Publications by authors named "Ralph E Harbach"

80 Publications

A case of two spellings for Humboldt's blue-winged mosquito (Diptera: Culicidae).

Zootaxa 2021 May 19;4974(1):191192. Epub 2021 May 19.

J. Linsley Gressitt Center for Entomological Research, Bishop Museum, Honolulu, Hawai'i 96817, USA..

While adding taxon names to the Systema Dipterorum database (Evenhuis Pape 2021), one of us (NLE) discovered that Humboldt (1819) had spelled the proposed name of a nominal mosquito species in two ways. He described the species, which was found in swampy places along the Magdalena River near Tenerife, Colombia, as Culex cyanopennis on page 340 and afterwards referred to it as Culex cyanopterus on pages 345 and 349. Both names have the same meaning: cyano- (Gr. kyanos, dark blue), pennis (L. penna, feather, wing) and pteron (Gr. feather, wing). The species was named for the perceived color of the wings: "Alæ cæruleæ, splendore semi-metallico…" (wings blue, a bright semi-metallic). On page 345, Humboldt states, translated from the French: "We have been informed in the Rio de la Magdalena that in Simitì no other Culex than the jejen [je·jén: Sp., gnat, mosquito] was known in the past. You can spend the night there quietly, because the jejen is not a nocturnal insect. Since the year 1801, the big blue-winged mosquito (Culex cyanopterus) has shown itself in such abundance that the poor inhabitants of Simitì do not know how to get a peaceful sleep." Thus, in addition to having the same meaning, the two names are associated with the same locality.
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http://dx.doi.org/10.11646/zootaxa.4974.1.10DOI Listing
May 2021

Culex (Culex) longitubus, A New Species of the Mimeticus Subgroup (Diptera: Culicidae) From Bhutan.

J Med Entomol 2021 May 13. Epub 2021 May 13.

Department of Life Sciences, Natural History Museum, Cromwell Road, London SW7 5BD,UK.

Culex longitubus Somboon, Namgay & Harbach is described as a new species of the Mimeticus Subgroup of the subgenus Culex. The larva is most similar to the larva of Cx. tianpingensis Chen from China, but is distinguished by the length of the siphon and the anal papillae, the form of the comb scales and pecten spines, and the development of setae 7-P, 13-T, 1-X, and 4-X. The adults have wing markings and male genitalia similar to those of species of the Mimeticus Complex. Phylogenetic analysis of COI sequences revealed that the new species is closely related to Cx. murrelli Lien of the Mimulus Complex. The immature stages of the new species were found in stagnant pools and marshes at high altitudes in several districts of Bhutan.
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http://dx.doi.org/10.1093/jme/tjab080DOI Listing
May 2021

Culex/ mosquitoes (Diptera: Culicidae) recorded along the Nile River in central and northern Sudan, with a key for the identification of all species of the genus known to occur in the country.

Zootaxa 2021 Apr 20;4963(3):zootaxa.4963.3.1. Epub 2021 Apr 20.

Vector Genetics and Control Laboratory, Department of Zoology, Faculty of Science, University of Khartoum, Khartoum, Sudan..

Despite the importance of Culex species as major vectors of Rift Valley fever virus, West Nile virus and the microfilariae that cause lymphatic filariasis, information on these mosquitoes in Sudan is limited to works published 65 years ago in the former Anglo-Egyptian Sudan, where some species were only recorded from areas of the territory now known as South Sudan. In this paper, we provide updated information on Culex mosquitoes collected indoors during surveillance studies conducted along the Nile River in central and northern areas of Sudan between 2012 and 2019. Of 3,411 female mosquitoes collected in Khartoum and northern states along the river, 2,560 (75%) were specimens of Culex belonging to 12 species: Cx. (Culex) antennatus (Becker, 1903), Cx. (Cux.) laticinctus Edwards, 1913, Cx. (Cux.) neavei Theobald, 1906, Cx. (Cux.) pipiens Linnaeus, 1758, Cx. (Cux.) perexiguus Theobald, 1903, Cx. (Cux.) poicilipes (Theobald, 1903), Cx. (Cux.) quinquefasciatus Say, 1823, Cx. (Cux.) simpsoni Theobald, 1905, Cx. (Cux.) sinaiticus Kirkpatrick, 1925, Cx. (Cux.) theileri Theobald, 1903, Cx. (Cux.) tritaeniorhynchus Giles, 1901 and Cx. (Culiciomyia) macfiei Edwards, 1923. This is the first record for Cx. tritaeniorhynchus and Cx. macfiei in central Sudan. The relative abundance of each species varied in different areas and seasons, but Cx. antennatus and Cx. quinquefasciatus were the most abundant indoor resting species. We provide an updated dichotomous key for the identification of the adults of Culex mosquitoes known to occur in the Republic of the Sudan.
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http://dx.doi.org/10.11646/zootaxa.4963.3.1DOI Listing
April 2021

Culex bhutanensis, a new species of the Mimeticus Subgroup of the nominotypical subgenus of the genus Culex Linnaeus (Diptera: Culicidae) from Bhutan.

Acta Trop 2021 May 19;217:105868. Epub 2021 Feb 19.

Department of Life Sciences, Natural History Museum, Cromwell Road, London SW7 5BD, UK.

