criteria such as "higher" and "lower" states of development or distinctions between "main" and "side" branches—only family relationships indicated by cladistic methods are considered relevant. From this point of view, several common characteristics still support the Articulata hypothesis — segmented body; paired appendages on each segment; pairwise arrangement of waste-elimination organs in each segment; and above all, a rope-ladder-like nervous system based on a double nerve strand lying along the belly. An alternative concept, most widely accepted today, is the so-called Ecdysozoa hypothesis. This places the annelids and Panarthropoda in two very different groups: the former in the Lophotrochozoa and the latter in the Ecdysozoa. Mitochondrial gene sequences also provide support for this hypothesis. Proponents of this hypothesis assume that the aforementioned similarities between annelids and velvet worms either developed convergently or were primitive characteristics passed unchanged from a common ancestor to both the Lophotrochozoa and Ecdysozoa. For example, in the first case, the rope-ladder nervous system would have developed in the two groups independently, while in the second case, it is a very old characteristic, which does not imply a particularly close relationship between the annelids and Panarthropoda. The Ecdysozoa concept divides the taxon into two, the Panarthropoda into which the velvet worms are placed, and the sister group Cycloneuralia, containing the threadworms (Nematoda), horsehair worms (Nematomorpha) and three rather obscure groups: the mud dragons (Kinorhyncha); penis worms (Priapulida); and brush-heads (Loricifera).


title: Onychophora

text: Particularly characteristic of the Cycloneuralia is a ring of "circumoral" nerves around the mouth opening, which the proponents of the Ecdysozoa hypothesis also recognise in modified form in the details of the nerve patterns of the Panarthropoda. Both groups also share a common skin-shedding mechanism (ecdysis) and molecular biological similarities. One problem of the Ecdysozoa hypothesis is the velvet worms' subterminal position of their mouths: Unlike in the Cycloneuralia, the mouth is not at the front end of the body, but lies further back, under the belly. However, investigations into their developmental biology, particularly regarding the development of the head nerves, suggest that this was not always the case, and that the mouth was originally terminal (situated at the tip of the body). This is supported by the fossil record.


title: Onychophora

text: The "stem-group arthropod" hypothesis is very widely accepted, but some trees suggest that the onychophorans may occupy a different position; their brain anatomy is more closely related to that of the chelicerates than to any other arthropod. The modern velvet worms form a monophyletic group, incorporating all the descendants of their common ancestor. Important common derivative characteristics (synapomorphies) include, for example, the mandibles of the second body segment and the oral papillae and associated slime glands of the third; nerve strands extending along the underside with numerous cross-linkages per segment; and the special form of the tracheae. By 2011, some 180 modern species, comprising 49 genera, had been described; the actual number of species is probably about twice this. According to more recent study, 82 species of Peripatidae and 115 species of Peripatopsidae have been described thus far. However, among the 197 species, 20 are "nomina dubia", due to major taxonomic inconsistencies. The best-known is the type genus "Peripatus", which was described as early as 1825 and which, in English-speaking countries, stands representative for all velvet worms. All genera are assigned to one of two families, the distribution ranges of which do not overlap but are separated by arid areas or oceans:


title: Onychophora

text: Certain fossils from the early Cambrian bear a striking resemblance to the velvet worms. These fossils, known collectively as the lobopodians, were marine and represent a grade from which arthropods, tardigrades, and Onychophora arose.<ref name="Cite doi|10.1038/nature13576"></ref>


title: Onychophora

text: Historically, all fossil Onychophora and lobopods were lumped into the taxon Xenusia, further subdivided by some authors to the Paleozoic Udeonychophora and the Mesozoic/Tertiary Ontonychophora; living Onychophora were termed Euonychophora. Importantly, few of the Cambrian fossils bear features that distinctively unite them with the Onychophora; none can be confidently assigned to the onychophoran crown or even stem group. The exceptions are "Hallucigenia" and related taxa such as "Collinsium ciliosum", which bear distinctly onychophoran-like claws. It is not clear when the transition to a terrestrial existence was made, but it is considered plausible that it took place between the Ordovician and late Silurian—approximately —via the intertidal zone.


