Insights from entire mitochondrial genome sequences into the phylogeny of ticks of the genera Haemaphysalis and Archaeocroton with the elevation of the subgenus Alloceraea Schulze, 1919 back to the status of a genus.

We used entire mitochondrial (mt) genome sequences (14.5-15 kbp) to resolve the phylogeny of the four main lineages of the Haematobothrion ticks: Alloceraea, Archaeocroton, Bothriocroton and Haemaphysalis. In our phylogenetic trees, Alloceraea was the sister to Archaeocroton sphenodonti, a tick of an archetypal reptile, the tuatara, from New Zealand, to the exclusion of the rest of the species of Haemaphysalis. The mt genomes of all four of the Alloceraea species that have been sequenced so far had a substantial insert, 132-312 bp, between the tRNA-Glu (E) gene and the nad1 gene in their mt genomes. This insert was not found in any of the other eight subgenera of Haemaphysalis. The mt genomes of 13 species of Haemaphysalis from NCBI GenBank were added to the most recent data set on Haemaphysalis and its close relatives to help resolve the phylogeny of Haemaphysalis, including five new subgenera of Haemaphysalis not previously considered by other authors: Allophysalis (structurally primitive), Aboimisalis (structurally primitive), Herpetobia (structurally intermediate), Ornithophysalis (structurally advanced) and Segalia (structurally advanced). We elevated Alloceraea Schulze, 1919 to the status of genus because Alloceraea Schulze, 1919 is phylogenetically distinct from the other subgenera of Haemaphysalis. Moreover, we propose that the subgenus Allophysalis is the sister to the rest of the Haemaphysalis (14 subgenera) and that the 'structurally primitive' subgenera Hoogstraal and Kim comprise early diverging lineages. Our matrices of the pairwise genetic difference (percent) of mt genomes and partial 16S rRNA sequences indicated that the mt genome sequence of Al. kitaokai (gb# OM368280) may not be Al. kitaokai Hoogstraal, 1969 but rather another species of Alloceraea. In a similar way, the mt genome sequence of H. (Herpetobia) nepalensis Hoogstraal, 1962 (gb# NC_064124) was only 2% genetically different to that of H. (Allophysalis) tibetensis Hoogstraal, 1965 (gb# OM368293): this indicates to us that they are the same species. Alloceraea cretacea may be better placed in a genus other than Alloceraea Schulze, 1919. Reptiles may have been the host to the most recent common ancestor of Archaeocroton and Alloceraea.

Description of a new species of Ixodes Latreille, 1795 (Acari: Ixodidae), parasite of rodents (Rodentia: Muridae) in Malaysia and Vietnam.

Ixodes abramovi n. sp. (Acari: Ixodidae) is described based on females ex various rodents (Rodentia: Muridae) from Malaysia and Vietnam. Females of this new species are similar to those of some other Oriental species of the subgenus Ixodes Latreille, 1795 such as I. granulatus Supino, 1897 and I. werneri Kohls, 1950, from which they can be distinguished by the size of scutum, length of cornua, shape and length of auriculae and length of spurs on coxae.

Redescription of adults and first description of the larva of Haemaphysalis (Rhipistoma) adleri Feldman-Muhsam, 1951 (Acari: Ixodidae), parasite of carnivorans in Western Asia.

The adults of Haemaphysalis (Rhipistoma) adleri Feldman-Muhsam, 1951 (Acari: Ixodidae) are redescribed and the larva of this species is described for the first time here. The adults of H. adleri that we studied were collected from various canid, felid and hyaenid carnivorans (Carnivora: Canidae, Felidae, Hyaenidae) as well as a hedgehog (Erinaceomorpha: Erinaceidae) in Iraq, Israel and West Bank. The males, females and larvae of H. adleri can be differentiated from Haemaphysalis (Rhipistoma) species occurring in the Palearctic portion of West Asia and Egypt as well those in the H. asiatica subgroup by the length of idiosomal setae, development and size of spurs on palpi, dental formula on the hypostome and size of spur on coxae. A lectotype of H. adleri has been designated and the geographic distribution and hosts of this tick species are discussed.

Reinstatement of Haemaphysalis (Rhipistoma) dentipalpis Warburton & Nuttall, 1909 (Acari: Ixodidae) as a valid species and redescription of adults of H. (R.) asiatica (Supino, 1897), parasites of carnivorans in Oriental Asia.