Culex bhutanensis Somboon, Namgay & Harbach is described as a new species of the Mimeticus Subgroup of the subgenus Culex Linnaeus. Based on morphological distinctions and phylogenetic analysis of COI sequences, the new species is a member of the Mimulus Complex. The adults have wing markings similar to those of Cx. mimulus Edwards and Cx. murrelli Lien, but are distinguished by the presence of a line of pale scales or scattered pale scales along the posterior margin of the proximal portion of the costa. The larva is most similar to the larva of Cx. orientalis Edwards from Japan and South Korea, but is distinguished by the form of the comb scales, pecten spines and seta 2-X. The pupa differs from the pupae of other species of the subgroup in having seta 8-C with more branches. The immature stages of the new species were found in rice paddies, stagnant pools and marshes at high altitudes in several districts of Bhutan.
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http://dx.doi.org/10.1016/j.actatropica.2021.105868DOI Listing
May 2021

Reinertia, a New Subgenus of the Genus Aedes Meigen and Its Type Species Aedes (Reinertia) suffusus (Diptera: Culicidae), Newly Recorded From Bhutan.

J Med Entomol 2021 05;58(3):1138-1148

Department of Life Sciences, Natural History Museum, London, UK.

A new subgenus, Reinertia Somboon, Namgay & Harbach, of the genus Aedes Meigen and its type species, Ae. suffusus Edwards, are described from specimens reared from larvae and pupae found in a tree hole in Bhutan. The scutum of the adults is mostly covered with narrow pale falcate scales. The proboscis, maxillary palpus, tibiae, and tarsi are dark-scaled. The gonocoxite of the male genitalia bears a unique setose basomesal sclerite. The larva closely resembles larvae of the subgenus Downsiomyia Vargus in having setae 4-6-C with numerous branches and inserted more or less on level with seta 7-C, abdominal seta 12-I is present and the comb is composed of 6-10 spine-like scales arranged in an irregular row. Surprisingly, Reinertia shares features of the adult habitus, male genitalia, and larva with the Palearctic subgenus Dahliana Reinert, Harbach & Kitching. However, in phylogenetic analyses of the mitochondrial COI gene of species representing 38 subgenera of Aedes and six other genera of the tribe Aedini Neveu-Lemaire, Reinertia was not associated with Dahliana or Downsiomyia. In both maximum likelihood and Bayesian analyses of the data, Ae. suffusus was recovered as the weakly supported sister of a clade composed of five species of the subgenus Protomacleaya Theobald. In the absence of strong support, and because Protomacleaya is an unnatural group of species that resemble each other phenetically by virtue of what they lack, Ae. suffusus cannot be placed in the subgenus Protomacleaya. Thus, the morphological and molecular data attest the uniqueness of Ae. suffusus and its recognition as a monobasic subgeneric lineage.
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http://dx.doi.org/10.1093/jme/tjaa277DOI Listing
May 2021

A Multiplex PCR Based on Mitochondrial Sequences for Identification of Members of the Complex (Diptera: Culicidae) in Thailand and Other Countries in the Region.

Insects 2020 Jul 2;11(7). Epub 2020 Jul 2.

Center of Insect Vector Study, Department of Parasitology, Faculty of Medicine, Chiang Mai University, Chiang Mai 50200, Thailand.

A multiplex-PCR assay based on mitochondrial cytochrome c oxidase subunit I () sequences was developed for identification of five members of the Barbirostris Complex which occur in Thailand: s.s., , , and species A3. was not included in the assay due to the lack of unequivocal sequences. Allele-specific primers were designed for specific nucleotide segments of sequences of each species. Mismatch method and addition of long GC tail were applied for some primers. The assay provided products of 706 bp for s.s., 238 bp for , 611 bp for , 502 bp for and 365 bp for A3. The assay was tested using 111 wild-caught female mosquitoes from Bhutan, Cambodia, Indonesia (Sulawesi) and Thailand. The results of the multiplex PCR were in complete agreement with sequencing; however, one of three specimens from Bhutan and all 11 specimens from Indonesia were not amplifiable by the assay due to their distinct sequences. This, together with the distinct rDNA sequences of these specimens, suggests the presence of at least two additional new species in the Barbirostris Complex.
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http://dx.doi.org/10.3390/insects11070409DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC7412068PMC
July 2020

Molecular and morphological evidence of sibling species in Anopheles baileyi Edwards (Diptera: Culicidae) in Bhutan and Thailand.

Acta Trop 2020 Sep 28;209:105549. Epub 2020 May 28.

Department of Life Sciences, Natural History Museum, Cromwell Road, London SW7 5BD, UK.

This paper reports the results of a molecular and morphological study of Anopheles baileyi in Bhutan and Thailand. Phylogenetic analyses of ribosomal (ITS2) and mitochondrial DNA (COI) sequences revealed the presence of four genetically distinct clades, three in Bhutan (Clades I, II and III) and one in Thailand (Clade IV). Most of the larvae in the Bhutanese clades differed from those in the Thai clade in having seta 4-C branched, whereas it is single in the latter. The adults of each clade showed variation of wing markings and overlapping characters. The combination of characteristics of thoracic setae 1,2-P and abdominal seta 3-I was found to be useful for distinguishing the larvae. Pupae were inseparable. We provisionally recognize mosquitoes of Clades I, II, III and IV as members of a sibling species complex, the Baileyi Complex, denoted as species A, B, C and D, respectively. Species A is most likely the type form of An. baileyi s.s. because it was found adjacent to the type locality (Yatung, Tibet), and the others are unrecognized species.
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http://dx.doi.org/10.1016/j.actatropica.2020.105549DOI Listing
September 2020

Integrated systematics of Anopheles subpictus (Diptera: Culicidae) in the Oriental Region, with emphasis on forms in Thailand and Sulawesi, Indonesia.