title: Onychophora

text: The low preservation potential of the non-mineralised Onychophora means that they have a sparse fossil record. Stem-group members include "Helenodora" (Carboniferous), "Tertiapatus dominicanus", and "Succinipatopsis balticus" (Tertiary). A Carboniferous fossil from Montceau-les-Mines, France, "Antennipatus" possesses clear onychophoran affinities, but its preservation prohibits differentiating between its placement on the stem or crown of the two extant families, or on the onychophoran stem-group. Crown group representatives are known only from amber—there is a single, partial specimen from the Cretaceous, and a more comprehensive record in Eocene deposits from . However, some of these amber-borne specimens lack slime papillae and separate feet, and thus may belong in the stem group.


title: Onychophora

text: The vagaries of the preservation process can make fossils difficult to interpret. Experiments on the decay and compaction of onychophora demonstrate difficulties in interpreting fossils; certain parts of living onychophora are visible only in certain conditions:


title: Onychophora

text: More significantly, features seen in fossils may be artefacts of the preservation process: For instance, "shoulder pads" may simply be the second row of legs coaxially compressed onto the body; branching "antennae" may in fact be produced through decay.


title: Roman Gabriel

text: Roman Ildonzo Gabriel Jr. (born August 5, 1940) is a former American football quarterback who played in the National Football League (NFL). He was the second overall pick in the 1962 NFL Draft and played for the Los Angeles Rams for eleven seasons, then five seasons for the Philadelphia Eagles. He is notable for being the first NFL quarterback of Filipino-American descent.


title: Roman Gabriel

text: Gabriel was born to Edna Mae Wyatt and Roman Ildonzo Gabriel Sr., a Filipino immigrant, in Wilmington, North Carolina. Gabriel grew up poor and suffered from asthma, but he played high school football at New Hanover High School and graduated in 1958. He went on to star at quarterback at North Carolina State University in Raleigh.


title: Roman Gabriel

text: A two-time All-American and two-time ACC Player of the Year (1960–61), Gabriel finished his college career holding virtually every Wolfpack passing record. An academic All-American, Gabriel saw his jersey retired after his senior season and then presented to him by North Carolina governor Terry Sanford on January 20, 1962, at halftime of an NC State-Maryland basketball game in Reynolds Coliseum. As captain of his team, Gabriel set 22 school and nine conference football records. He threw for 2,961 yards and 19 touchdowns.


title: Roman Gabriel

text: Known for his arm strength, he also played baseball and was voted the best amateur athlete in the Carolinas. In a three-year career, he passed for 20 touchdowns and ran for 15. The Atlantic Coast Conference's 50th Anniversary Football Team was announced in 2003 and Gabriel was among the top 50 players in the history of the ACC to be listed. Gabriel was inducted into the College Football Hall of Fame in 1989.


title: Roman Gabriel

text: Gabriel was the number one 1962 AFL Draft pick, chosen by the Oakland Raiders, and was the number 2 1962 NFL Draft pick, selected by the Los Angeles Rams. Gabriel signed with the Rams and went on to a distinguished professional career.


title: Roman Gabriel

text: Gabriel wore the number 18 with the Rams and the number 5 with the Eagles. In the professional ranks, Gabriel went on to play 16 seasons in the NFL, splitting time with the Los Angeles Rams (1962–72) and the Philadelphia Eagles (1973–77). He was awarded the NFL Most Valuable Player Award in 1969 and earned Pro Bowl spots in 1967, 1968, 1969, and 1973. When he retired, he ranked as the Rams' all-time passing leader with 22,223 yards and 154 touchdowns (1,705 com./3,313 att.) and threw for 7,221 yards and 45 touchdowns (661 com./1,185 att.) with the Eagles. In 1973, he led the NFL with 3,219 yards and 23 touchdown passes, for which he was awarded the NFL Comeback Player of the Year Award. As of the end of the 2016 NFL season, he still holds the Rams' career records for touchdown passes (154), passes attempted (3,313), and wins by a starting quarterback (74).


title: Roman Gabriel

text: From 1962 through 1965, Gabriel had a difficult time securing a starting quarterback job. Ram coaches gave Zeke Bratkowski or Bill Munson the nod over Gabriel. However, due to other quarterbacks slumping or being injured, Gabriel did get to start 23 games from 1962 through 1965. The team's record in those games was 11–11–1. Although his record as a starter was average, the other Rams quarterbacks who started the other 33 games combined record was 4–27–2. Gabriel's significant wins include a 1965 victory to beat the eventual NFL champion Green Bay Packers and the 11–3 Cleveland Browns.


