QUR'AN AS A COMPENDIOUS {THE PROLOGUE OF THE BASIC CONCEPT}

Allah (the exalted and the glorious) vehemently engraved everything in his noble book known as Qur'an.
The Qur'an comprises of everything of knowledge and wisdom that is beyond one's fictitious. According
to Dr. Dogãn in his book of Usulul tafsir he qouted as follows:

The Qur’ān is the basis for alI islamic sciences and disciplines, without it they would not exist, in the
early days of Islam religious commandments were memorized and transmitted by oral tradition rather
than transcribed. Religious rules and regulations in relation to creed, worship, and daily life were stored
within the memories of the early Muslims. Since religious commandments were vital issues in a
Muslim’s personal and social life, scholars gave these issues priority and compiled books on religious
commandments using the Qur’ān as guidance. Using the Qur’ān as their main source, Islamic jurists
collected and codified books on Islamic law and its rules and principles pertaining to all fields of life,
Hadith scholars (muhaddithun) established the Prophetic traditions and preserved them in collections,
theologians dealt with issues concerning Muslim belief, exegetes of the Qur’ān dedicated themselves to
studying its meaning and interpretation including issues that would later be called ‘Qur’anic sciences’,
linguists used the Qur’ān as a base for their philological arguments, and qurrā (professionals in Qur’ān
recitation) used it for their studies.In the third century of Islam, most Islamic disciplines established their
methodology (usūl). Usūl al-Tafsīr or ‘Ulūm al-Qur’ān was one such discipline established in this period.
Previously, the science of the Qur’ān came under the Hadith science and was recognized as the ‘book of
exegesis’ (kitāb al-tafsīr). The two were later separated and each science established its own
methodology. The sciences of the Qur’ān focused on the knowledge of those sciences that had a direct
relationship with the recitation, history, understanding and implementation of the Qur’ān. In order to
interpret the Qur’ān, it is essential to have knowledge of many subjects including, and not limited to, a
thorough knowledge of Arabic grammar, Arabic rhetoric, and principles of jurisprudence (fiqh), the
various readings of the Qur’ān, the sciences of the exegesis, the Prophetic traditions and many more.
‘Ulūm al-Qur’ān is understood by early scholars as the sciences and disciplines contained within the
Qur’ān.

____________________________

Zarkashi, Al-Burhan fil ulumul Qur'an, 1/17.

The Qur’ān is an inexhaustible source of all kinds of knowledge as well as being the subject of study for
exegetical works. The sciences, which are derived from the Qur’ān or applied to it, were divided by
scholars into many disciplines and collectively they created the Qur’anic sciences.

Allah SWT said:

‫َو َي ْو َم َن ْب َع ُث ِفى ُك ِّل ُأ َّم ٍة َش ِهي ًدا َع َل ْي ِه ْم ِّم ْن َأ ْن ُف ِس ِه ْم ۖ  َو ِج ْئ َنا ِب َك َش ِهي ًدا َع ٰلى ٰٓه ُؤَل ٓا ِء ۚ  َو َن َّزْل َنا َع َل ْي َك ا ْل ِك ٰت َب ِت ْب ٰي ًنا ِّل ُك ِّل َش ْى ٍء َو ُه ًدى َو َر ْح َم ًة َو ُب ْش ٰرى ِل ْل ُم ْس ِل ِمي َن‬

"And [mention] the Day when We will resurrect among every nation a witness over them from
themselves. And We will bring you, [O Muhammad], as a witness over your nation. And We have sent
down to you the Book as clarification for all things and as guidance and mercy and good tidings for the
Muslims." An Nahl 89:

Allah SWT said:

‫َو ِع ْن َد ُهۥ َم َفا ِت ُح ا ْل َغ ْي ِب َلا َي ْع َل ُم َه ٓا ِإ َّلا ُه َو ۚ  َو َي ْع َل ُم َما ِفى ا ْل َب ِّر َوا ْل َب ْح ِر ۚ  َو َما َت ْس ُق ُط ِم ْن َو َر َق ٍة ِإ َّلا َي ْع َل ُم َها َوَلا َح َّب ٍة ِفى ُظ ُل ٰم ِت ا ْل َأ ْر ِض َوَلا َر ْط ٍب َوَلا َيا ِب ٍس ِإ َّلا ِفى ِك ٰت ٍب ُّم ِبي ٍن‬

"And with Him are the keys of the unseen; none knows them except Him. And He knows what is on the
land and in the sea. Not a leaf falls but that He knows it. And no grain is there within the darknesses of
the earth and no moist or dry [thing] but that it is [written] in a clear record."

An'aam 6: Verse 59)

Commentary

"And with Him are the keys of the Unseen. None but He knows them. ..."

Then, for a more emphasis and explanation, it adds:

"... And He knows whatever is in the land and the sea....."

The Arabic term /barr/ means 'a vast place', which is usually used for a (dry) land. The term /bahr/
originally also means 'a vast place' where a plenty of water is found. It is often applied for the seas and,
sometimes, for the great streams.

However, Allah’s knowledge upon whatever exists in the lands and the seas is something which hints to
Allah’s omniscience, and that His knowledge encompasses all things.

That is, He knows the movements of millions of million living creatures, small and big, in the depths of
the seas. He knows the true number of the cells of every human being as well as his blood cells.

He knows the mystic circulations of all electrons inside atoms. And, finally, He knows all thoughts and
contemplations pass through different parts and levels of our minds and those which penetrate into the
depths of our soul.

Yes, He knows equally all of these things! Again, in the second phrase, in order to emphasize on Allah’s
Omniscience, it has particularly pointed to this aspect, and says:

"and no leaf (of a tree) drops down but He knows it, nor a grain in the darkness of the earth ..."

In fact, it has taken two sensitive subjects with which no human being can be thoroughly acquainted
even if he lives millions of years and that the technical sets be surprisingly completed.

Who knows that, in each day, which seeds the winds separate from the plants throughout the earth, and
on which spots of the ground they scatter them? Which electronic mind can count exactly the number of
the leaves that separate from the branches of trees of the forests only during a day?
A glance upon the trees of a forest, especially in fall, when the leaves drop ceaselessly and make up
beautiful scenery, proves this fact well that such sciences will never be within the reach of human
beings.

The fall of the leaves of trees, in fact, is the moment of their death, and the fall of seeds into the
concealed holes of the land is the first step of their lives. It is only He Who is aware of the system of
death and life.

The statement of this subject has two effects: a philosophical effect and a training effect. Its
philosophical effect is that it nullifies the imagination of those who restrict Allah’s knowledge to general
principles and believe that He is not aware of the details of this world. It clearly specifies that Allah is
aware of both all general principles and the details.

Its training effect is also clear, because having belief in the vast knowledge of Allah, that He is
Omniscience, tells us that the whole secrets of our entity, our deeds and speeches, our intentions and
our thoughts are utterly clear for His Pure Essence.

With such a belief, how is it possible that a person be careless of his own condition and does not control
his own deeds, speeches, and intentions?