Acta Trop 2020 May 11;208:105503. Epub 2020 May 11.

Center of Insect Vector Study, Department of Parasitology, Faculty of Medicine, Chiang Mai University, Chiang Mai, Thailand. Electronic address:

The Anopheles subpictus complex consists of four species informally designated, based on fixed inversions of polytene chromosomes and morphology, as species A, B, C and D in India. However, recent studies revealed the presence of only species A and B in Sri Lanka. Little is known about the specific identity of the taxon in other countries in Asia. This paper reports the results of a molecular and morphological study of An. subpictus in Thailand and South Sulawesi, Indonesia. The maxillary palpi of most females from Thailand have the apical pale band longer than the subapical dark band, seta 7-I of pupae branched and short, and eggs with 18-25 float ridges. These characters do not agree with those described for species A, B, C and D in India. The females of An. subpictus from South Sulawesi usually have the subapical dark band of the maxillary palpus equal in length to the apical pale band. Phylogenetic analyses of sequences of the internal transcribed spacer 2 (ITS2) region of rDNA and the cytochrome c oxidase subunit I (COI) gene of mtDNA of specimens from Thailand, and South Sulawesi, and from various localities in GenBank, were conducted. ITS2 sequences of specimens from all localities in Thailand were identical, except for a small divergence in specimen from Phang Nga Province. Three distinct COI clades were detected in specimens from Chiang Mai Province in northern Thailand. However, crossing experiments between the three clades revealed no genetic incompatibility, suggesting that they were conspecific. ITS2 and COI sequences of most specimens from Thailand fell in clades other than those of An. subpictus species A and B and An. subpictus from Indonesia (East Nusa Tenggara, Java, South Sulawesi) and the Philippines. ITS2 sequences from South Sulawesi and East Nusa Tenggara were very similar, and fell in a clade consisting of specimen from Phang Nga in southern Thailand and sequences of some specimens from Cambodia and Vietnam, but their COI sequences were distinct. DNA sequences and morphological differences suggest the presence of two species within An. subpictus in Thailand, and more than one species in Indonesia.
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http://dx.doi.org/10.1016/j.actatropica.2020.105503DOI Listing
May 2020

Molecular and morphological evidence for sibling species within Anopheles (Anopheles) lindesayi Giles (Diptera: Culicidae) in Bhutan.

Acta Trop 2020 Jul 10;207:105455. Epub 2020 Apr 10.

Center of Insect Vector Study, Department of Parasitology, Faculty of Medicine, Chiang Mai University, Chiang Mai, Thailand. Electronic address:

This paper reports the results of a comparative molecular and morphological study of An. lindesayi collected from various districts of Bhutan and An. l. cameronensis from Thailand, compared with GenBank accessions and publications for An. l. japonicus from Japan, South Korea and China, An. l. pleccau from Taiwan, and An. lindesayi from India. Phylogenetic analyses based on ribosomal (ITS2) and mitochondrial (COI) DNA sequences using the Maximum Likelihood method revealed five genetically distinct clades (A, B, C, D and E) in Bhutan. Specimens in Clade A correspond to the original description of An. lindesayi, particularly in wing markings, the pattern of basal pale scales on the hindfemur and the single seta 4-C of larvae, and their COI sequences were closely related to one Indian sequence. Larvae of Clades B, C, D and E are similar in having seta 4-C branched rather than single. The adults of Clades C, D and E (B not available) are distinguishable from those of Clade A and other subspecies. Specimens of Clade C are unique in having a long pale spot on wing vein R and the subcosta, scattered pale scales on several veins and a dark spot at the tip of vein R. The adults of Clades D and E are similar in having a dark spot at the tip of vein R and no scattered pale scales on all other veins. We provisionally recognize mosquitoes of Clades A, B, C, D and E as species A, B, C, D and E, respectively, of the Lindesayi Complex. Species A is An. lindesayi sensu stricto and the others are unnamed species. Concomitantly, the previous concept of the "Lindesayi Complex", which included An. lindesayi, An. menglangensis, An. nilgiricus and An. wellingtonianus, is now recognized as the Lindesayi Subgroup of the Lindesayi Group (Anopheles Series, subgenus Anopheles) with the five sibling species of An. lindesayi comprising a more apposite Lindesayi Complex within the subgroup.
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http://dx.doi.org/10.1016/j.actatropica.2020.105455DOI Listing
July 2020

Description of Aedes (Hulecoeteomyia) bhutanensis n. sp. (Diptera: Culicidae) from Bhutan.

Acta Trop 2020 Mar 24;203:105280. Epub 2019 Dec 24.

Department of Life Sciences, Natural History Museum, Cromwell Road, London SW7 5BD, UK.

A new species of the subgenus Hulecoeteomyia of the genus Aedes reared from pupae found with two larvae in a small pool on a log in mountainous forest of Bhutan is diagnosed and formally named Aedes bhutanensis Somboon & Harbach, n. sp. The specific status of the species is supported by differential morphological characters of females, larvae and pupae, and sequences of the mitochondrial COI gene. Dichotomous keys are provided for identification of the adult females and larvae of the species of the subgenus.
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http://dx.doi.org/10.1016/j.actatropica.2019.105280DOI Listing
March 2020

Molecular identification of mosquitoes of the Anopheles maculatus group of subgenus Cellia (Diptera: Culicidae) in the Indonesian Archipelago.

Acta Trop 2019 Nov 5;199:105124. Epub 2019 Aug 5.