title: Roman Gabriel

text: When George Allen took over for Harland Svare to coach the Rams in 1966, one of his first moves was to make Gabriel the primary starter. Gabriel started all fourteen games and the Rams went 8–6, their first winning season since 1958. In 1967 the Rams went 11–1–2 and made the playoffs as NFL Coastal Division champions. Gabriel was named the AP Offensive Player of the Week the last two weeks of the season. In week 13, needing a win to keep their playoff hopes alive, Gabriel was 20-for-36 with three touchdowns (including the game-winner in the last minute) in a 27–24 come from behind win over the defending champion Green Bay Packers. The next week, in a game against the Baltimore Colts that would decide the division title, Gabriel completed 18-of-22 passes with three touchdowns as the Rams won 34–10. The 1967 Rams finished as the highest-scoring team in the NFL but were eliminated from the playoffs by the Packers 28–7. Gabriel threw for 2,779 yards and 25 touchdowns and was a second-team All-Pro and a Pro Bowler.


title: Roman Gabriel

text: The following season they were in another neck and neck battle for the Coastal Division title with the Colts. Going into the thirteenth game of the season, the Rams needed a win to stay within one-half game of the Colts, who would be coming to Los Angeles the following week for the season finale. However, the Rams took a 17–16 loss to the Chicago Bears and soon finished in second place.


title: Roman Gabriel

text: In 1969, the Rams opened the season with an eleven-game winning streak (still a team record), before suffering their first loss to the 10–1 Minnesota Vikings in Los Angeles by a score of 20–13. With the division clinched and the undefeated record gone, coach Allen decided to rest many of his starters and the Rams lost their last two games to finish 11–3. In a rematch with the Vikings in the playoffs in Minnesota, the Rams lost, 23–20. For the season, Gabriel threw 24 touchdowns and only seven interceptions and was named the NFL's Most Valuable Player by the AP and NEA, the Player of the Year by the UPI and was voted All-Pro and to the Pro Bowl.


title: Roman Gabriel

text: In 1970, the league realigned, putting the Rams in contention with the San Francisco 49ers for the new NFC West Division title. After an upset loss at home to the lowly New York Jets (who were without the injured Joe Namath) in which Gabriel threw three interceptions, the Rams won three straight games, including a crucial 30–13 win over the 49ers to take over first place. Going into the thirteenth week of the season, the Rams participated in the first Monday Night Football game in the city of Los Angeles. However, despite over 300 yards passing from Gabriel, the Rams lost 28–23 to the Detroit Lions, putting the Rams back in 2nd place and left them a half-game behind the Lions for the wild card playoff spot. The Rams won their finale at the New York Giants 31–3 (eliminating the Giants from playoff contention) but failed to make the playoffs as the Lions won to secure the wild card spot, and the 49ers won to clinch the NFC Western Division title. From 1967 to 1970, Gabriel led the Rams to a 41–14–4 overall record and was named to three Pro Bowls during that four-year span.


title: Roman Gabriel

text: In 1971, the veteran Rams began to show their age and Gabriel missed parts of every game due to knee and shoulder injuries. In addition, coach George Allen left for the Redskins after a long-running dispute with general manager Dan Reeves. Still, the Rams, despite playing the league's toughest schedule, faced almost the same situation as in 1970. After falling behind the 49ers in the seventh week of the season, they rallied back to take the division lead going into another matchup in Los Angeles on Monday night for the thirteenth week of the season, for which the matchup was against Washington (now coached by Allen). After falling behind 31–10, Gabriel led the Rams back to within 31–24 and was driving to a possible tying score when he was intercepted; it was returned for a touchdown and the Redskins won. Once again, the Rams fell into second place behind the 49ers and behind the Redskins for the wild card berth. Despite winning in Pittsburgh in the final week of the year, the 49ers came from behind to beat the Lions, 31–27, and win the division.


title: Roman Gabriel

text: In 1972, Gabriel's knee and shoulder injury problems deteriorated. After making 89 consecutive starts over eight seasons, he missed two games and lost playing time in all twelve other games. Still, after a Monday night win in San Francisco in the twelfth week, the Rams regained first place. But losses to the Cardinals and Lions in the final two weeks (combined with a previous loss to the woeful Saints), doomed their season. The Rams finished 6–7–1 and coach Tommy Prothro was fired.