So, at the end of the verse, it says:

"Nor anything wet or dry but it is (noted) in a clear Book.". And this verse is enough as an inference for
those who reasoned to imagine the knowledge engraved in the holy Qur'an base on their capacity of
imagination.And also the Qur’anic sciences primarily cover; the occasions of revelation (asbāb an-nuzūl),
the verses which were revealed in Mecca or Medina, the various forms of Qur’ān reading (ahruf), the
abrogating and abrogated rulings and verses (nāsikh-mansūkh), the various classifications of verses
(muhkam, mutashābih, mutlaq, muqayyad, etc.), the inimitable style of the Qur’ān (i’jāz), the
interpretation (tafsīr), the grammatical analysis of the verses and the knowledge of words that have
fallen out of usage in modern times (gharīb al-Qur’ān). A detailed knowledge of all Qur’anic sciences is
essential in order to properly interpret the Qur’ān. Therefore, to name this subject the ‘Methodology of
Interpretation’ (Usūl al-Tafsīr) instead of ‘the Qur’anic sciences’ would be more befitting. Although the
terms ‘Usūl al-Tafsīr’ (the interpretation of the Qur’ān) and ‘Ulūm al-Qur’ān (the sciences and disciplines
of the Qur’ān) are often used interchangeably, in the case of themethodology of interpretation there are
some topics which are omitted or scarcely touched upon, such as the history and development of the
oral and written Qur’anic scripture. As such, it is more correct to regard ‘Usūl al-Tafsīr as a subset of
‘Ulūm al-Qur’ān. Not all of the sciences related to the Qur’ān play a role in explaining its meanings, for
example, the counting of the verses. The different classical authorities of Qur’anic recitation have
different methods of counting the verses of each surah. The differences in these methods have no
impact on the tafsīr of the Qur’ān even though they fall under ‘Ulūm al-Qur’ān. According to Abū Bakr b.
‘Arābī (468 – 543 A.H. / 1076 – 1148 C.E.), Qur’anic sciences encompass three areas; 1) Tawḫīd (unity
and knowledge of God, His names, attributes, essential qualities, etc. within the scope of Kalām), 2)
Tadhkīr (knowledge of ẕāhir (outer) and bātin (inner) meanings together with the afterlife (wā’d ‘giving
good tidings’ and wa’īd ‘warnings’), paradise and hell, etc. these are the topics of Tasawwuf, 3) Aḫkām
(knowledge of ḫalal (lawful) and ḫaram (unlawful) which is the topic of fiqh).

______________________________

Zarkashī, Burhān, I.38

According to imam Tabarī (224 – 310 A.H. /838–923 C.E.), he categorizes these sciences differently as
tawḫīd, akhbār (reports, history) and diyānah (religiosity).

______________________________

Zarkashī, ibid. I.40-1

Early scholars understood that the importance of the science is related with the importance of its
content, therefore, understanding the Qur’ān is the most important study among all other Islamic
sciences as the Qur’ān is prevailing and preeminent.

THE HISTORY OF QUR'ANIC SCIENCE {ULUMUL~QUR'AN}

There are many different words that have been used to express the Qur’anic sciences such as ‘fahm al-
Qur’ān’ (comprehending the Qur’ān), ’ilm al-Qur’ān’ (knowledge of the Qur’ān) and ‘fiqh al-Qur’ān’
(understanding the Qur’ān). ‘Ilm al-Qur’ān (the science of the Qur’ān) was used for the first time in
history by ‘Umar al-Wāqidī (130-207 A.H./748-822 C.E.) in his book ‘al-Rāghib Fi ‘Ilm al-
Qur’ān wa Ghalat ar-Rijāl’. Unfortunately, we don’t have this book today so it is impossible to
know how he discussed the notion of the Qur’anic sciences. From the early period to the present
day, the works on the understanding of the Qur’ān (Usūl al-Tafsīr) as one subset of the sciences of the
Qur’ān (‘Ulūm al-Qur’ān). The emergence of the technical ‘sciences of the Qur’ān’ has been credited
to the twelfth, thirteenth, and even to the beginning of the eleventh centuries. ‘Ilm al-Tafsīr
provides the infrastructure for the mufassir (interpreter/exegete) in preparation for interpreting
the Qur’ān. Usūlal-Tafsīr primarily affords mufassir with a prescriptive step-by-step guide to interpreting
the Qur’ān.‘Ulūm al-Qur’ān is more comprehensive and contains both the ‘ilm and usūl of tafsīr.
Therefore many exegeses are also called ‘Ulūm al Qur’ān.Over time, ‘Usūl al-Tafsīr (the methodology of
interpretation) gained prominence and ‘Ulūm al-Qur’ān lost its primacy. This is an example of one
such shift in Qur’anic studies. Some scholars consider disciplines which serve
purpose in the interpretation of the Qur’ān, such as Hadith, ‘Usūl al-Fiqh (the methodology
of) and others, as part of ‘Ulūm al-Qur’ān. Other scholars expand the content covered by ‘Ulūm al-
Qur’ān by including myriad disciplines within its scope. According to these scholars, such as the notion
of revelation, the collection of the Qur’ān, the occasions of revelation and verses are directly related
to the ‘Ulūm al-Qur’ān, but other disciplines such as jurisprudence and theology are indirectly
within its scope. Additionally, social and scientific discipline such as psychology, sociology, politics,
biology, physics and medicine that are dealt with and illuminated in the Qur’ān are also to be
considered as one dimension of the Qur’anic sciences.
The historic development of ‘Ulūm al Qur’ān conjures the question of why was it developed so late. The
reason for this can be extrapolated by examining Prophetic history. Prophet Muhammad (may Allah
bestows peace, mercy and blessings upon him and his household members) warned his community
against using their own opinion when interpreting the Qur’ān. Therefore, there were only a few
Companions who were deemed qualified to undertake its interpretation. The other reason is that those
early works were not under the title of ‘Ulūm al-Qur’ān. Nevertheless it is true to say that the sciences
of the Qur’ān were, in fact, birthed during the time of the Prophet (may Allah bestows peace, mercy and
blessings upon him and his household members) who explained the meaning of the verses and gave
synonyms or explained metaphors in the verses. An example of this is when the following verse was
revealed:They who believe and do not mix their belief with injustice, those will have security, and they
are rightly guided.

________________________

Qur’ān, 6: 82The Companions asked: ‘O Messenger of God! Who amongst us does not do injustice?’
Their understanding was that the verse referred to those believers who did not commit any injustice, or
sin. The Prophet (may Allah bestows peace, mercy and blessings upon him and his household members)
replied: ‘The injustice referred to in this verse is associating partners with God’.

________________________

Bukhari, Saḫīḫ, Kitāb at-Tafsīr, 153.The Companions were so enthusiastic in seeking this knowledge that
they were able not only to explain the meaning of any verse in the Qur’ān, but also give its history and
the cause of its revelation. Ibn Mas’ud reports:I swear by Allah, besides whom there is no other god,
there is no chapter in the Qur’ān that I don’t know where it was revealed. And there is not a single verse
in the Qur’ān that I don’t know the reason behind its revelation. If there were any person who knew
more about the Qur’ān than I know, and it was possible for me to reach him, I would ride on my camel
to him to gain this knowledge.