Center of Insect Vector Study, Department of Parasitology, Faculty of Medicine, Chiang Mai University, Chiang Mai, Thailand. Electronic address:

This study reports the molecular differentiation of females of Anopheles maculatus s.l. collected in eight localities on five islands in the Indonesian Archipelago: Hargowilis and Hargotirto villages of Central Java Province, North Kalimantan Province, Sabang off the northern tip of Sumatra Province, Sumba Island of East Nusa Tenggara Province and Sulawesi Province. Analyses based on rDNA (ITS2 and D3) and mtDNA (COII) sequences revealed the presence of An. greeni for the first time in North Kalimantan, and at least one novel (previously unrecognized) species of the Maculatus Group in Central Java (Hargowilis). Despite the similarity of rDNA markers of specimens of An. maculatus s.l. from Central Java and Sulawesi, their COII sequences are highly divergent (3.3%), which might indicate the presence of a further new species. Specimens of An. maculatus s.l. from the other localities had identical rDNA sequences to most An. maculatus s.s. from mainland Southeast Asia, but moderate divergence in their COII sequences (1.2-2.1%). The latter might indicate there are further novel species within the Maculatus Complex. However, as the divergence at COII may be the result of geographical structuring within species related to the historical biogeography of the region, further studies are needed to shed light on this possibility.
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http://dx.doi.org/10.1016/j.actatropica.2019.105124DOI Listing
November 2019

Assessment and an updated list of the mosquitoes of Saudi Arabia.

Parasit Vectors 2019 Jul 19;12(1):356. Epub 2019 Jul 19.

Department of Life Sciences, Natural History Museum, Cromwell Road, London, SW7 5BD, UK.

Background: Mosquito-borne pathogens are important causes of diseases in the Kingdom of Saudi Arabia. Knowledge of the mosquito fauna is needed for the appropriate control of the vectors that transmit the pathogens and prevent the diseases they cause. An important first step is to have an up-to-date list of the species known to be present in the country. Original occurrence records were obtained from published literature and critically scrutinized to compile a list of the mosquito species that occur within the borders of the Kingdom.

Results: Fifty-one species have been recorded in the Kingdom; however, the occurrence of two of these species is unlikely. Thus, the mosquito fauna of the Kingdom comprises 49 species that include 18 anophelines and 31 culicines. Published records are provided for each species. Problematic records based on misidentifications and inappropriate sources are discussed and annotated for clarity.

Conclusion: Integrated morphological and molecular methods of identification are needed to refine the list of species and accurately document their distributions in the Kingdom.
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http://dx.doi.org/10.1186/s13071-019-3579-4DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC6642568PMC
July 2019

Lutzia (Metalutzia) chiangmaiensis n. sp. (Diptera: Culicidae), Formal Name for the Chiang Mai (CM) Form of the Genus Lutzia in Thailand.

J Med Entomol 2019 09;56(5):1270-1274

Department of Life Sciences, Natural History Museum, London, UK.

The Chiang Mai (CM) form of the genus Lutzia, subgenus Metalutzia, in Thailand is diagnosed and formally named Lt. chiangmaiensis Somboon & Harbach, n. sp. The species is contrasted with related species, and information is provided on its bionomics and distribution.
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http://dx.doi.org/10.1093/jme/tjz072DOI Listing
September 2019

A detailed review of the mosquitoes (Diptera: Culicidae) of Iran and their medical and veterinary importance.

Acta Trop 2019 Jun 18;194:106-122. Epub 2019 Mar 18.

Department of Life Sciences, Natural History Museum, London, UK.

Mosquitoes (Diptera: Culicidae) are the most significant arthropods of medical importance because of the burden of diseases, such as malaria, encephalitis and filariasis, which are caused by pathogens and parasites they transmit to humans. In 2007, the most recently published checklist of Iranian mosquitoes included 64 species representing seven genera. Public databases were searched to the end of August 2018 for publications concerning the diseases in Iran caused by mosquito-borne pathogens. Pertinent information was extracted and analyzed, and the checklist of Iranian mosquitoes was updated. Six arboviral diseases, two bacterial diseases, four helminthic diseases and two protozoal diseases occur in Iran. The agents of these diseases are biologically or mechanically known or assumed to be transmitted by mosquitoes. The updated checklist of Iranian mosquitoes includes 69 species representing seven or 11 genera depending on the generic classification of aedines. There is no published information about the role of mosquitoes in the transmission of the causal agents of avian malaria, avian pox, bovine ephemeral fever, dengue fever, Rift Valley fever, Sindbis fever, Deraiophoronema evansi infection, lymphatic filariasis, anthrax and tularemia in Iran. There is just one imported case of lymphatic filariasis, which is not endemic in the country. It seems arthropods do not play an important role in the epidemiology of anthrax and ixodid ticks are the main vectors of the tularemia bacterium. In view of the recent finding of only a few adults and larvae of Aedes albopictus in southeastern Iran and the absence of Ae. aegypti, it is not possible to infer the indigenous transmission of the dengue fever virus in Iran. Considering the importance of mosquito-borne diseases in the country, it is necessary to improve vector and vector-borne disease surveillance in order to apply the best integrated vector management interventions as a part of the One Health concept.
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http://dx.doi.org/10.1016/j.actatropica.2019.03.019DOI Listing
June 2019

Genetic and morphological evidence for a new species of the Maculatus Group of Anopheles subgenus Cellia (Diptera: Culicidae) in Java, Indonesia.

Parasit Vectors 2019 Mar 14;12(1):107. Epub 2019 Mar 14.