title: Roman Gabriel

text: After the 1972 season, the Rams hired Chuck Knox as their new coach and obtained John Hadl to be the quarterback. After he threatened to accept a $100,000 contract with the Las Vegas Casinos of the Southwestern Football League in April 1973, Gabriel was traded from the Rams to the Philadelphia Eagles for Harold Jackson, Tony Baker, a 1974 first-round selection (11th overall–John Cappelletti) and first- and third-round picks in 1975 (11th and 67th overall–Dennis Harrah and Dan Nugent) on June 8, 1973.


title: Roman Gabriel

text: Gabriel improved a 2–11–1 Eagles team to a 5–8–1 record. Gabriel was voted to the Pro Bowl for the fourth time and was voted the "Comeback Player of the Year" by "Pro Football Weekly". For the 1973 season, Gabriel led the Eagles with 270 completions, 460 attempts, and 3,219 yards, and 23 touchdowns (all were league highs) as the Eagle offense was the most prolific passing game in the NFL. Gabriel played through 1977 but his final two years were in a backup role. In his last season, he backed up Ron Jaworski, who had played for the Rams from 1973 to 1976.


title: Roman Gabriel

text: In his career, he had a winning record of 86–64–7 and passed for over 29,000 yards and 201 touchdowns. He is the only quarterback from his era to still rank high in the "lowest interception percentage" category in NFL passing statistics. The Professional Football Researchers Association named Gabriel to the PFRA Hall of Very Good Class of 2013.


title: Roman Gabriel

text: After his playing career, he had a brief two-year career as a member of the NFL on CBS broadcasting team from 1978 to 1979.


title: Roman Gabriel

text: Gabriel was the last football coach at Cal Poly Pomona, where from 1980 to 1982 his teams compiled an 8–24 record. On November 26, 1982, he resigned to become offensive coordinator with the Boston Breakers of the USFL.Cal Poly-Pomona terminated its football program.


title: Roman Gabriel

text: Gabriel was head coach of the Raleigh-Durham Skyhawks of the World League of American Football. He was the only coach who did not win a game in the inaugural 1991 season. The Skyhawks disbanded shortly thereafter.


title: Roman Gabriel

text: Roman Gabriel was a frequent guest on television talk shows of the era, including "The Merv Griffin Show", "The Virginia Graham Show", "The Rosey Grier Show", and "The Joey Bishop Show".


title: Roman Gabriel

text: Gabriel had a brief career in movies, playing a prison guard in Otto Preminger's 1968 spoof "Skidoo" and a Native American named "Blue Boy" in the 1969 John Wayne and Rock Hudson film, "The Undefeated." He had previously appeared as a headhunter in the November 14, 1966 "Topsy-Turvey" episode of CBS' "Gilligan's Island." With several of his Rams teammates, he made a cameo appearance as a football player in the 1965 "Perry Mason" episode, "The Case of the 12th Wildcat", as well as in 1970 on an "Ironside" episode, "Blackout". He appeared twice on "Rowan & Martin's Laugh-In". He was also on a 1978 episode of "Wonder Woman", "The Deadly Sting".


title: Jed Mercurio

text: Gerald Gary "Jed" Mercurio (born September 1966) is a British television writer, producer, director and novelist. A former hospital doctor and Royal Air Force officer, Mercurio has been ranked among UK television's leading writers. In 2017, Mercurio was awarded a Fellowship of the Royal Television Society and the Baird Medal by RTS Midlands.


title: Jed Mercurio

text: His works for television include the series "Line of Duty", "Bodyguard", "Bodies" (based on his 2002 novel), "The Grimleys", and "Cardiac Arrest". His books are "Bodies" (2002), "Ascent" (2007), "American Adulterer" (2009) and, for children, "The Penguin Expedition" (2003).


title: Jed Mercurio

text: Mercurio was born in Nelson, Lancashire, but grew up in Cannock, Staffordshire. His parents were Italian immigrants, with his father working as a coal miner. He studied at the University of Birmingham Medical School. During his third year, in August 1988, he was commissioned as a pilot officer in the Royal Air Force Medical Branch. He received flying training with the University Air Squadron, with the intention of specialising in aviation medicine. He was promoted to flying officer in June 1991, but resigned his commission in February 1992.