______________________

Buhkari, Saḫīḫ, Book of the Virtues of the Qur’ān, 4716.The other reason pertaining to the late
development of the Qur’anic sciences is that, for a long time, ta’wil and tafsīr were used interchangeably
to explain the Qur’ān. There are many books of different names, attributable to early generations, that
dealt with the sciences of the Qur’ān such as, I’rāb al-Qur’ān (linguistic interpretation), Ma’āni al-Qur’ān
(the meanings of the words used in the verses), Wujūh wa al-Nazāir (the different meanings of one word
and the different words which have the same meaning), Gharīb al-Qur’ān (strange or unusual words),
Majāz al-Qur’ān (metaphoric expressions), al-Nāsikh wa al-Mansūkh (abrogating and abrogated verses),
Rasm al-Muṣḫāf (the writing of the Qur’ān), etc. There are early books pertaining to the sciences of
Qur’ān such as, Qatāda (118 A.H. / 726 C.E.), Mansūkh, Zuhrî (124 A.H. / 742 C.E.), an-Nāsikh wal-
Mansūkh, Muslim b. Maysara al-Khurasanī (135 A.H. / 757 C.E.), al-Nāsikh wal-Mansūkh, Ata b. Abī
Rabah (114 A.H. / 732 C.E.), Qarīb Al~Qur'an’ān, Muqātil b. Sulayman (150 A.H. / 763 C.E.), al-Ashbah
wal-Nazāir and Mutashābih al-Qur’ān, Musa al-Azdî al-A’war (170 A.H. / 786 C.E.) and Wujūh wal-Nazāir.
It must also be mentioned that, in addition to these books, many Hadith collections, such as the Saḫīḫ
Bukhari and Muslim collections, included sections on various topics of the sciences of the Qur’ān.

to this approach, some scholars were of the opinion that the sciences of the Qur’ān developed very
late, even after the development of the methodology of fiqh and Hadith. One example is Abū al-Faraj
Ibn al-Jawzī who held that the early scholars neglected to produce a work on the ‘Ulūm al-Qur’ān similar
to the methodology of Hadith. To address this Abū alFaraj Ibn al-Jawzī (508 – 597 A.H.) wrote Funūn al-
Afnān Fi Uyūn ‘Ulūm al-Qur’ān, a comprehensive work on the sciences of the Qur’ān.

_______________________

Abu al-Faraj Abd al-Rahman Ibn al-Jawzī, Funūn al-Afnān fi Uyūn ‘Ulūm al-Qur’an, (ed.) Hasan Ziya al-Din
Itr, Beirut: Dar al-Basair al-Islamiyya 1987, 141

Zarkashī (1344/45 – 1392 C.E.) was also of the same opinion. He believed that Hadith, fiqh and other
disciplines had established at least the technical terms of their methodology whilst this was lacking in
the case of the sciences of the Qur’ān. His book al-Burhān Fi ‘Ulūm al-Qur’ān was an attempt to fill this
gap. According to Zarqānī (1367 A.H.), no work was published on ‘Ulūm al-Qur’ān before the fifth
century of Islam.

_____________________________

Zarqānī/Zurgani, Manāhil al-Irfan fi Ulūm al-Quran, I.27-8

His explanation for this was that there was no need to publish books on this subject as the sciences of
the Qur’ān were known to scholars. The first book in this field al-‘Aql wa Fahm al-Qur’ān (the intellect
and understanding the Qur’an), belongs to al-Hārith al-Muhāsibī (d.243 A.H. / 857 C.E.). In his book, al-
Muhāsibī explains the sciences of the Qur’ān through the topics of; naskh (the abrogation of the verses),
muhkam (clear verses), mutashābih (ambiguous verses), fadāil al-Qur’ān (the virtue of the Qur’ān), khalq
al-Qur’ān (the created or uncreated nature of the Qur’ān), uslūb al-Qur’ān (the style of the Qur’ān), etc.
Major chapters in his work include the reason and the understanding of the Qur’ān, the rejection of
innovators and heretics and the subjects related to the sciences of the Qur’ān.

_________________________

See for detailed explanation Adil Öksüz, Tefsir Usûlü Açısından Hâris el-Muhâsibî’nin Fehmu’l-Kur’an’ı,
SAÜ Sosyal Bilimler Enstütüsü (unpublished masters theses) 1996.

The other works of this period are Abū al- Ḫasan al-Ash’arī’s (d. 324 A.H.) alMukhtazan fî ‘Ulūm al-
Qur’ān (apparently a voluminous work, which does not exist today and is infrequently referenced), Abū
al-Mutarrif b. Isa Ashbagh (d. 402 A.H.) and Wāḫidī’s (d.468 A.H.) Asbāb an-Nuzūl, Hibat Allah b. Salama
(d. 410 A.H.) and Abū Tālib al-Makkī’s (d.386 A.H.) Nāsikh wal-Mansūkh, Hasan b. Furak (d. 410 A.H.) and
Qādi Abd al-Jabbā Al~Hamadânî’s (d. 415 A.H.) Muhkam and Mutashābih, Ali b. Ibrahim b. Sa’id al-Hūfī
or Hawfi’s (d. 430 A.H.) al-Burhān fi ‘Ulūm al-Qur’ān. Ibn al-Anbari’s (d. 328 A.H.) Ajāib ‘Ulūm al-Qur’ān
and Muhammad b. Halaf al-Marzaban’s (d. 309 A.H.) ‘al-Hāwi fi ‘Ulūm Al~Qur'an’ān’.The seventh
century of Islam witnessed major contributions to the sciences of the Qur’ān. Many books containing
comprehensive analysis of every branch of ‘Ulūm al-Qur’ān were published in this period. For example,
Rāghib al-Isfahānī (d. 503 A.H.) wrote Muqaddimāt alTafsīr, (also known as Muqaddimāt Jami al-Tafāsir)
of which there are various publications. Ibn al-Jawzī (d. 597 A.H.) wrote several books on the subject
including, Ajāib ‘Ulūm Al~Qur'an’ān and al-Mujtaba fi ‘Ulūm al-Qur’ān. Other contributing authors
include, Najm al-Din Sulayman b. Abd al-Qawi al-Tufi’s (716 A.H. / 1216 C.E.) book al-Ikthīr fi Ilm al-Tafsīr
and Ibn Taymiyya’s (728 A.H. / 1327 C.E.) book titled Muqaddima fi Usūl al-Tafsīr.It can be said that al-
Zarkashī’s Burhān and al-Suyūtī’s Itqān represent the culmination of centuries of Islamic studies on the
Qur’ān. Even today, they (especially the Itqān) remain the main sources for those writing ‘new’
handbooks on the sciences of the Qur’ān. The sciences of the Qur’ān has some more modern works
including; Tahir al-Jazâirî’s (1388 A.H. / 1919 C.E.) al-Tibyân li Ba’d al-Mabâhith al-Mutaallaqati bil-
Qur’ān, Muhammad Abd al-Azîm al-Zarqānī’s (1367 A.H. / 1948 C.E.) Manahil al-Irfan fi ‘ulūm al-Qur’ān,
Subh al-Salih’s Mabahith fi ‘ulūm al-Qur’ān, Muhammad b. Muhammad Abū Shahba’s alMadkhal ila
dirasa al-Qur’ān al-Karim, Manna’ Khalil al-Qattan’s Mabâhith fî ‘ulūm Al~Qur'an’ān, Abdallah Mahmud
Shahhâta’s Ulūm al-Qur’ān, Muhammad Adnan Zarzur’s Ulūm al-Qur’ān, Muhammad Ali al-Sabūnî’s al-
Tibyan fi ‘ulūm al-Qur’ān and Ismail Cerrahoglu’s Tefsir Usūlu.