Haemaphysalis (Rhipistoma) dentipalpis Warburton & Nuttall, 1909 (Acari: Ixodidae) is reinstated here as a valid species and the male is redescribed whereas the female is described for the first time. The adults of H. dentipalpis that we studied were collected from various felid and viverrid carnivorans (Carnivora: Felidae, Viverridae) in Indonesia and Malaysia. For comparative purposes, the male and female of H. (R.) asiatica (Supino, 1897) are redescribed. The adults of H. asiatica that we studied were from various felid and viverrid carnivorans (Carnivora: Felidae, Viverridae) as well as a treeshrew (Scandentia: Tupaiidae) in Thailand and Vietnam. The males and females of both H. dentipalpis and H. asiatica can be differentiated by the pattern of punctations on the conscutum and scutum and the shape and size of the posterodorsal and posteroventral spurs on palpal segment II.

Description of a new species of Ixodes Latreille, 1795 (Acari: Ixodidae), parasite of shrew tenrecs (Afrotheria: Tenrecidae) and rodents (Rodentia: Muridae) on Madagascar.

Ixodes (Afrixodes) ambohitantelensis n. sp. (Acari: Ixodidae) is described based on females ex endemic shrew tenrecs (Afrosoricida: Tenrecidae) and an introduced rodent (Rodentia: Muridae) from Madagascar. Females of this new species are similar to those of other species of the subgenus Afrixodes Morel, 1966, known from Madagascar, from which they can be distinguished by the size of scutum, size of scutal setae, shape of alloscutal setae, development of genital apron, size of auriculae, size of anterior angle of basis capituli, size of palpi, dental formula on hypostome, development of syncoxae, and size and development of spurs on coxae I and IV.

Identification of a new species of Ixodes Latreille, 1795 (Acari: Ixodidae), parasite of hog badgers (Carnivora: Mustelidae) in China.

Ixodes hunanensis n. sp. (Acari: Ixodidae), is identified based on the morphological characteristics and molecular biological analyses of males and females ex hog badger, Arctonyx collaris Cuvier (Carnivora: Mustelidae) from China. Adults of this new species are similar to those of other species of the subgenus Pholeoixodes Schulze, 1942, from which they can be distinguished by the shape of basis capituli, development of cornua, size of porose areas, shape, and size of spurs on coxae and phylogenetic analyses of the cox1 and 16S rRNA sequences.

Haemaphysalis hoodi (Acari: Ixodidae) on a human from Yaoundé, Cameroon, and its molecular characterization.

The genus Haemaphysalis Koch, 1844 (Acari: Ixodidae) is the second-largest genus, with more than 170 described species that primarily parasitize mammals and birds (Guglielmone et al. 2014, Guglielmone et al. 2020). Haemaphysalis species are three-host ticks, mainly distributed in southern and southeastern Asia and tropical Africa (Guglielmone et al. 2014). The present study identified a tick, Haemaphysalis hoodi Warburton & Nuttall, 1909, collected from a human in Yaoundé, Cameroon. This tick species feed on birds in sub-Saharan Africa. To the best of our knowledge, this is the second record of H. hoodi from humans. In addition, 16S ribosomal RNA and cytochrome oxidase I sequences were generated for this species for the first time. Screening pan-Rickettsia-PCR infection gave a negative result.

Adaptive radiation and speciation in Rhipicephalus ticks: A medley of novel hosts, nested predator-prey food webs, off-host periods and dispersal along temperature variation gradients.