title: Jed Mercurio

text: During practice as a hospital physician, Mercurio answered an advertisement in the "British Medical Journal" and, despite little writing experience, scripted the BBC medical drama "Cardiac Arrest" under the pseudonym John MacUre. Subsequently he retired from medicine to pursue a writing career under his own name.


title: Jed Mercurio

text: Mercurio's writing debut, "Cardiac Arrest" (1994–96), caused controversy due to its realistic depiction of hospital life. The series was twice nominated in the Best Original Drama category by the Writers' Guild of Great Britain and topped a poll of UK medical professionals as the most realistic medical drama of all time. Mercurio served as medical adviser on the second series of "Cardiac Arrest", which he cites as his apprenticeship in producing/directing.


title: Jed Mercurio

text: Jed Mercurio entered a new genre when he created "The Grimleys", a comedy series set in the Black Country, which ran for three series (1999–2001) on ITV.


title: Jed Mercurio

text: Mercurio adapted his first novel, "Bodies" (2002), into an award-winning television series. Mercurio assumed the producer/showrunner position for the first time on "Bodies" (2004–06), a role he has fulfilled on all his subsequent original series. "The Times" ranked "Bodies" in "Shows of the Decade" and "The Guardian" placed it in "The Greatest TV Dramas of All Time." The series won the Royal Television Society Award for Best Drama Series, was twice nominated for the BAFTA Award for Best Drama Series and Mercurio was also twice nominated as Best Drama Writer by the Royal Television Society.


title: Jed Mercurio

text: His critically acclaimed second novel, "Ascent" (2007), was ranked among the "1000 Novels Everyone Must Read".


title: Jed Mercurio

text: Mercurio's next original drama series, "Line of Duty" (first broadcast 26 June 2012), premiered as BBC2's then best-performing drama series in 10 years with 4.1 million viewers. A well-received second series (first broadcast 12 February 2014) resulted in the BBC commissioning two further series for production in 2015 and 2016. The first series was nominated for a Royal Television Society Award for Best Drama Series and earned Mercurio his third nomination as Best Drama Writer by the Royal Television Society and The Writers' Guild of Great Britain. In 2014, "Line of Duty" was named in the Top 50 BBC2 Shows of All-Time, and named best television drama of the year by "The Observer", and in 2016 ranked among the best BBC shows of all time and among the best police series of all time. In 2015, the second series won the Royal Television Society Award for Best Drama Series. and was nominated for four BAFTA Awards. Series 3 of "Line of Duty" became the most watched drama series broadcast on BBC2 in the multichannel era. Series 4 transferred to BBC One, consisting of six episodes broadcast from 26 March 2017.


title: Jed Mercurio

text: Mercurio's third medical drama series, "Critical", ran for 13 episodes on Sky1 from 24 February 2015 to predominantly positive reviews, but was cancelled on 15 July 2015 after one season.


title: Jed Mercurio

text: Mercurio created and wrote "Bodyguard", serving as showrunner on the six-part series starring Richard Madden and Keeley Hawes. The series began broadcasting on BBC One on 26 August 2018, achieving the highest viewing figures for a new BBC drama in the multichannel era.


title: Jed Mercurio

text: Mercurio was appointed Officer of the Order of the British Empire (OBE) in the 2021 New Year Honours for services to television drama.


title: Jed Mercurio

text: More recently, he and Jimmy Mulville, via HTM Television inked a first look deal with 20th Television in an effort to expand it to the US.


title: Jed Mercurio

text: Mercurio's long-time partner is Elaine Cameron, a producer and script editor, and they have two children; Jack and Molly Mercurio.


title: Battle of Cold Harbor

text: The Battle of Cold Harbor was fought during the American Civil War near Mechanicsville, Virginia, from May 31 to June 12, 1864, with the most significant fighting occurring on June 3. It was one of the final battles of Union Lt. Gen. Ulysses S. Grant's Overland Campaign, and is remembered as one of American history's bloodiest, most lopsided battles. Thousands of Union soldiers were killed or wounded in a hopeless frontal assault against the fortified positions of Confederate Gen. Robert E. Lee's army.