Center of Insect Vector Study, Department of Parasitology, Faculty of Medicine, Chiang Mai University, Chiang Mai, Thailand.

Background: Anopheles maculatus, a species of the Maculatus Group of subgenus Cellia (Diptera: Culicidae), is an important vector of human malarial protozoa in Java, Indonesia. However, the identity of this species in Indonesia has been questionable because published reports and records are based mainly on morphological identification, which is unreliable for distinguishing members of the Maculatus Group due to overlapping characters.

Methods: We performed morphological assessments, metaphase karyotype preparations, phylogenetic analyses of ITS2 and cox2 sequence data and cross-mating experiments to determine whether the Javanese form and An. maculatus (s.s.) from Thailand were conspecific.

Results: The adults of the Java strain are similar to those of An. maculatus (s.s.), but the larvae and pupae exhibit significant differences. The metaphase karyotype of Javanese specimens includes a long acrocentric X chromosome and a small telocentric Y chromosome, which are distinct from other members of the Maculatus Group. Cross-mating of the Java strain with An. maculatus (s.s.) revealed genetic incompatibility. Phylogenetic analysis of ITS2 and cox2 sequences revealed that the Java strain forms a single clade that is distinct from clades of other members of the group (Kimura 2-parameter, K2P, genetic distances 3.1-19.2% and 1.6-9.6%, respectively).

Conclusions: This study provides evidence that the Javanese form of An. maculatus is not conspecific with An. maculatus (s.s.) and constitutes a previously unrecognized species of the Maculatus Group.
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http://dx.doi.org/10.1186/s13071-019-3358-2DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC6419379PMC
March 2019

Morphological and molecular evidence for a new species of Lutzia (Diptera: Culicidae: Culicini) from Thailand.

Acta Trop 2019 Mar 26;191:77-86. Epub 2018 Dec 26.

Center of Insect Vector Study, Department of Parasitology, Faculty of Medicine, Chiang Mai University, Chiang Mai, 50200, Thailand. Electronic address:

We found a species of LutziaTheobald (1903) (Diptera: Culicidae) in Chiang Mai Province and other provinces in northern Thailand which bears morphological and DNA sequence differences from the three species of Lutzia, subgenus Metalutzia Tanaka, previously recorded from Thailand, namely Lt. fuscana (Wiedemann), Lt. halifaxii (Theobald) and Lt. vorax Edwards. The adults of the Chiang Mai form (CM form) have abdominal banding patterns similar to those of Lt. vorax from Japan (which includes the type locality of this species), but differ in having the mediocubital crossvein usually positioned before rather than beyond the radiomedial crossvein. The thoracic and abdominal integument of Lt. vorax larvae is covered with relatively short pointed spicules whereas it is covered by denser, longer and more acutely pointed spicules in the CM form. Some differences are also found in the development of thoracic seta 1-M, and abdominal setae 8-II and 8-III. The pupa of the CM form clearly differs from the pupa of Lt. vorax in having setae 1 and 5 of abdominal segments V and VI branched rather than single. The characters of the wing, larva and pupa of the CM form are similar to those of Lt. fuscana and Lt. halifaxii. However, whereas the phallosome of males of the CM form is similar to males of Lt. vorax and Lt. halifaxii, it is distinct from males of Lt. fuscana. Phylogenetic analysis of mitochondrial cytochrome c oxidase subunits I and II sequences revealed that the CM form falls outside a clade comprised of specimens of Lt. vorax from Japan, Korea, Thailand and Bhutan (Kimura 2-parameter, K2P, genetic distances 3.9-5.6, and 5.1-6.6, respectively). However, the two gene sequences of the CM form are not clearly distinct from clades comprised of sequences from specimens of Lt. fuscana and Lt. halifaxii (K2P 0.2-2.4%). However, based on the combination of morphological and molecular data, the current study provides evidence that the CM form is a previously unrecognized species of the genus Lutzia.
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http://dx.doi.org/10.1016/j.actatropica.2018.12.029DOI Listing
March 2019

Two new species of the subgenus Sabethinus of Sabethes (Diptera: Culicidae) from Costa Rica, first confirmation of members of the subgenus in Central America.

Authors:
Ralph E Harbach

Zootaxa 2018 Jun 6;4429(2):269-280. Epub 2018 Jun 6.

Department of Life Sciences, Natural History Museum, Cromwell Road, London SW7 5BD, UK..

Two new species of Sabethes subgenus Sabethinus (Diptera: Culicidae) from Costa Rica are formally named Sa. andreae and Sa. theresae. Descriptions consisting of differential and diagnostic morphological characters are provided for adults, male genitalia and fourth-instar larvae. The male genitalia and the fourth-instar larva and pupa of both species are illustrated. The species are compared with the four previously described species known to occur only in South America, i.e. Sa. idiogenes, Sa. intermedius, Sa. melanonymphe and Sa. xhyphydes. A new country record for Colombia is reported for Sa. xhyphydes and keys are provided to distinguish the adults and larvae of the six species.
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http://dx.doi.org/10.11646/zootaxa.4429.2.3DOI Listing
June 2018

A checklist of the Anopheles mosquito species (Diptera: Culicidae) in Bhutan.

Acta Trop 2018 Dec 10;188:206-212. Epub 2018 Sep 10.