BENEFITS OF STUDYING THE SCIENCES OF QUR'AN

The study of the Qur’anic sciences is associated with innumerable benefits including an appreciation of
its richness and wealth of knowledge, and the enabling of a more comprehensive understanding of the
book itself. Through the study of various fields within the ‘Ulūm al-Qur’ān such as the history of
revelation and collection, one is able to understand the terms used in the interpretation of the Qur’ān
and benefit from this knowledge. This, for the believer, results in an increased and strengthened faith
that can withstand any deception and false claims made by biased people. The affirmations of the
Qur’ān’s authenticity render the believer’s heart tranquil. The Qur’anic challenge to both the human
race and jinn to produce just one chapter similar to one of its chapters (a challenge made by God in
response to claims that the book was written by man) and throughout, history failed to produce any text
similar to even the shortest chapter within the Qur’ān, thus establishing its authenticity. History testifies
to this reality and the Qur’ān both predicts and affirms that no one will ever be able to present such a
chapter.The Qur’anic sciences enable its students to defend the Qur’ān against its extreme critics and
their false claims. Students achieve a profound knowledge of the Qur’ān by studying these sciences and
by interpreting its verses in a way meaningful to their time and conditions. This knowledge is the key to
unlocking the meanings contained within the Qur’ān and without it, satisfactory interpretation is
impossible.

THE ORIGIN OF THE WORD QUR'AN

The majority of Islamic scholars agree that the word ‘Qur’ān’ is an infinitive form of the verb ‘qa-ra-a’
meaning ‘reading’ or ‘reciting’. Therefore, it literally means ‘a thing recited by adding letters and words
to one another’. This verb has another infinitive form, qar’u, which means ‘to collect’. Based on this
form, some scholars are of the opinion that ‘Qur’ān’ means ‘the thing which collects’. There are other
scholars who affirm that the word ‘Qur’ān’ was not derived from any existing word. They hold that it is
the proper name given to the book which God sent to His final Messenger, Prophet Muhammad (may
Allah bestows peace, mercy and blessings upon him and his household members).

________________________

Rāghib al-Isfahānī, Mufradaat, p. 402

The Qur’ān has other titles, each of which describes one of its aspects and therefore can be regarded as
one of its attributes. Some of them are, the Book, the Criterion, the Remembrance, the Advice, the Light,
the Guidance, the Healer, the Noble, the Mother of the Book, the Truth, the Admonishment, the Good
Tiding, the Book Gradually Revealed, the Knowledge, and the Clear.

WHAT KIND OF THE BOOK QUR'AN WAS?

Since the Qur’ān is the direct word of God it can be described by Him best. So The Qur’ān describes itself
as;

Alif. Lam. Mim. This is the (most honoured, matchless) Book: there is no doubt about it (its divine
authorship and it being a collection of pure truths). It is guidance for the God-revering, pious who keep
their duty to God.

Commentary

One obvious meaning of this verse is that this Book, the Qur'an, is undoubtedly from God. Another
possible meaning is that nothing contained in it can be subject to doubt. Books which deal with
supernatural questions, with matters that lie beyond the range of sense perception, are invariably based
on conjecture and their authors, despite their brave show of competence, are therefore not immune
from a degree of scepticism regarding their statements. This Book, which is based wholly on Truth, a
Book which is the work of none other than the All-Knowing God Himself is distinguishable from all other
books. Hence, there is no room for doubt about its contents despite the hesitation some people might
express either through ignorance or folly.

3. This means that while the Book is potentially for all, only those who possess certain qualities can
benefit from it. The first such quality is piety: those who want to benefit should be disposed to
distinguish between good and evil, and to shun evil and do good. Those who lead an animal existence,
who never to consider whether their actions are either good or bad, whose cynically follow the
prevailing winds, who are helplessly tossed about by the animal desires that dominate their minds, such
persons are all together incapable of deriving any benefit from the guidance embodied in the Qur'an.

4. This is the second prerequisite for deriving benefit from the Qur'an. Ghayb signifies the verities which
are hidden from man's senses and which are beyond the scope of man's ordinary observation and
experience, for example the existence and attributes of God, the angels. the process of revelation,
Paradise, Hell and so on. 'Belief in the ghaib' means having faith in such matters, based on an absolute
confidence in the Messengers of God and despite the fact that it is impossible to experience them.
According to this verse, Qur'anic guidance can prove helpful only to those prepared to affirm the truths
of the suprasensory realm. People who make their belief in these questions conditional upon sensory
perception of the object of belief, and who are not prepared even to consider the possibility of the
existence of things that cannot be weighed or measured, cannot profit from this Book.

5. This is the third requirement. It is pointed out that those to whom belief means merely the
pronouncement of a formula, who think that a mere verbal confession of faith is enough and that it
makes no practical demands on them, can derive no guidance from the Qur'an. To benefit from the
Qur'an it is essential that a man's decision to believe should be followed immediately by practical
obedience to God.

Prayer is the first and continuing sign of practical obedience. No more than a few hours can pass after a
man has embraced Islam than the mu'adhin calls to Prayer and it becomes evident whether or not the
profession of faith has been genuine. Moreover, the mu'adhin calls to Prayer five times every day and
whenever a man fails to respond to his call it becomes clear that he has transgressed the bounds of
practical obedience. An abandonment of Prayer amounts to an abandonment of obedience. Obviously, if
a man is not prepared to follow the directives of his guide, it is immaterial whether or not true guidance
is available to him.

It should also be noted that the expression 'establishment of Prayer' has a wider meaning than mere
performance of Prayer. It means that the system of Prayer should be organized on a collective basis. If
there is a person in a locality who prays individually but no arrangements are made for congregational
Prayer, it cannot be claimed that Prayer is established in that locality.

6. This, the fourth prerequisite for a person to benefit from the Qur'an, demands that the person
concerned should neither be niggardly nor a worshipper of money. On the contrary, he should be willing
to pay the claims on his property of both God and man, and should not flinch from making financial
sacrifices for the sake of his convictions.

7. The fifth requirement is that one should believe in the Books revealed by God to His Prophets in the
various ages and regions of the world, in the Book revealed to Muhammad (peace be on him) as well as
in those revealed to the other Prophets who preceded him. The door of the Qur'an is closed to all those
who do not consider it necessary for man to receive guidance from God. It is also closed to those who,
even if they believe in the need for such guidance, do not consider it necessary to seek it through the
channel of revelation and prophethood, but would rather weave their own set of ideas and concepts
and regard them as equivalent to Divine Guidance.