title: Battle of Cold Harbor

text: On May 31, as Grant's army once again swung around the right flank of Lee's army, Union cavalry seized the crossroads of Old Cold Harbor, about 10 miles northeast of the Confederate capital of Richmond, Virginia, holding it against Confederate attacks until the Union infantry arrived. Both Grant and Lee, whose armies had suffered enormous casualties in the Overland Campaign, received reinforcements. On the evening of June 1, the Union VI Corps and XVIII Corps arrived and assaulted the Confederate works to the west of the crossroads with some success.


title: Battle of Cold Harbor

text: On June 2, the remainder of both armies arrived and the Confederates built an elaborate series of fortifications 7 miles long. At dawn on June 3, three Union corps attacked the Confederate works on the southern end of the line and were easily repulsed with heavy casualties. Attempts to assault the northern end of the line and to resume the assaults on the southern were unsuccessful.


title: Battle of Cold Harbor

text: Grant said of the battle in his "Personal Memoirs", "I have always regretted that the last assault at Cold Harbor was ever made. ... No advantage whatever was gained to compensate for the heavy loss we sustained." The armies confronted each other on these lines until the night of June 12, when Grant again advanced by his left flank, marching to the James River. In the final stage, Lee entrenched his army within besieged Petersburg before finally retreating westward across Virginia.


title: Battle of Cold Harbor

text: Grant's Overland Campaign was one of a series of simultaneous offensives the newly appointed general in chief launched against the Confederacy. By late May 1864, only two of these continued to advance: Maj. Gen. William T. Sherman's Atlanta Campaign and the Overland Campaign, in which Grant accompanied and directly supervised the Army of the Potomac and its commander, Maj. Gen. George G. Meade. Grant's campaign objective was not the Confederate capital of Richmond, but the destruction of Lee's army. President Abraham Lincoln had long advocated this strategy for his generals, recognizing that the city would certainly fall after the loss of its principal defensive army. Grant ordered Meade, "Wherever Lee goes, there you will go also." Although he hoped for a quick, decisive battle, Grant was prepared to fight a war of attrition. Both Union and Confederate casualties could be high, but the Union had greater resources to replace lost soldiers and equipment.


title: Battle of Cold Harbor

text: On May 5, after Grant's army crossed the Rapidan River and entered the Wilderness of Spotsylvania, it was attacked by Lee's Army of Northern Virginia. Although Lee was outnumbered, about 60,000 to 100,000, his men fought fiercely and the dense foliage provided a terrain advantage. After two days of fighting and almost 29,000 casualties, the results were inconclusive and neither army was able to obtain an advantage. Lee had stopped Grant, but had not turned him back; Grant had not destroyed Lee's army. Under similar circumstances, previous Union commanders had chosen to withdraw behind the Rappahannock, but Grant instead ordered Meade to move around Lee's right flank and seize the important crossroads at Spotsylvania Court House to the southeast, hoping that by interposing his army between Lee and Richmond, he could lure the Confederates into another battle on a more favorable field.


title: Battle of Cold Harbor

text: Elements of Lee's army beat the Union army to the critical crossroads of Spotsylvania Court House and began entrenching, a tactic that became increasingly essential for the outnumbered defenders. Meade was dissatisfied with Maj. Gen. Philip Sheridan's Union cavalry's performance and released it from its reconnaissance and screening duties for the main body of the army to pursue and defeat the Confederate cavalry under Maj. Gen. J.E.B. Stuart. Sheridan's men mortally wounded Stuart in the tactically inconclusive Battle of Yellow Tavern (May 11) and then continued their raid toward Richmond, leaving Grant and Meade without the "eyes and ears" of their cavalry.


title: Battle of Cold Harbor

text: Near Spotsylvania Court House, fighting occurred on and off from May 8 through May 21, as Grant tried various schemes to break the Confederate line. On May 8, Union Maj. Gens. Gouverneur K. Warren and John Sedgwick unsuccessfully attempted to dislodge the Confederates under Maj. Gen. Richard H. Anderson from Laurel Hill, a position that was blocking them from Spotsylvania Court House. On May 10, Grant ordered attacks across the Confederate line of earthworks, which by now extended over 4 miles (6.5 km), including a prominent salient known as the Mule Shoe. Although the Union troops failed again at Laurel Hill, an innovative assault attempt by Col. Emory Upton against the Mule Shoe showed promise.