Rhipicephalus are a species-diverse genus of ticks, mainly distributed in the Afrotropics with some species in the Palearctic and Oriental regions. Current taxonomic consensus comprise nine informal species groups/lineages based on immature morphology. This work integrates biogeographic, ecological and molecular lines of evidence to better understand Rhipicephalus evolution. Phylogenetic analysis based on four genes (12S, 16S, 28S-D2 and COI) recovered five distinct clades with nine descendant clades that are generally congruent with current taxonomy, with some exceptions. Historical biogeography is inferred from molecular divergence times, ancestral distribution areas, host-use and climate niches of four phylogenetically significant bioclimatic variables (isothermality, annual, seasonal and diurnal temperature range). Novel hosts enabled host-linked dispersal events into new environments, and ticks exploited new hosts through nested predator-prey connections in food webs. Diversification was further induced by climate niche partitioning along gradients in temperature range during off-host periods. Ancestral climate niche estimates corroborated dispersal events by indicating hypothetical ancestors moved into environments with different annual and seasonal temperature ranges along latitudinal gradients. Host size for immature and adult life stages was important for dispersal and subsequent diversification rates. Clades that utilise large, mobile hosts (ungulates and carnivores) early in development have wider geographic ranges but slower diversification rates, and those utilising small, less mobile hosts (rodents, lagomorphs and afroinsectivores) early in development have smaller ranges but higher diversification rates. These findings suggest diversification is driven by a complex set of factors linked to both host-associations (host size, ranges and mobility) and climate niche partitioning along annual and seasonal temperature range gradients that vary with latitude. Moreover, competitive interactions can reinforce these processes and drive speciation. Off-host periods facilitate adaptive radiation by enabling host switches along nested predator-prey connections in food webs, but at the cost of environmental exposure that partitions niches among dispersing progenitors, disrupting geneflow and driving diversification. As such, the evolution and ecological niches of Rhipicephalus are characterised by trade-offs between on- and off-host periods, and these trade-offs interact with nested predator-prey connections in food webs, host-use at different life stages, as well as gradients in annual and seasonal temperature ranges to drive adaptive radiation and speciation.

Reinstatement of Dermacentor tricuspis (Schulze, 1933) n. comb., n. stat. (Acari: Ixodidae) as a valid species, synonymization of D. atrosignatus Neumann, 1906 and description of a new species from Indonesia, Malaysia and Thailand.

Re-examination of the holotype of Dermacentor atrosignatus Neumann, 1906 (Acari: Ixodidae) stored in the Natural History Museum (London, UK) revealed that this taxon is identical with D. auratus Supino, 1897 and should be treated as a junior synonym of the latter species. A correct name for the distinct species previously identified as D. atrosignatus Neumann, 1906 sensu Wassef & Hoogstraal, 1984 should be D. tricuspis (Schulze, 1933) n. comb., n. stat. Adults of D. tricuspis are redescribed here. Re-examination of extensive holdings of Oriental Dermacentor Koch, 1844 ticks stored in the United States National Tick Collection revealed that a morphologically distinct new species of this genus, namely D. falsosteini D. Apanaskevich, M. Apanaskevich & Nooma n. sp. should be recognized. Adults of D. tricuspis and D. falsosteini n. sp. can be distinguished from other species of Oriental Dermacentor and each other by the colour pattern of the conscutum and scutum, the pattern of punctations on the pseudoscutum and scutum, the shape of female genital structures and spurs on coxa I. Dermacentor tricuspis is recorded from Indonesia, Malaysia, the Philippines and Thailand where the adults were mostly collected from various species of wild pigs (Artiodactyla: Suidae) and vegetation; few adults were available from other mammals (Artiodactyla: Bovidae; Carnivora: Canidae, Felidae, Ursidae; Pholidota: Manidae), as well as humans and reptiles (Squamata: Elapidae, Varanidae). One male was reared from a nymph collected on a rodent (Rodentia: Muridae). Dermacentor falsosteini n. sp. is found in Indonesia, Malaysia and Thailand where the adults were collected from bearded pig, Sus barbatus Müller, wild boar, S. scrofa Linnaeus, unidentified wild pig, Sus sp. (Artiodactyla: Suidae), Malayan tapir, Tapirus indicus Desmarest (Perissodactyla: Tapiridae), human and vegetation.

Morphological and molecular identification of medically important questing Dermacentor species collected from some recreational areas of Peninsular Malaysia.