Center of Insect Vector Study, Department of Parasitology, Faculty of Medicine, Chiang Mai University, Chiang Mai, Thailand. Electronic address:

The present paper records, for the first time, the Anopheles fauna of Bhutan, determined from surveys conducted from 2007 to early 2018. Adult mosquitoes were collected mainly on cattle bait and occasionally in human landing catches. Collections of immature stages were performed in various aquatic habitats. Larvae were preserved or reared to adults. Identification was based on morphological characters using available keys. A total of 30 species were identified, including nine species of subgenus Anopheles and 21 species of subgenus Cellia. Distribution and collection data are provided with notes on the locations and habitats of the species. Anopheles pseudowillmori is suspected to be a vector of malarial parasites in the plains and hilly forested areas of the country because it is widely distributed and the most common species collected in human landing catches. Notes also include observed morphological variation observed in An. baileyi and An. lindesayi, which differ from the type forms. Corrections are made for previous reports of Anopheles in Bhutan. The need for further surveys and molecular identification of members of species complexes and morphological variants is emphasized.
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http://dx.doi.org/10.1016/j.actatropica.2018.09.006DOI Listing
December 2018

An Anopheles by Any Other Name …?

Authors:
Ralph E Harbach

J Med Entomol 2018 08;55(5):1069-1070

Department of Life Sciences, Natural History Museum, London, UK.

The physical characteristics of a nematoceran fly that evokes the image of a mosquito of the genus Anopheles Meigen (Diptera: Culicidae) has existed since the genus was defined in the early part of the twentieth century. Is that image likely to change with the recent proposal to elevate four Neotropical subgenera to generic status based on relationships generated by phylogenetic analyses of mitochondrial protein-coding genes? Reasons for retaining the traditional concept of Anopheles, with its traditionally recognized subgenera, are presented. However, as the ranking of taxa as genera or subgenera is subjective, the choice of rank of a genus-group name may be considered a subjective decision and a matter of user preference.
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http://dx.doi.org/10.1093/jme/tjy108DOI Listing
August 2018

First Records of and in Quintana Roo State, Mexico.

J Am Mosq Control Assoc 2018 06;34(2):124-127

Collections of mosquitoes were conducted for the surveillance of species of medical importance in the state of Quintana Roo, Mexico, in June of 2017. Species collected included () and () , both new records for the state. was previously recorded in Mexico State and in Chiapas State. With the addition of these species, the mosquito fauna of Quintana Roo State now includes 81 species. Quintana Roo and Tamaulipas State with 82 species are the most mosquito-rich states in Mexico, based on currently available records. The specimens collected in this study were deposited in the Culicidae collection of the Universidad Autónoma Agraria Antonio Narro Unidad Laguna.
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http://dx.doi.org/10.2987/18-6743.1DOI Listing
June 2018

Controversies over the scientific name of the principal mosquito vector of yellow fever virus - expediency versus validity.

J Vector Ecol 2018 06;43(1):1-14

Department of Life Sciences, Natural History Museum, London, UK.

The history of the scientific name of the yellow fever mosquito, the vector of yellow fever virus, ranges from 1757 to the early twenty-first century. In his 1757 work Iter Palaestinum, Frederic Hasselquist gave the name Culex aegypti to a mosquito species responsible for fierce attacks on humans in Egypt. That name was never later ascribed to Hasselquist as author, but to Linnaeus, although the name never appeared in any of Linnaeus' publications. In Cuba, at the end of the nineteenth century, the vector of the unknown infectious agent of yellow fever was first identified as Culex mosquito and later more validly named Stegomyia fasciata. Mosquito taxonomists differed strongly about the name of the mosquito through much of the twentieth century. Interventions by the International Commission on Zoological Nomenclature imposed a biologically invalid specific name, and in the early twenty-first century a phylogenetic analysis of the culicid tribe Aedini restored the genus Stegomyia from a century earlier. That action was short-lived. A phylogenetic reassessment resulted in the return of Stegomyia to subgeneric rank in Aedes; thus, the name of the yellow fever mosquito survives in the traditional classification of convenience as the trinomial Aedes (Stegomyia) aegypti (Linnaeus).
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http://dx.doi.org/10.1111/jvec.12277DOI Listing
June 2018

History of the discovery of the mode of transmission of yellow fever virus.

J Vector Ecol 2017 12;42(2):208-222

Department of Life Sciences, Natural History Museum, London, UK.

This essay documents and examines the historical circumstances and events surrounding the discovery of the mode of transmission of yellow fever virus in Cuba. Close scrutiny of the articles published by Walter Reed and his colleagues in 1900, 1901 and 1902 reveals their limitations as historic documents. Fortunately, other sources of information from that period survive in letters and documents written by individuals involved in the quest for the mode of transmission. Examination and comparison of those sources of information unveiled a fascinating story which reveals that misunderstandings engendered by published articles accorded merit where it was not fully due.
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http://dx.doi.org/10.1111/jvec.12261DOI Listing
December 2017

Natural Plasmodium vivax infections in Anopheles mosquitoes in a malaria endemic area of northeastern Thailand.

Parasitol Res 2017 Dec 29;116(12):3349-3359. Epub 2017 Oct 29.

Department of Parasitology, Chiang Mai University, Chiang Mai, 50200, Thailand.