title: Battle of Cold Harbor

text: Grant used Upton's assault technique on a much larger scale on May 12 when he ordered the 15,000 men of Maj. Gen. Winfield S. Hancock's corps to assault the Mule Shoe. Hancock was initially successful, but the Confederate leadership rallied and repulsed his incursion. Attacks by Maj. Gen. Horatio G. Wright on the western edge of the Mule Shoe, which became known as the "Bloody Angle," involved almost 24 hours of desperate hand-to-hand fighting, some of the most intense of the Civil War. Supporting attacks by Warren and by Maj. Gen. Ambrose Burnside were unsuccessful. In the end, the battle was tactically inconclusive, but with almost 32,000 casualties on both sides, it was the costliest battle of the campaign. Grant planned to end the stalemate by once again shifting around Lee's right flank to the southeast, toward Richmond.


title: Battle of Cold Harbor

text: Grant's objective following Spotsylvania was the North Anna River, about south. He sent Hancock's Corps ahead of his army, hoping that Lee would attack it, luring him into the open. Lee did not take the bait and beat Grant to the North Anna. On May 23, Warren's V Corps crossed the river at Jericho Mills, fighting off an attack by A.P. Hill's corps, while Hancock's II Corps captured the bridge on the Telegraph Road. Lee then devised a plan, which represented a significant potential threat to Grant: a five-mile (8 km) line that formed an inverted "V" shape with its apex on the river at Ox Ford, the only defensible crossing in the area. By moving south of the river, Lee hoped that Grant would assume that he was retreating, leaving only a token force to prevent a crossing at Ox Ford. If Grant pursued, the pointed wedge of the inverted V would split his army and Lee could concentrate on interior lines to defeat one wing; the other Union wing would have to cross the North Anna twice to support the attacked wing.


title: Battle of Cold Harbor

text: The Union Army assaulted the tip of the apex at Ox Ford and the right wing of the V. However, Lee, incapacitated in his tent by diarrhea, could not effect the attack he hoped to make. Grant realized the situation he was faced with and ordered his men to stop advancing and to build earthworks of their own. The Union general remained optimistic and was convinced that Lee had demonstrated the weakness of his army. He wrote to the Army's chief of staff, Maj. Gen. Henry W. Halleck: "Lee's army is really whipped. ... I may be mistaken but I feel that our success over Lee's army is already assured."


title: Battle of Cold Harbor

text: As he did after the Wilderness and Spotsylvania, Grant planned another wide swing around Lee's flank, marching east of the Pamunkey River to screen his movements from the Confederates. His army disengaged on May 27 and crossed the river. Lee moved his army swiftly in response, heading for Atlee's Station on the Virginia Central Railroad, a point only 9 miles north of Richmond. There, his men would be well-positioned behind a stream known as Totopotomoy Creek to defend against Grant if he moved against the railroads or Richmond. Lee was not certain of Grant's specific plans, however; if Grant was not intending to cross the Pamunkey in force at Hanovertown, the Union army could outflank him and head directly to Richmond. Lee ordered cavalry under Maj. Gen. Wade Hampton to make a reconnaissance in force, break through the Union cavalry screen, and find the Union infantry.


title: Battle of Cold Harbor

text: On May 28, Hampton's troopers encountered Union cavalry under Brig. Gen. David McM. Gregg in the Battle of Haw's Shop. Fighting predominately dismounted and utilizing earthworks for protection, neither side achieved an advantage. The battle was inconclusive, but it was one of the bloodiest cavalry engagements of the war. Hampton held up the Union cavalry for seven hours, prevented it from achieving its reconnaissance objectives, and had provided valuable intelligence to Lee about the disposition of Grant's army.


title: Battle of Cold Harbor

text: After Grant's infantry had crossed to the south bank of the Pamunkey, Lee saw an opportunity on May 30 to attack Warren's advancing V Corps with his Second Corps, now commanded by Lt. Gen. Jubal Early. Early's divisions under Maj. Gens. Robert E. Rodes and Stephen Dodson Ramseur drove the Union troops back in the Battle of Bethesda Church, but Ramseur's advance was stopped by a fierce stand of infantry and artillery fire. On that same day, a small cavalry engagement at Matadequin Creek (the Battle of Old Church) drove an outnumbered Confederate cavalry brigade to the crossroads of Old Cold Harbor, verifying to Lee that Grant intended to move toward that vital intersection beyond Lee's right flank, attempting to avoid another stalemate on the Totopotomoy Creek line.