Questing is a situation when a tick is seeking to get closer or ambush its potential host. However, information on questing tick species in Malaysia is still lacking, thus the association with tick-borne diseases (TBD) is not completely understood. The aim of this study was to investigate the tick species from five most frequently visited recreational areas in Pahang and Terengganu states, which were recorded to have high potential of TBD cases. By implementing handpick method, a total of 18 males and 15 females belonging to five Dermacentor Koch, 1844 species, were collected, namely D. compactus Neumann 1901, D. tricuspis (Schulze, 1933), D. auratus Supino 1897, D. steini (Schulze, 1933), and D. falsosteini Apanaskevich, Apanaskevich & Nooma respectively. The specimens were collected and identified based on morphological characters prior to obtaining the molecular data of COI and 16S rDNA. The D. compactus was the most abundant species collected in this study, while D. falsosteini was the least. All species were distinctly separated on the Neighbor Joining and Maximum Parsimony tree topologies and supported with high bootstrap values. Furthermore, a low intraspecific variation (0.00 - 0.01) was observed amongst the individuals of the same species in both genes. Meanwhile, each Dermacentor species was genetically different, with interspecific values ranging from 0.13-0.19 and 0.11-0.20 for COI and 16S rDNA. These findings had successfully recorded the tick species that were potentially associated with TBD, and which might be circulated among humans and animals. This study also has some implications on the diversity and geographical extension of Dermacentor ticks, thus should warrant further investigation as a potential vector of tick-borne diseases and public health importance.

Description of a new species of Haemaphysalis Koch, 1844 (Acari: Ixodidae) from the H. (Rhipistoma) asiatica subgroup, parasite of an endemic Malagasy carnivoran (Carnivora: Eupleridae).

Haemaphysalis (Rhipistoma) galidiae n. sp. (Acari: Ixodidae) is described herein based on males and a single female. Adults of H. galidiae n. sp. were collected from the ring-tailed vontsira, Galidia elegans Geoffroy Saint-Hilaire (Carnivora: Eupleridae) on Madagascar. Males and the single female of this new species morphologically are most close to H. obtusa Dönitz, 1910, known from various Malagasy euplerid carnivorans, including G. elegans, as well as introduced carnivorans, but can be differentiated from it by the shape of the female genital aperture, shape of posterolateral margin of palpal segment II ventrally in both male and female and shape and size of spurs on palpal segment II, coxa I and trochanter I in both sexes.

Description of three new species of Ixodes Latreille, 1795 (Acari: Ixodidae), parasites of tenrecs (Afrotheria: Tenrecidae) on Madagascar.

Ixodes soarimalalae n. sp., Ixodes uilenbergi n. sp. and Ixodes uncus n. sp. (Acari: Ixodidae), are described based on females ex various species of tenrecs (Afrosoricida: Tenrecidae) from Madagascar. Females of all of these new species are similar to those of other species of the subgenus Afrixodes Morel, 1966 known from Madagascar, from which they can be distinguished and from one other by the size of scutum, size of scutal setae, shape of alloscutal setae, shape of genital aperture, development of genital apron, size of auriculae, size of anterior angle of basis capituli, size of palpi, dental formula on hypostome and size and development of spurs on coxa I.

Description of two new species of Haemaphysalis Koch, 1844 (Acari: Ixodidae) from the H. (Rhipistoma) spinulosa subgroup, parasites of carnivores and other mammals in Africa.

Haemaphysalis (Rhipistoma) latitudinis n. sp. (Acari: Ixodidae) and H. (R.) setosa n. sp. are described based on adults. Adults of H. (R.) latitudinis n. sp. were mostly collected from carnivores (Carnivora: Canidae, Felidae, Herpestidae, Mustelidae, Viverridae), few specimens were found on rodents (Rodentia: Muridae), bat (Chiroptera: Molossidae) and domestic cattle (Artiodactyla: Bovidae) in Benin, Central African Republic, Côte d'Ivoire, Democratic Republic of the Congo, Ethiopia, Ghana, Kenya, Nigeria, South Sudan and Uganda. Adults of H. (R.) setosa n. sp. were mostly found on carnivores (Carnivora: Felidae, Herpestidae, Viverridae), few specimens were found on rodents (Rodentia: Muridae), hare (Lagomorpha: Leporidae), hyrax (Hyracoidea: Procaviidae) and human (Primates: Hominidae) in Kenya. Males and females of both species can be differentiated from each other and other H. spinulosa-like ticks by their size, pattern of punctations on conscutum/scutum, size of setae, shape of genital structures, size and shape of spiracular plates, dorsal cornua, posterodorsal and posteroventral spurs on palpal segment II and spur on coxae and dentition on hypostome.

Description of five new species of Ixodes Latreille, 1795 (Acari: Ixodidae) and redescription of I. luxuriosus Schulze, 1935, I. steini Schulze, 1935 and I. zaglossi Kohls, 1960, parasites of marsupials, rodents and echidnas in New Guinea Island.