There was recently an outbreak of malaria in Ubon Ratchathani Province, northeastern Thailand. In the absence of information on malaria vector transmission dynamics, this study aimed to identify the anopheline vectors and their role in malaria transmission. Adult female Anopheles mosquitoes were collected monthly by human-landing catch in Na Chaluai District of Ubon Ratchathani Province during January 2014-December 2015. Field-captured mosquitoes were identified to species using morphology-based keys and molecular assays (allele-specific polymerase chain reaction, AS-PCR), and analysed for the presence of Plasmodium falciparum and Plasmodium vivax using an enzyme-linked immunosorbent assay (ELISA) for the detection of circumsporozoite proteins (CSP). A total of 1,229 Anopheles females belonging to 13 species were collected. Four anopheline taxa were most abundant: Members of the Anopheles barbirostris complex, comprising 38% of the specimens, species of the Anopheles hyrcanus group (18%), Anopheles nivipes (17%) and Anopheles philippinensis (12%). The other nine species comprised 15% of the collections. Plasmodium infections were detected in two of 668 pooled samples of heads/thoraces, Anopheles dirus (1/29) and An. philippinensis (1/97). The An. dirus pool had a mixed infection of P. vivax-210 and P. vivax-247, whereas the An. philippinensis pool was positive only for the latter protein variant. Both positive ELISA samples were confirmed by nested PCR. This study is the first to incriminate An. dirus and An. philippinensis as natural malaria vectors in the area where the outbreak occurred. This information can assist in designing and implementing a more effective malaria control programme in the province.
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http://dx.doi.org/10.1007/s00436-017-5653-1DOI Listing
December 2017

(Eckstein, 1918) (Diptera, Culicidae), a new country record for England, contrasted with (Meigen, 1838).

Zookeys 2017 27(671):119-130. Epub 2017 Apr 27.

Medical Entomology & Zoonoses Ecology group, Emergency Response Department, Public Health England, Porton Down, Salisbury, SP4 0JG, UK.

We report the discovery of (Eckstein, 1918) in the New Forest of southern England, bringing to 36 the number of mosquito species recorded in Britain. Because it seems that this species has been misidentified previously in Britain as the morphologically similar (Meigen, 1838), the two species are contrasted and distinguished based on distinctive differences exhibited in the adult and larval stages. The pupa of is unknown, but the pupa of is distinguished from the pupae of other species of by modification of the most recent key to British mosquitoes. The history of the mosquito fauna recorded in the UK is summarized and bionomical information is provided for the two species.
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http://dx.doi.org/10.3897/zookeys.671.12447DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC5523212PMC
April 2017

Discovery of a single male Aedes aegypti (L.) in Merseyside, England.

Parasit Vectors 2017 Jun 24;10(1):309. Epub 2017 Jun 24.

Department of Biology, Edge Hill University, St. Helens Road, Ormskirk, Lancashire, L39 4QP, UK.

Background: The mosquito Aedes aegypti (L.) is found in tropical and sub-tropical regions where it is the major vector of dengue fever, yellow fever, chikungunya and more recently Zika virus. Given its importance as a vector of arboviruses and its propensity to be transported to new regions, the European Centre for Disease Prevention and Control (ECDC) has placed Ae. aegypti on a list of potentially invasive mosquito species. It was previously reported in the United Kingdom (UK) in 1865 and 1919 but did not establish on either occasion. It is now beginning to reappear in European countries and has been recorded in the Netherlands (not established) and Madeira (Portugal), as well as southern Russia, Georgia and Turkey.

Results: During summer 2014, a single male Ae. aegypti was captured during mosquito collections in north-western England using a sweep net. Morphological identification complimented by sequencing of the ITS2 rDNA, and cox1 mtDNA regions, confirmed the species. Following confirmation, a programme of targeted surveillance was implemented around the collection site by first identifying potential larval habitats in greenhouses, a cemetery, a farm and industrial units. Despite intensive surveillance around the location, no other Ae. aegypti specimens were collected using a combination of sweep netting, larval dipping, mosquito magnets, BG sentinel traps and ovitraps. All species collected were native to the UK.

Conclusion: The finding of the single male Ae. aegypti, while significant, presents no apparent disease risk to public health, and the follow-up survey suggests that there was no established population. However, this report does highlight the need for vigilance and robust surveillance, and the requirement for procedures to be in place to investigate such findings.
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http://dx.doi.org/10.1186/s13071-017-2251-0DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC5483247PMC
June 2017

An updated checklist of the Culicidae (Diptera) of Morocco, with notes on species of historical and current medical importance.

J Vector Ecol 2017 06;42(1):94-104

Department of Life Sciences, Natural History Museum, London, UK.

An updated checklist of the mosquito species (Diptera: Culicidae) recorded in Morocco from 1916 to 2016 is provided, including synonyms and synonymous usage for each species. Forty-three species belonging to seven genera are recorded so far: Anopheles (9), Aedes (12) Coquillettidia (2), Culex (12), Culiseta (5), Orthopodomyia (1) and Uranotaenia (2). Traditional and equivalent names in the polyphyletic concept of Aedes are provided for the aedine species. The historical importance and current potential threat of mosquitoes to human health in Morocco is reviewed.
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http://dx.doi.org/10.1111/jvec.12243DOI Listing
June 2017

Culex (Culiciomyia) sasai (Diptera: Culicidae), senior synonym of Cx. spiculothorax and a new country record for Bhutan.

Acta Trop 2017 Jul 13;171:194-198. Epub 2017 Apr 13.