title: Battle of Cold Harbor

text: Lee received notice that reinforcements were heading Grant's way from Bermuda Hundred. The 16,000 men of Maj. Gen. William F. "Baldy" Smith's XVIII Corps were withdrawn from Maj. Gen. Benjamin Butler's Army of the James at Grant's request, and they were moving down the James River and up the York to the Pamunkey. If Smith moved due west from White House Landing to Old Cold Harbor, southeast of Bethesda Church and Grant's left flank, the extended Federal line would be too far south for the Confederate right to contain. Smith's men arrived at White House May 30–31. One brigade was left behind on guard duty, but about 10,000 men arrived to join Grant's army about 3 p.m. on June 1.


title: Battle of Cold Harbor

text: Lee also received reinforcements. Confederate President Jefferson Davis directed Gen. P.G.T. Beauregard to send the division of Maj. Gen. Robert F. Hoke, over 7,000 men, from below the James River. (The first troops of Hoke's division arrived at Old Cold Harbor on May 31, but were unable to prevent the Union cavalry from seizing the intersection.) With these additional troops, and by managing to replace many of his 20,000 casualties to that point in the campaign, Lee's Army of Northern Virginia had 59,000 men to contend with Meade's and Grant's 108,000. But the disparity in numbers was no longer what it had been—Grant's reinforcements were often raw recruits and heavy artillery troops, pulled from the defenses of Washington, D.C., who were relatively inexperienced in infantry tactics, while most of Lee's had been veterans moved from inactive fronts, and who were soon entrenched in impressive fortifications.


title: Battle of Cold Harbor

text: Grant's Union forces totaled approximately 108,000 men. They consisted of the Army of the Potomac, under Maj. Gen. George Meade, and the XVIII Corps, on temporary assignment from the Army of the James. The six corps were:


title: Battle of Cold Harbor

text: Lee's Confederate Army of Northern Virginia comprised about 59,000 men and was organized into four corps and two independent divisions:


title: Battle of Cold Harbor

text: The battle was fought in central Virginia, in what is now Mechanicsville, over the same ground as the Battle of Gaines's Mill during the Seven Days Battles of 1862. Some accounts refer to the 1862 battle as the First Battle of Cold Harbor, and the 1864 battle as the Second Battle of Cold Harbor. Union soldiers were disturbed to discover skeletal remains from the first battle while entrenching. Cold Harbor was not a port city, despite its name. Rather, it described two rural crossroads named for the Cold Harbor Tavern (owned by the Isaac Burnett family) which provided shelter (harbor) but no hot meals. Old Cold Harbor stood two miles east of Gaines's Mill, and New Cold Harbor a mile southeast. Both were approximately northeast of Richmond, capital of the Confederacy. From these crossroads, the Union army was positioned to receive reinforcements sailing up the Pamunkey River, and could attack either the Confederate capital or its Army of Northern Virginia.


title: Battle of Cold Harbor

text: The cavalry forces that had fought at Old Church continued to face each other on May 31. Lee sent a cavalry division under Maj. Gen. Fitzhugh Lee to reinforce Brig. Gen. Matthew Butler and secure the crossroads at Old Cold Harbor. As Union Brig. Gen. Alfred T. A. Torbert increased pressure on the Confederates, Robert E. Lee ordered Anderson's First Corps to shift right from Totopotomoy Creek to support the cavalry. The lead brigade of Hoke's division also reached the crossroads to join Butler and Fitzhugh Lee. At 4 p.m. Torbert and elements of Brig. Gen. David McM. Gregg's cavalry division drove the Confederates from the Old Cold Harbor crossroads and began to dig in. As more of Hoke's and Anderson's men streamed in, Union cavalry commander Maj. Gen. Philip Sheridan became concerned and ordered Torbert to pull back toward Old Church.


title: Battle of Cold Harbor

text: Grant continued his interest in Old Cold Harbor as an avenue for Smith's arrival and ordered Wright's VI Corps to move in that direction from his right flank on Totopotomoy Creek. He ordered Sheridan to return to the crossroads and secure it "at all hazards." Torbert returned at 1 a.m. and was relieved to find that the Confederates had failed to notice his previous withdrawal.