Ixodes acer n. sp., Ixodes giluwensis n. sp., Ixodes mirzai n. sp., Ixodes planiscutatus n. sp. and Ixodes stellae n. sp. (Acari: Ixodidae), are described based on females ex various marsupials (Dasyuromorphia: Dasyuridae, Diprotodontia: Petauridae, Peramelemorphia: Peramelidae), rodents (Rodentia: Muridae) and a bird (Galliformes: Megapodiidae) from Indonesia (Papua Province) and Papua New Guinea. Females of all new species are similar to those of I. luxuriosus Schulze, 1935, I. steini Schulze, 1935 and I. zaglossi Kohls, 1960, but can be distinguished from them and each other by the distinctness of lateral carinae and cervical grooves on the scutum, length of setae on the idiosoma, sculpture of the basis capituli and palpal segment I dorsally, size of the dorsal cornua, shape and sculpture of the palpal segment I ventrally, size of ventral spur on the palpal segment I, dental formula on the hypostome and measurements of various structures and their proportions. For comparative purposes the female of I. luxuriosus, I. steini and I. zaglossi is redescribed and lectotypes for I. luxuriosus and I. steini have been designated. Studied females of I. luxuriosus, I. steini and I. zaglossi were found on marsupials (Dasyuromorphia: Dasyuridae; Diprotodontia: Macropodidae, Petauridae, Phalangeridae, Pseudocheiridae; Peramelemorphia: Peramelidae), rodents (Rodentia: Muridae) and echidnas (Monotremata: Tachyglossidae) in Indonesia (Papua Province) and Papua New Guinea.

Description of a new species of Dermacentor Koch, 1844 (Acari: Ixodidae) from the mountains of Laos and Thailand.

Dermacentor pasteuri n. sp. (Acari: Ixodidae) is described based on adults ex wild boar and vegetation from Laos and Thailand. Adults of D. pasteuri n. sp. are similar to those of D. compactus Neumann, 1901, but can be distinguished by the shape of conscutum in the male, development of dorsal cornua in the female, size and shape of spurs on coxae and coloration of leg segments in both sexes.

Description of four new species of Haemaphysalis Koch, 1844 (Acari: Ixodidae) from the H. (Rhipistoma) spinulosa subgroup, parasites of carnivores and rodents in Africa.

Haemaphysalis (Rhipistoma) bochkovi n. sp., H. (R.) burkinae n. sp., H. (R.) horaki n. sp. and H. (R.) walkerae n. sp. (Acari: Ixodidae), are described based on males and females. Adults of H. bochkovi n. sp. were collected from the marsh mongoose, Atilax paludinosus (Cuvier) (Carnivora: Herpestidae) and the spotted-necked otter, Hydrictis maculicollis (Lichtenstein) (Carnivora: Mustelidae) in Côte d'Ivoire and Liberia; adults of H. burkinae n. sp. were collected from the rusty-spotted genet, Genetta maculata (Gray), the Haussa genet, Genetta thierryi Matschie (Carnivora: Viverridae) and the Cape hare, Lepus capensis Linnaeus (Lagomorpha: Leporidae) in Burkina Faso; adults of H. horaki n. sp. were collected from the Cape dune mole-rat, Bathyergus suillus (Schreber) (Rodentia: Bathyergidae) in South Africa; adults of H. walkerae n. sp. were collected from the striped polecat, Ictonyx striatus (Perry) (Carnivora: Mustelidae), the bat-eared fox, Otocyon megalotis (Desmarest) (Carnivora: Canidae), the aardwolf, Proteles cristata (Sparrman) (Carnivora: Hyaenidae) and the East African spring hare, Pedetes surdaster (Thomas) (Rodentia: Pedetidae) in Kenya. Males and females of these new species can be differentiated from each other and other H. spinulosa-like ticks by their size, pattern of punctations on conscutum/scutum, length of setae, shape and size of genital structures, spiracular plates, basis capituli, dorsal cornua, posterosdorsal and posteroventral spurs of palpal segment II and spur on coxae.

Two new species of Haemaphysalis Koch, 1844 (Acari: Ixodidae) in the H. (Rhipistoma) spinulosa subgroup, parasites of carnivores and hedgehogs in Africa.

