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.

The first cryptic genus of Ixodida, Cryptocroton n. gen. for Amblyomma papuanum Hirst, 1914: a tick of North Queensland, Australia, and Papua New Guinea.

We describe a new genus Cryptocroton n. gen. for Amblyomma papuanum Hirst, 1914, a tick of North Queensland, Australia, and Papua New Guinea.

Dermacentor (Indocentor) auratus Supino 1897: Potential geographic range, and medical and veterinary significance.

Dermacentor (Indocentor) auratus Supino, 1897 occurs in many regions of Southeast Asia and South Asia. In many regions of Southeast Asia and South Asia, targeted tick sampling and subsequent screening of collected D. auratus ticks have detected pathogenic bacteria and viruses in D. auratus. These disease-causing pathogens that have been detected in D. auratus include Anaplasma, Bartonella, Borrelia, Rickettsia (including spotted fever group rickettsiae), African swine fever virus, Lanjan virus, and Kyasanur forest disease virus. Although D. auratus predominantly infests wild pigs, this tick is also an occasional parasite of humans and other animals. Indeed, some 91 % of human otoacariasis cases in Sri Lanka were due to infestation by D. auratus. With the propensity of this tick to feed on multiple species of hosts, including humans, and the detection of pathogenic bacteria and viruses from this tick, D. auratus is a tick of medical, veterinary, and indeed zoonotic concern. The geographic range of this tick, however, is not well known. Therefore, in the present paper, we used the species distribution model, BIOCLIM, to project the potential geographic range of D. auratus, which may aid pathogen and tick-vector surveillance. We showed that the potential geographic range of D. auratus is far wider than the current geographic distribution of this tick, and that regions in Africa, and in North and South America seem to have suitable climates for D. auratus. Interestingly, in Southeast Asia, Borneo and Philippines also have suitable climates for D. auratus, but D. auratus has not been found in these regions yet despite the apparent close proximity of these regions to Mainland Southeast Asia, where D. auratus occurs. We thus hypothesize that the geographic distribution of D. auratus is largely dependent on the movement of wild pigs and whether or not these wild pigs are able to overcome dispersal barriers. We also review the potential pathogens and the diseases that may be associated with D. auratus and provide an updated host index for this tick.

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 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.

A novel Rickettsia, Candidatus Rickettsia takensis, and the first record of Candidatus Rickettsia laoensis in Dermacentor from Northwestern Thailand.

Three hundred and forty-four tick samples were collected from vegetation at Taksin Maharat National Park, Tak province, northwestern Thailand. They were morphologically identified and molecularly confirmed by 16S rRNA and COI genes as Dermacentor laothaiensis (n = 105), D. steini (n = 139), and D. auratus (n = 100). These ticks were examined for the spotted fever group rickettsiae (SFGRs) using PCR and DNA sequencing of six genes; 17-kDa, gltA, 16S rRNA, ompA, ompB, and sca4. Of these ticks, 6.10% (21/344) gave positive results for the presence of SFGRs. Phylogenetic analyses of the SFGRs clearly indicated that a novel genotype assigned as Candidatus Rickettsia takensis was detected in D. laothaiensis (19/105) and at lesser frequency in D. steini (1/139). Furthermore, Candidatus Rickettsia laoensis was also found at a low frequency in D. auratus (1/100), the first record in Thailand. Although, the pathogenicities of these SFGRs remain unknown, our findings suggest potential risks of SFGRs being transmitted via ticks near the border between Thailand and Myanmar, a gateway of daily migrations of local people and visitors both legal and illegal.

Crimean-Congo hemorrhagic fever virus in Central, Eastern, and South-eastern Asia.