This door is also closed to those who believe in Divine books as such, but confine this belief to those
books accepted by their forefathers, and spurn Divine Guidance revealed to anyone born beyond their
own racial and national boundaries. The Qur'an excludes all such people and is prepared to open the
source of its grace only to those who believe that mankind does require Divine Guidance, who
acknowledge that this guidance does not come to people individually but reaches them through
Prophets and Divine Books and who are not given to racial or national chauvinism but are devotees of
Truth alone, and are therefore prepared to submit to Divine Guidance wherever it be found.
8. Belief in the After-life is the sixth and last requirement. The term al-Akhirah embraces a whole set of
ideas: (i) that man is not an irresponsible being, but is answerable to God for all his conduct in this
world; (ii) that the present order of the world is not timeless, but will come to an end at an appointed
hour known only to God; (iii) that when this world comes to an end God will bring into being another
world in which He will resurrect, at one and the same moment, all the human beings ever born on earth.
He will gather them together, examine their conduct and grant each one just reward for his actions; (iv)
that those who are accounted good in God's judgement will be sent to Heaven, and those judged by Him
as evil-doers will be consigned to Hell; (v) that the real measure of success and failure is not one's
prosperity in the present life, but one's success or failure according to God's judgement in the Next.
Those who do not accept this set of beliefs can derive no benefit from the Qur'an. For if a man is merely
in a state of doubt and hesitation with regard to these matters - let alone disbelieving them - he cannot
advance even one step forward along the path charted out by the Qur'an.

And also this verse (i.e. verse 2 of surah baqara) can further be explained as illustrated below:

1. Alif Lam Mim}

The Individual Letters

The individual letters in the beginning of some Surahs are among those things whose knowledge Allah
has kept only for Himself.

These Letters testify to the Miraculous Qur’an

The wisdom behind mentioning these letters in the beginning of the Surahs, regardless of the exact
meanings of these letters, is that they testify to the miracle of the Qur’an. Indeed, the servants are
unable to produce something like the Qur’an, although it is comprised of the same letters with which
they speak to each other. Every Surah that begins with these letters demonstrates the Qur’an’s miracle
and magnificence, and this fact is known by those well-versed in such matters.

{2. That is Book in which there is no Rayb, guidance for the Muttaqin}

There is no Doubt in the Qur’an

Several Companions of the Messenger of Allah, peace and blessings of Allah be upon him, narrated that,
“In which there is no Rayb”, means about which there is no doubt. Similarly, Allah said in Surat As-
Sajdah,

“Alif Lam Mim. The revelation of the Book (this Qur’an) in which there is no doubt, is from the Lord of all
that exists” (32:1-2).

Guidance is granted to Those Who have Taqwa

Hidayah – correct guidance – is only granted to those who have Taqwa – fear of Allah. Allah said ,
“Say: It is for those who believe, a guide and a healing. And as for those who disbelieve, there is
heaviness (deafness) in their ears, and it (the Qur’an) is blindness for them. They are those who are
called from a place far away (so they neither listen nor understand)” (41:44), and,

“And We send down of the Qur’an that which is a healing and a mercy to those who believe (in Islamic
Monotheism and act on it), and it increases the Zalimin (wrongdoers) in nothing but loss” (17:82).

This is a sample of the numerous Ayat indicating that the believers, in particular, benefit from the
Qur’an. That is because the Qur’an is itself a form of guidance, but the guidance in it is only granted to
the righteous, just as Allah said,

“O mankind! There has come to you a good advice from your Lord (i. e. the Qur’an, enjoining all that is
good and forbidding all that is evil), and a healing for that (disease of ignorance, doubt, hypocrisy and
differences) which is in your breasts, – a guidance and a mercy (explaining lawful and unlawful things)
for the believers” (10:57).

Ibn ‘Abbas and Ibn Mas’ud and other Companions of the Messenger of Allah, peace and blessings of
Allah be upon him, said, “guidance for the Muttaqin (the pious and righteous persons”, means, a light
for those who have Taqwa.

The Meaning of Al-Muttaqin

Ibn Abbas said about “guidance for the Muttaqin” that it means, “They are the believers who avoid Shirk
with Allah and who work in His obedience.” Ibn Abbas also said that Al-Muttaqin means, “Those who
fear Allah’s punishment, which would result if they abandoned the true guidance that they recognize
and know. They also hope in Allah’s mercy by believing in what He revealed.” Further, Qatadah said
that: “Al-Muttaqin’, are those whom Allah has described in His statement: “Who believe in the Ghayb
and perform the Salah” (2:3), and the following Ayat. Ibn Jarir stated that the Ayah (2:2) includes all of
these meanings that the scholars have mentioned, and this is the correct view. Also, At-Tirmidhi and Ibn
Majah narrated that `Atiyah As-Sa`di said that the Messenger of Allah, peace and blessings of Allah be
upon him, said,

“The servant will not acquire the status of the Muttaqin until he abandons what is harmless out of fear
of falling into that which is harmful.” (Tuhfat Al-Ahwadi 7:147)

There are Two Types of Hidayah (Guidance)

Huda here means the faith that resides in the heart, and only Allah is able to create it in the heart of the
servants. Allah said,

“Verily, you (O Muhammad ) guide not whom you like” (28:56),

“Whomsoever Allah sends astray, none can guide him” (7:186), and,

“He whom Allah guides, he is the rightly guided; but he whom He sends astray, for him you will find no
Wali (guiding friend) to lead him (to the right path)” (18:17).
“Not upon you (Muhammad ) is their guidance” (2:272)

Huda also means to explain the truth, give direction and lead to it. Allah, the Exalted, said,

“And verily, you (O Muhammad ) are indeed guiding (mankind) to the straight path (i.e. Allah’s religion
of Islamic Monotheism)” (42: 52),

“You are only a warner, and to every people there is a guide” (13:7), and,

“And as for Thamud, We showed and made clear to them the path of truth (Islamic Monotheism)
through Our Messenger (i.e. showed them the way of success), but they preferred blindness to
guidance” (41:17).

testifying to this meaning. Also, Allah said,

“And shown him the two ways (good and evil).” (90:10)

This is the view of the scholars who said that the two ways refer to the paths of righteousness and evil,
which is also the correct explanation. And Allah knows best.

Meaning of Taqwa

The root meaning of Taqwa is to avoid what one dislikes. It was reported that Umar bin Al- Khattab
asked Ubayy bin Ka`b about Taqwa. Ubayy said, “Have you ever walked on a path that has thorns on it?”
Umar said, “Yes.” Ubayy said, “What did you do then?” He said, “I rolled up my sleeves and struggled.”
Ubayy said, “That is Taqwa.”The Qur’ān is a collection of revelations, compiled as a book, which were
conveyed to Prophet Muhammad (may Allah bestows peace, mercy and blessings upon him and his
household members) by the angel Gabriel over a period of twenty-three years. Whenever the Prophet
(may Allah bestows peace, mercy and blessings upon him and his household members) received a verse,
he instructed scribes to write it down and they collectively wrote every single verse of the Qur’ān. After
the Prophet (may Allah bestows peace, mercy and blessings upon him and his household members)
passed away, the Qur’ān was transmitted by the most authentic narrations (mutawātir) and has been
preserved without a single change up until the present day. It is the verbatim Word of God (Allah) which
has multiple aspects and its recitation is considered as worship in Islam. The Qur’ān is the holy book for
Muslims but it also represents the best example of Arab literature. In fact, the Qur’ān is the oldest
written form of Arabic literature.The Qur’ān is beyond comparison with regards to its style, parables,
method of explaining realities, its sciences, its description of God by His names, attributes and essential
qualities, its guidance methodology, the information about the unseen world and many other areas.
Bediuzzaman Said Nursi (1878 – 1960 C.E.) depicted this reality in his statement:The Qur’ān is an eternal
translation of the great Book of the Universe and everlasting translator of it multifarious tongues reciting
the Divine laws of the universe’s creation and operation; the interpreter of the books of the visible,
material world and the World of the Unseen; the discloser of the immaterial treasuries of the Divine
Names hidden on the earth and in the heavens; the key to the truths lying behind events; the World of
the Unseen’s tongue in the visible, material one; the treasury of the All-Merciful One’s favours and the
All-Glorified One’s eternal addresses coming from the World of the Unseen beyond the veil of this
visible world; the sun of Islam’s spiritual and intellectual world, as well as its foundation and plan; the
sacred map of the worlds of the Hereafter; the expounder, lucid interpreter, articulate proof, and clear
translator of the Divine Essence, Attributes, Names and essential Qualities; the educator and trainer of
the world of humanity and the water and light of Islam, which is the true and greatest humanity; and the
true guide of humanity leading them to happiness. For humanity, it is both a book of law, and a book of
prayer, and a book of wisdom, and a book of worship and servanthood to God, and a book of command
and call to God, and a book of invocation, and a book of thought and reflection. It is a comprehensive,
holy book containing books for all spiritual needs of humanity; a heavenly book that, like a sacred
library, offers numerous booklets from which all saints, eminently truthful people, all discerning and
verifying scholars, and those well-versed in knowledge of God have derived their own specific ways, and
which illuminate each way and answer their followers’ needs.