Department of Parasitology, Faculty of Medicine, Chiang Mai University, Chiang Mai 50200, Thailand. Electronic address:

Culex (Culiciomyia) spiculothorax was described from Thailand based on the presence of spiculation on the thorax of larvae. Adult females are characterized but are indistinguishable from those of related species, such as Cx. pallidothorax. Phylogenetic analysis of mitochondrial oxidase subunit I (COI) sequences revealed that specimens identified as Cx. spiculothorax from Thailand, Japan and Bhutan form a single clade with Cx. sasai from Japan (Kimura 2-parameter genetic distances 0-0.9%) that is clearly distinct from clades comprised of other species of subgenus Culiciomyia. Attempts to collect Cx. sasai from several locations in Japan were unsuccessful - only larvae with thoracic vesicular-like spicules identified as Cx. spiculothorax were collected. Careful examination of specimens collected near the type locality of Cx. sasai revealed the presence of spicules on the thorax. Based on these findings, Cx. spiculothorax is formally synonymized with Cx. sasai, which replaces the former as the species present in Thailand and is a new country record for Bhutan.
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http://dx.doi.org/10.1016/j.actatropica.2017.04.003DOI Listing
July 2017

Anopheles prachongae, a new species of the Gigas Complex of subgenus Anopheles (Diptera: Culicidae) in Thailand, contrasted with known forms of the complex.

Zootaxa 2017 Feb 21;4236(1):zootaxa.4236.1.3. Epub 2017 Feb 21.

Department of Life Sciences, Natural History Museum, Cromwell Road, London SW7 5BD, UK..

Anopheles (Anopheles) prachongae, a new species of the Gigas Complex from northern Thailand, is described and illustrated in the adult, pupal and larval stages, and bionomics and chaetotaxy tables are provided for the immature stages. The species is distinguished from Anopheles baileyi, the only other species of the complex known to occur in Thailand, and contrasted with other taxa of the complex that occur in the Oriental Region. Available morphological data indicate that An. gigas sumatrana is unique and is therefore formally afforded species status. The three other Sumatran subspecies may be conspecific. The taxonomic status of the non-Sumatran subspecies, i.e. crockeri (Borneo), formosus (Philippines), refutans (Sri Lanka) and simlensis (south-central Asia), is questioned but their status is unchanged pending further study.
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http://dx.doi.org/10.11646/zootaxa.4236.1.3DOI Listing
February 2017

An effective method for the identification and separation of Anopheles minimus, the primary malaria vector in Thailand, and its sister species Anopheles harrisoni, with a comparison of their mating behaviors.

Parasit Vectors 2017 02 21;10(1):97. Epub 2017 Feb 21.

Department of Parasitology, Faculty of Medicine, Chiang Mai University, Chiang Mai, 50200, Thailand.

Background: Species of the Anopheles minimus complex are considered to be the primary vectors of malaria in South and Southeast Asia. Two species of the complex, Anopheles minimus and Anopheles harrisoni, occur in Thailand. They are sympatric and difficult to accurately distinguish based on morphological characters. The aim of this study was to investigate the potential of antennal sensory organs to distinguish these two species. Additionally, we investigated their ability to mate in cages of different sizes, as well as the possible mechanism(s) that evokes stenogamous behavior.

Methods: Large sensilla coeloconica present on the antennae of females of An. minimus and An. harrisoni were counted under a conventional light microscope and various types of antennal sensilla were examined under a scanning electron microscope (SEM). Determinations of mating ability were carried out in 20 and 30 cm cages with a density resting surface (DRS) of 7.2. The insemination rate, frequency of clasper (gonocoxopodite) movement of the male genitalia during induced copulation and duration of mating of the two species were compared.

Results: The mean numbers of large sensilla coeloconica on antennal flagellomeres 1-8 and the mean number of large sensilla coeloconica on each flagellum in An. minimus (26.25) and An. harrisoni (31.98) were significantly different. Females of both species bear five types of antennal sensilla: chaetica, trichodea, basiconica, coeloconica and ampullacea. Marked differences in the structure of the large sensilla coeloconica were observed between the two species. Furthermore, only An. minimus could copulate naturally in the small cages. The frequency of clasper movement in the stenogamous An. minimus was significantly higher than in An. harrisoni, but there was no difference in the duration of mating.

Conclusions: To our knowledge, this study is the first to examine and discover the usefulness of large sensilla coeloconica on the antennae of females and the frequency of clasper movement in males for distinguishing the sibling species An. minimus and An. harrisoni. The discovery provides an effective and relatively inexpensive method for their identification. Additionally, the greater frequency of clasper movement of An. minimus might influence its ability to mate in small spaces.
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http://dx.doi.org/10.1186/s13071-017-2035-6DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC5320799PMC
February 2017

A review of the mosquito species (Diptera: Culicidae) of Bangladesh.

Parasit Vectors 2016 10 22;9(1):559. Epub 2016 Oct 22.

Department of Life Sciences, Natural History Museum, Cromwell Road, London, SW7 5BD, UK.

Background: Diseases caused by mosquito-borne pathogens remain an important source of morbidity and mortality in Bangladesh. To better control the vectors that transmit the agents of disease, and hence the diseases they cause, and to appreciate the diversity of the family Culicidae, it is important to have an up-to-date list of the species present in the country. Original records were collected from a literature review to compile a list of the species recorded in Bangladesh.

Results: Records for 123 species were collected, although some species had only a single record. This is an increase of ten species over the most recent complete list, compiled nearly 30 years ago. Collection records of three additional species are included here: Anopheles pseudowillmori, Armigeres malayi and Mimomyia luzonensis.

Conclusions: While this work constitutes the most complete list of mosquito species collected in Bangladesh, further work is needed to refine this list and understand the distributions of those species within the country. Improved morphological and molecular methods of identification will allow the refinement of this list in years to come.
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http://dx.doi.org/10.1186/s13071-016-1848-zDOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC5075421PMC
October 2016
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