Crimean-Congo hemorrhagic fever (CCHF), caused by Crimean-Congo hemorrhagic fever virus (CCHFV), is endemic in Africa, Asia, and Europe, but CCHF epidemiology and epizootiology is only rudimentarily defined for most regions. Here we summarize what is known about CCHF in Central, Eastern, and South-eastern Asia. Searching multiple international and country-specific databases using a One Health approach, we defined disease risk and burden through identification of CCHF cases, anti-CCHFV antibody prevalence, and CCHFV isolation from vector ticks. We identified 2313 CCHF cases that occurred in 1944-2021 in the three examined regions. Central Asian countries reported the majority of cases (2,026). In Eastern Asia, China was the only country that reported CCHF cases (287). In South-eastern Asia, no cases were reported. Next, we leveraged our previously established classification scheme to assign countries to five CCHF evidence levels. Six countries (China, Kazakhstan, Kyrgyzstan, Tajikistan, Turkmenistan, and Uzbekistan) were assigned to level 1 or level 2 based on CCHF case reports and the maturity of the countries' surveillance systems. Two countries (Mongolia and Myanmar) were assigned to level 3 due to evidence of CCHFV circulation in the absence of reported CCHF cases. Thirteen countries in Eastern and South-eastern Asia were categorized in levels 4 and 5 based on prevalence of CCHFV vector ticks. Collectively, this paper describes the past and present status of CCHF reporting to inform international and local public-health agencies to strengthen or establish CCHFV surveillance systems and address shortcomings.

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.

A new subgenus, Australixodes n. subgen. (Acari: Ixodidae), for the kiwi tick, Ixodes anatis Chilton, 1904, and validation of the subgenus Coxixodes Schulze, 1941 with a phylogeny of 16 of the 22 subgenera of Ixodes Latreille, 1795 from entire mitochondrial genome sequences.

A new subgenus, Australixodes n. subgen., is described for the kiwi tick, Ixodes anatis Chilton, 1904. The subgenus Coxixodes Schulze, 1941 is validated for the platypus tick, Ixodes (Coxixodes) ornithorhynchi Lucas, 1846, sister group of the subgenus Endopalpiger Schulze, 1935. A phylogeny from mitochondrial genomes of 16 of the 22 subgenera of Ixodes (32 spp.) indicates, for the first time, the relationships of the subgenera of Ixodes Latreille, 1795, the largest genus of ticks.

Description of eight new species of Ixodes Latreille, 1795 (Acari: Ixodidae) and redescription of I. auritulus Neumann, 1904, parasites of birds in the Australasian, Nearctic and Neotropical Regions.

Ixodes brevisetosus n. sp. from Papua New Guinea, Ixodes contrarius n. sp. from Costa Rica, Ixodes guglielmonei n. sp. from Argentina and Chile, Ixodes insulae n. sp. from Australia, Ixodes moralesi n. sp. from Guatemala and Panama, Ixodes rio n. sp. from Brazil, Ixodes robbinsi n. sp. from Canada and the United States and Ixodes tinamou n. sp. from Peru (Acari: Ixodidae) are described based on females, nymphs and larvae from various avian hosts (Charadriiformes: Charadriidae; Falconiformes: Falconidae; Galliformes: Cracidae, Odontophoridae, Phasianidae; Passeriformes: Acanthizidae, Cardinalidae, Emberizidae, Fringillidae, Furnariidae, Melanocharitidae, Orthonychidae, Paridae, Passeridae, Petroicidae, Thamnophilidae, Troglodytidae, Turdidae; Strigiformes: Strigidae; Tinamiformes: Tinamidae) and a rodent (Rodentia: Muridae). Females, nymphs and larvae of all new species are similar to those of Ixodes auritulus Neumann, 1904, but can be distinguished by the shape of the scutum and basis capituli, relative length of idiosomal setae, auriculae and internal projection on palpal segment I, degree of development of dorsoapical and mesodorsal spurs on palpal segment I, and measurements and proportions of various structures. Females, nymphs and larvae of I. auritulus s. str. are redescribed based on specimens from passerine birds (Furnariidae, Troglodytidae, Turdidae) in Chile. The subspecies I. auritulus zealandicus Dumbleton, 1961 is elevated to species status, Ixodes zealandicus Dumbleton, 1961. Based on our extensive survey and morphological analyses, we evaluate the actual species richness in the I. auritulus species group and discuss future directions in the systematics of this group of ticks.

The Genus Haemaphysalis (Acari: Ixodidae) in Laos: An Update of Species Records and a Review of Taxonomic Challenges.

Haemaphysalis Koch, 1844, is the largest genus of ticks in Southeast Asia, but little information is available concerning the species present in Laos. Recent research has yielded records for 10 Haemaphysalis species in Laos, including 5 new records, as well as 3 morphological entities of uncertain status. Further morphological and molecular studies are needed to clarify our taxonomic understanding of this genus in Southeast Asia.

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.

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.