___________________________

Said Nursi, Words, The Twenty-fifth Word, p. 388The Qur’ān differs from all other divinely revealed
books in regard to its time of revelation, its effect on people, satisfying their worldly and spiritual needs
in every aspect, its style, language and transmission. God endowed Moses and Jesus with miracles that
were most suitable for their time and people; He gave the miracles of the staff and the bright hand to
Moses, and gave the miracles of raising the dead and healing certain illnesses to Jesus. The Qur’ān is the
chief miracle of Prophet Muhammad (may Allah bestows peace, mercy and blessings upon him and his
household members) and its eloquence is its most notable attribute. At the time of the Prophet (may
Allah bestows peace, mercy and blessings upon him and his household members) most people of the
Arabian Peninsula were illiterate and they preserved their tribal pride, history, and proverbs in oral
poetry. Eloquence was esteemed and highly prized so new or meaningful, unique expressions were
memorized for their poetic form and eloquence and then handed down to posterity. Eloquence was so
important that wars between tribes were started and ended on the words of prominent literary figures.
The wall of the Ka’ba was adorned with gold inscriptions of the odes of seven poets, called the ‘Seven
Suspended Poems’. The Qur’anic revelation challenged the literary figures of the Arabian Peninsula: ‘If
you doubt what We have sent down on Our servant, produce a surah like it’. As previously mentioned,
this challenge has not been met by anyone up until now.The Qur’ān speaks to the layman and the
profound sage simultaneously, making the Qur’ān both universal and timeless. It is extremely difficult, if
not impossible, to address different people with different intellectual capacities at the same time
adequately, yet the Qur’ān (being guidance for all people and the Word of God) comprehensively
addresses all people without ignoring or causing injustice to anyone. The Qur’ān aims to guide all people
to the truth and has four main purposes and themes: demonstrating God’s existence and unity,
establishing Prophethood, proving and elucidating the afterlife with all its aspects and dimensions and
promulgating the worship of God and the essentials of justice.

____________________________

Ali Unal, The Qur’ān with Annotated Interpretation, foreword

The Qur’ān is also a source of healing; its application in life provides a cure for almost all psychological
and social illnesses. Additionally it is a book of cosmology, epistemology, ontology, sociology,
psychology, and law. It was revealed to regulate human life in this world and it is not bound or limited to
any time, place or people. It is for all times and for all peoples, universal and timeless. The most
distinguished miracle of the Qur’ān is in its letters, words and verses. The Qur’ān uses the Arabic
alphabet and brings letters together to create words, verses and surah. It would seem rather easy to
merge letters together and produce similar words to that of the Qur’ān. However, as history has
attested, even the most eloquent Arab poets such as Walid, Labīd, Ā’shā and Ka’b bin Zuhayr could not
produce similar words, let alone similar chapters, because the Qur’ān is a miracle and inimitable. The
Prophet (may Allah bestows peace, mercy and blessings upon him and his household members) was
unlettered as attested in the Qur’ān:

You did not recite before it any scripture, nor did you inscribe one with your right hand. Otherwise the
falsifiers would have had cause for doubt.

__________________________

Qur’ān, 29: 48

Despite their lack of belief in the Qur’ān, polytheist leaders in Mecca could not resist listening to its
recitation, due to its appealing nature and their appreciation of poetry. They would secretly go to the
Ka’ba to listen to the Prophet (may Allah bestows peace, mercy and blessings upon him and his
household members) reciting during prayer. ‘Umar b. al-Khattāb was prepared to kill the Prophet (may
Allah bestows peace, mercy and blessings upon him and his household members) yet upon hearing the
recitation from Surah Ta-ha his heart was illuminated with the words of the Qur’ān and he
consequentially converted to Islam.

________________________

Ibn Hishām, Sīrah, vol.2 p. 187

Utba ibn Rabia set out to the Prophet’s (may Allah bestows peace, mercy and blessings upon him and his
household members) house with the sole purpose of defeating the Prophet (may Allah bestows peace,
mercy and blessings upon him and his household members) in a debate. The Prophet (may Allah
bestows peace, mercy and blessings upon him and his household members) listened silently to all Utbas’
arguments. When he had finished, the Prophet (may Allah bestows peace, mercy and blessings upon
him and his household members) recited the first thirteen verses of Surah Fussilat. Utba listened in
shock and was greatly affected. When he returned to his friends, who were awaiting news of the
outcome of the debate, Utba responded: ‘I have listened to such words from him that I have never
heard before. This is neither poetry, nor magic nor the words of soothsayers. It is not like of any of
these. O Quraysh, pay attention and listen to me, leave him alone. If he cannot be successful the Arabs
will destroy him, but if he is successful, his success is your success’.

__________________________
Ibn Hishām, Sīrah, 1/294

Once Tufayl ibn Amir came to Mecca and was told by the polytheist leaders: ‘O Tufayl, you came to our
land; people who listen to Muhammad (may Allah bestows peace, mercy and blessings upon him and his
household members) have separated from us, and our unity is being dissolved because of him. His
words are very effective, like magic which separates a man from his father, his brother and even his
wife. We are worried about you and your tribe, be careful and never listen to him’. Tufayl was scared by
their warnings and whenever he went to Ka’ba, he refrained from listening to Muhammad (may Allah
bestows peace, mercy and blessings upon him and his household members). He would even close his
ears so as to not hear the words of the Prophet (may Allah bestows peace, mercy and blessings upon
him and his household members). One day he said to himself, ‘I am in a false belief, what is the harm in
listening to his words, if the words are true; I have the capacity to understand it’. After that he listened
to the Qur’ān and consequently converted to Islam. He was the first Muslim guide in his tribe.