Haemaphysalis (Rhipistoma) princeps n. sp. (Acari: Ixodidae) and H. (R.) camicasi n. sp. are described based on adults. Adults of H. princeps n. sp. were mostly collected from various carnivores (Carnivora: Canidae, Felidae, Herpestidae, Mustelidae, Viverridae) and hedgehogs (Erinaceomorpha: Erinaceidae); few specimens were found on hare (Lagomorpha: Leporidae), various rodents (Rodentia: Muridae) and an antelope (Artiodactyla: Bovidae) in Benin, Burkina Faso, Cameroon, Chad, Côte d'Ivoire, Ghana, Mali, Nigeria, Senegal, South Sudan and Sudan. Few adults of H. princeps n. sp. were reared from nymphs collected on rodents (Rodentia: Muridae) and shrew (Soricomorpha: Soricidae). Adults of H. camicasi n. sp. were also mostly collected from various carnivores (Carnivora: Canidae, Felidae, Herpestidae, Mustelidae, Viverridae) and hedgehogs (Erinaceomorpha: Erinaceidae); few specimens were found on different rodents (Rodentia: Muridae, Nesomyidae, Sciuridae), hare (Lagomorpha: Leporidae) and bushbaby (Primates: Galagidae) in Benin, Burkina Faso, Cameroon, Central African Republic, Chad, Côte d'Ivoire, Nigeria, Senegal, South Sudan and Togo. Males and females of both species can be differentiated from each other and other H. spinulosa-like ticks by their size, pattern of punctations on conscutum/scutum, size of setae, shape of genital structures, size and shape of spiracular plates, dorsal cornua, posterodorsal and posteroventral spurs on palpal segments II and spur on coxae.

Description of a new species of Dermacentor Koch, 1844 (Acari: Ixodidae) from Laos and Thailand.

Dermacentor laothaiensis n. sp. (Acari: Ixodidae) is described based on adults ex wild boar and vegetation from Laos and Thailand. Adults of D. laothaiensis n. sp. are similar to those of D. bellulus (Schulze, 1935) and D. steini (Schulze, 1933) but can be distinguished by the conscutum shape, colour pattern of the conscutum and scutum, the density of punctations on the pseudoscutum and scutum and the shape of female genital structures.

Description of a new species of Amblyomma Koch, 1844 (Acari: Ixodidae), parasite of deer (Artiodactyla: Cervidae) and wild pigs (Artiodactyla: Suidae) in the Philippines.

Amblyomma anicornuta n. sp. (Acari: Ixodidae) is described based on adults and nymphs ex deer (Artiodactyla: Cervidae) and wild pigs (Artiodactyla: Suidae) from Luzon, Philippines. Adults of A. anicornuta n. sp. are similar to those of several Asian and Australasian species of Amblyomma Koch, 1844 with a 4/4 dental formula on the hypostome but can be distinguished by the colouration and pattern of punctations on the conscutum in the male and scutum in the female, the absence of a marginal groove on the conscutum in the male, the possession of long, thick, prominent setae on the alloscutum in the female, projections on anal valves and sclerotised ring around them in the male, a large median sclerite ventrally in the male, as well by the shape of the genital aperture in the female and the size and shape of spurs on coxae I-IV in both sexes. The nymph of A. anicornuta n. sp. is somewhat similar to that of A. babirussae Schulze, 1933 and A. geoemydae (Cantor, 1847) but can be distinguished by the colouration pattern on the scutum, the presence of dorsal cornua and the size of the spurs on coxae I-IV.

Description of a new species of Ixodes Latreille, 1795 (Acari: Ixodidae) and redescription of I. priscicollaris Schulze, 1932, parasites of New Guinea rodents (Rodentia: Muridae).

Ixodes goliath n. sp. (Acari: Ixodidae), is described based on females collected from the eastern hyomys, Hyomys goliath (Milne-Edwards) (Rodentia: Muridae) from Papua New Guinea. Females of I. goliath n. sp. are similar to those of I. priscicollaris Schulze, 1932 but can be distinguished by the overall size, porose areas sculpture and shape of palpal segment I ventrally. For comparative purposes, the female of I. priscicollaris is redescribed. Studied females of I. priscicollaris were found on murid rodents and Phascogale sp. (Dasyuromorphia: Dasyuridae) from Indonesia (Papua Province) and Papua New Guinea.