Description of the female, nymph and larva and mitochondrial genome, and redescription of the male of Ixodes barkeri Barker, 2019 (Acari: Ixodidae), from the short-beaked echidna, Tachyglossus aculeatus, with a consideration of the most suitable subgenus for this tick.

BACKGROUND: Ixodes barkeri, a tick with a distinctive ventrolateral horn-like projection on palpal segment 1, was described in 2019 from two male ticks from the Wet Tropics of Far North Queensland, Australia. However, females lie at the core of the taxonomy and subgenus classification of Ixodes; hence, we sought specimens of female ticks, successfully recovering females, plus nymphs and larvae. Mitochondrial genomes are also desirable additions to the descriptions of species of ticks particularly regarding subgenus systematics. So, we sequenced the mt genomes of I. barkeri Barker, 2019, and the possible relatives of I. barkeri that were available to us (I. australiensis Neumann, 1904, I. fecialis Warburton & Nuttall, 1909, and I. woyliei Ash et al. 2017) with a view to discovering which if any of the subgenera of Ixodes would be most suitable for I. barkeri Barker, 2019. RESULTS: The female, nymph, larva and mitochondrial genome of Ixodes barkeri Barker, 2019, are described for the first time and the male of I. barkeri is redescribed in greater detail than previously. So far, I. barkeri is known only from a monotreme, the short-beaked echidna, Tachyglossus aculeatus (Shaw, 1792), from the highland rainforests of the Wet Tropics of Far North Queensland, Australia. CONCLUSIONS: Our phylogeny from entire mitochondrial genomes indicated that I. barkeri and indeed I. woyliei Ash et al., 2017, another tick that was described recently, are best placed in the subgenus Endopalpiger Schulze, 1935.

Rediscovery of Ixodes confusus in Australia with the first description of the male from Australia, a redescription of the female and the mitochondrial (mt) genomes of five species of Ixodes.

We: (i) report the rediscovery of Ixodes (Sternalixodes) confusus Roberts, 1960 in Australia; (ii) redescribe the male and female of I. confusus; (iii) describe the mitochondrial (mt) genome of I. confusus from five ticks from four localities in Far North Queensland; and (iv) present the first substantial phylogeny of the subgenera of the Ixodes. The mt genomes of I. confusus, I. cornuatus, I. hirsti, I. myrmecobii and I. trichosuri are presented here for the first time. In our phylogeny from entire mt genomes (ca. 15 kb), the subgenus Endopalpiger was the sister-group to subgenera Sternalixodes plus Ceratixodes plus Exopalpiger whereas Exopalpiger was the sister to Sternalixodes plus Ceratixodes. [i.e. ((Endopalpiger) (Sternalixodes, Ceratixodes and Exopalpiger))]. Finally, we show that Ixodes anatis, the kiwi tick, may be closely related to the ticks of marsupials of Australia and Papua New Guinea.

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.

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.

Epidemiology of Crimean-Congo Hemorrhagic Fever (CCHF) in Africa-Underestimated for Decades.

Crimean-Congo hemorrhagic fever (CCHF) is endemic in Africa, but the epidemiology remains to be defined. Using a broad database search, we reviewed the literature to better define CCHF evidence in Africa. We used a One Health approach to define the impact of CCHF by reviewing case reports, human and animal serology, and records of CCHF virus (CCHFV) isolations (1956-mid-2020). In addition, published and unpublished collection data were used to estimate the geographic distribution of Hyalomma ticks and infection vectors. We implemented a previously proposed classification scheme for organizing countries into five categories by the level of evidence. From January 1, 1956 to July 25, 2020, 494 CCHF cases (115 lethal) were reported in Africa. Since 2000, nine countries (Kenya, Mali, Mozambique, Nigeria, Senegal, Sierra Leone, South Sudan, Sudan, and Tunisia) have reported their first CCHF cases. Nineteen countries reported CCHF cases and were assigned level 1 or level 2 based on maturity of their surveillance system. Thirty countries with evidence of CCHFV circulation in the absence of CCHF cases were assigned level 3 or level 4. Twelve countries for which no data were available were assigned level 5. The goal of this review is to inform international organizations, local governments, and healthcare professionals about shortcomings in CCHF surveillance in Africa to assist in a movement toward strengthening policy to improve CCHF surveillance.

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.