_________________________

Ibn Hishām, Sīrah, 1/382Sometimes listening to just one verse was enough for people to understand the
Qur’ān and appreciate its value. When Sa’sa the uncle of Farazdaq heard the following verse: ‘So
whoever does an atom’s weight of good will see it, and whoever does an atom’s weight of evil will see it’
he said, ‘it is enough for me’ and he became Muslim.

_________________________

Qurtubī, al-Jami’ al-Aḫkām al-Qur’ān, 20/153

Arabs achieved deepened levels of understanding through the Qur’ān. An example was when Asma’i
heard a pleasant poem from a village girl, he commented ‘What a beautiful poem, how eloquent it
is’.Thereupon the girl replied, ‘What a bad man you are, my words are not more eloquent than this
verse; ‘We inspired to the mother of Moses, ‘Suckle him; but when you fear for him, cast him into the
river and do not fear and do not grieve. Indeed, We will return him to you and will make him one of the
Messengers’’.

_______________________

Qur’ān, 28: 7

She added; this verse contains two commands, two prohibitions and two good tidings. {Qurtubī, al-Jami’
al-Aḫkām al-Qur’ān, 13/252}.

At the beginning of Islam, the Meccan leaders tried to refute the Qur’ān saying that Muhammad (may
Allah bestows peace, mercy and blessings upon him and his household members) was insane or a poet
or a soothsayer, but the majority of them eventually accepted Islam. In a short period of time, they
understood that the Qur’ān was a divine revelation, a miracle and impossible to challenge.
The Arabs were differentiating between real and false Prophethood. For example, Talha An~Namari
came to Yamama to speak with Musaylama {a false prophet from the Bani Hanifa tribe during the same
time as Muhammad}. After speaking with him, Talha said, ‘I testify that you are a liar, you are not a real
Prophet, and I testify that Muhammad (may Allah bestows peace, mercy and blessings upon him and his
household members) is the real and true Prophet (may Allah bestows peace, mercy and blessings upon
him and his household members). But the liar of the tribe Rabia is more beloved for me than the true
Prophet of the Mudar tribe’.

_________________________

Mustafa Sadiq ar-Rafi, I’jaz’ul Qur’ān, 195

Indeed, the impossibility of challenging or producing something similar to the Qur’ān was obvious to the
Arabs, and despite some of their efforts to do so, they stood defeated confessing their inability to do
this.

_________________________

Mustafa Sadiq ar-Rafi, I’jaz’ul Qur’ān, 187-212One day, a great philosopher Kindi was asked by his
friends to write something similar to the Qur’ān. He responded ‘I can do it for some part of the Qur’ān’.
After secluding himself and dedicating his time to the study of the Qur’ān in efforts to fulfil his friends’
challenge, he attested to his friend ‘I swear by God, I am incapable of doing this and I don’t think
anybody can do this either. When I opened the Qur’ān I encountered Surah Maida, the first verse is ‘O
you who have believed, fulfil all contracts. Lawful for you are the animals of grazing livestock except for
that which is recited to you in this Qur’ān, hunting not being permitted while you are in the state of
iḫrām. Indeed, God ordains what He intends’. This verse starts with the fulfilment of contracts, then it
prohibits breaking them, it regulates a general law about the meat of animals and then makes an
exception from this general rule. After that God declares his power and wisdom. All these things cannot
be phrased in two lines; no one can fit these things in small sentence, it is necessary to write volumes of
books to explain these things’.

____________________

Qurtubī, al-Jami’ al-Aḫkām al-Qur’ān, 8/31-32Finally, it is imperative to note the definition of the Qur’ān
given by the Prophet (may Allah bestows peace, mercy and blessings upon him and his household
members):The superiority of the Qur’ān over all other words and speeches is like God’s superiority over
His creatures. The Qur’ān is a definite decree distinguishing between truth and falsehood. It is not for
pastime. Whoever rejects it because of his or her despotism, God breaks his or her neck. It contains the
history of previous peoples, the tidings of those to come after you, and the judgment on the
disagreements between you.Whoever searches for guidance in something other than it, God leads him
or her astray. It is God’s strong rope. It is the wise instruction. It is the Straight Path. It is a book which
desires cannot deviate and tongues cannot confuse, and which scholars are not fed up with, never worn-
out by repetition, and has uncountable admirable aspects. It is such a book that they could not help but
say: ‘We have indeed heard a wonderful Qur’ān, guiding to what is right in belief and action and so we
have believed in it.’ Whoever speaks based on it speaks the truth; whoever judges by it, judges justly
and whoever calls to it, calls to the truth.

__________________________

Tirmidhi, Sunan, Thawab al-Qur’ān, 14.

The revelation of the Qur'an was considered by a Muslims to Muhammad's greatest miracle, and a
miracle for all the times. Unlike the miracles of other prophets which confirmed to be witness in their
own lifetime.

The holy Qur'an involves everything including warning to avoid the unlawful, and a command to do the
righteous good deeds, and the story of faithfully believed persons, and the story of the greatest
disbelievers. In fact is a guide for those who fear Allah.

So that Allah (the exalted and the merciful) says about Qur'an.

‫َت َبا َر َك ا َّل ِذى َن َّز َل ا ْل ُف ْر َقا َن َع ٰلى َع ْب ِد ِهۦ ِل َي ُكو َن ِل ْل ٰع َل ِمي َن َن ِذي ًرا‬

"Blessed is He who sent down the Criterion upon His Servant that he may be to the worlds a warner -"

(QS. Al Furqaan 25: Verse 1)

Commentary

1. The Arabic word tabaraka is very comprehensive, and cannot be understood fully and completely by
“blessed”, not even by a sentence. However, its meanings may be grasped by keeping in view the
remaining portion of (verses 1-2). Here it has been used to convey the following meanings:

(1) He is the most Beneficent: that is why He has bestowed the great blessing of Al-Furqan by degrees on
His servant so that he may admonish all mankind.

(2) He is the most Exalted and Great: for the Sovereignty of the heavens and the earth belongs to Him.

(3) He is the most Holy, Pure and Perfect: He is free from every tinge of shirk and has neither a partner
in His Godhead, nor needs a son to succeed Him; for He is Ever- Lasting.

(4) He is the Highest and the Most Supreme in rank: the Kingdom and the Dominion wholly and solely
belong to Him, and there is none who has any share in His Authority and Powers.

(5) He is the sole Creator of the universe and has created each and everything in it and predetermined
its destiny. For further details, see (E.N. 14 of Al-Mominoon) and (E.N. 19 of Al-Furqan).

2. Al-Furqan: The Criterion. The Quran has been called Al-Furqan because it is the Criterion for judging
right and wrong virtue and vice, truth and falsehood.
3. The word nazzala implies revelation of the Quran piecemeal by degrees. The wisdom of this
introductory remark will be explained in the study of verse 32, where the objection of the disbelievers of
Makkah as to why the Quran has not been sent down all at once, has been dealt with.

4. “Warner to all mankind”: To warn all mankind of the evil consequences of their heedlessness and
deviation. The warner may be Al-Furqan or the Prophet (peace be upon him) to whom it was revealed.
In fact, both were the warners because they were both sent for one and the same purpose. The
message of the Quran and Prophethood of Muhammad (peace be upon him) were not meant for any
particular country but for the whole world; and not for their own time, but for all times to come. This
has been stated at several places in the Quran. For instance: O Muhammad, say, O mankind, I am a
Messenger to all of you from Allah (Surah Al-Aaraf, Ayat 158). And this Quran has been revealed to me
so that I should thereby warn you all and to whom it may reach. (Surah Al Anaam, Ayat 19). We have
sent you only as a bearer of good news and as a warner to all mankind. (Surah Saba, Ayat 28). We have
sent you as a mercy to all mankind. (Surah Al Anbiya, Ayat 107). The Prophet (peace be upon him)
himself has stated this clearly in the Hadith. For instance, he said: I have been sent to all men, the red
and the black. Before me a Prophet was sent only to his own people, but 1 have been sent to all
mankind. (Bukhari, Muslim). I have been sent to all mankind, and I am the last of the Prophets. (Muslim).

5. “To Him belongs the Sovereignty of the heavens and the earth.” That is, He alone has an exclusive
right to it, and no one else has any right to it nor any share in it.

6. That is, He has neither any relation of direct parenthood to anyone, nor has He taken anyone as a son.
Therefore, none else in the universe is entitled to worship. He is Unique and there can be no partner in
His Godhead. Thus all those who associate with Him angels or jinns or saints as His offspring, are
ignorant. Likewise, those who believe that someone is His son, are also ignorant. They have no true
conception of the Greatness of Allah and consider Him to be weak and needy like human beings, who
require someone to become their inheritor. It is sheer ignorance and folly. For further details, see (E.Ns
66 to 68 of Surah Younus).

7. The Arabic word mulk means Sovereignty, Supreme Authority, and Kingship. Thus the sentence will
mean: Allah is the Absolute Ruler of the whole universe and there is none other who may have any right
to authority; therefore He alone is God. For, whenever a man takes anything else as his lord, he does so
under the presumption that his deity has the power to do good or bring harm and make or mar his
fortune; nobody will like to worship a powerless deity. Now when it is recognized that none but Allah
has the real power and authority in the universe, nobody will bow before anyone other than Him in
worship, nor will sing anybody else’s hymns, nor commit the folly of bowing in worship before anything
else except his real God, or recognize any other as his ruler, because “To Allah belongs the Sovereignty
of the heavens and the earth and to Him alone.”

8. There may be other translations of this also: He has ordained it in due proportion; or He has
appointed an exact measure for everything. But no translation can convey its real meaning, which is:
Allah has not only created everything in the universe but also determined its shape, size, potentialities,
characteristics, term of existence, the limitations and extent of its development and all other things
concerning it. Then, He has created the means and provisions to enable it to function properly in its own
separate sphere.

This is one of the most comprehensive verses of the Quran with regard to the doctrine of Tauhid.
According to traditions, the Prophet (peace be upon him) himself taught this verse to every child of his
family as soon as it was able to speak and utter a few words. Thus, this verse is the best means of
impressing the doctrine of Tauhid on our minds, and every Muslim should use it for educating his
children as soon as they develop understanding.

But according to famous work of tafsir ishraq Al-ma'ani it has been proved that:

1. The meaning of “tabaaraka” as given here reflects the understanding of Ibn `Abbas as in Ibn Jarir, and
of Zajjaj as in Qurtubi.

The report about Ibn `Abbas is in Ibn Abi Hatim also (Shawkani).

Linguistically however, as Zamakhshari, Razi and others point out, the word in the root is barakah which
means increase in (all kinds of) good and beneficence. In this case, tabaaraka would point to overflow of
benefits and goodness from Allah. It is outflow of good that resulted in the revelation of the Qur’an, and
hence the importance of knowledge (Razi). There have been other explanations, quite philosophic, by
Razi (though in greater detail by Alusi) while Qurtubi, as well as others inform us that the root word
leads us to another meaning: that of permanence and eternity of Allah’s Attributes.

Ibn al-Qayyim explains that the attribution of “barakah” to Allah means attribution of the Qualities of
mercy, power and honor to Him. In verb form the word becomes “tabaaraka” which is applicable only to
Allah and none else. Hence, he on whom He bestowed “barakah” is mubaarak, e.g., the mubaarak Book,
the mubaarak Messenger, etc. Here, however, as well as in all other places in the Qur’an, the word
“tabaaraka” has been used as an adjective and not as a verb (Badaa’i`).

Yusuf Ali adds: “Tabaaraka: the root meaning is "increase" or "abundance". Here that aspect of Allah's
dealing with His creatures is emphasised, which shows abundant goodness to all His creatures, in that
He sent the Revelation of His Will, not only in the unlimited Book of Nature, but in a definite Book in
human language, which gives clear directions and admonitions to all. The English word ‘blessed’ hardly
conveys that meaning.”

2. The textual word for “sending down” is “nazzala” which is a verb form used for expressing
exaggeration or emphasis. Here it has been used to emphasize the gradual, and therefore, fragmentized
sending down of the revelation as against the previous Scriptures that were sent down as one whole
(Ibn Kathir). As Allah said (3: 3), points out Razi:

{‫[ } َن َّز َل َع َل ْي َك ا ْل ِك َتا َب ِبا ْل َح ِّق ُم َص ِّد ًقا ِل َما َب ْي َن َي َد ْي ِه َو َأ ْن َز َل ال َّت ْو َرا َة َوا ْل ِإ ْن ِجي َل‬3 :‫] آل عمران‬

“He has sent down upon you the Book with the truth, confirming that which preceded it (as) He sent
down the Tawrah and the Injil.”
In this ayah, Allah used two different words for two different Revelations. He used nazzala for the
Qur’an, while anzala (which is for sending down in one installment) for Torah and Injeel.

4. Yusuf Ali comments on “al-furqaan”: “That by which we can judge clearly between right and wrong.
Here the reference is to the Qur’an, which has already been symbolised by light. This symbol is
continued here, and many contrasts are shown, in the midst of which we can distinguish between the
true and the false by Allah's Light, especially the contrast between righteousness and sin.þ”

A second meaning, as most commentators have pointed out, is that of a separator – from faraqa: to
separate – in this case, the separator of truth and falsehood.”

5. `Abd (slave) is an honorific title. The best that one can do is to be an `abd of Allah. When Allah uses
the term for a person, it means He holds that person in high repute. Hence, on all important occasions
Prophet Muhammad was referred to as an `abd in the Qur`an (Ibn Kathir).

6. Ibn Zayd has said that the pronoun of li-yakuna is for the Prophet. He was and remains a warner unto
the worlds. Allah said (35: 24),

{‫[ } َو ِإ ْن ِم ْن ُأ َّم ٍة ِإ َّلا َخ َلا ِفي َها َن ِذي ٌر‬24 :‫]فاطر‬

“And there hasn’t been a nation except a warner had been in it.”

And (26: 208),

{‫[ } َو َما َأ ْه َل ْك َنا ِم ْن َق ْر َي ٍة ِإ َّلا َل َها ُم ْن ِذ ُرو َن‬208 :‫]الشعراء‬

“And We did not destroy a town but it had its warners.”

Initially, there was only one warner: the Messenger. After he had passed away, he became a warner
unto everyone who received the message after him. The Qur’an said (6: 19),

{‫[ } َو ُأو ِح َي ِإ َل َّي َه َذا ا ْل ُق ْر آ ُن ِل ُأ ْن ِذ َر ُك ْم ِب ِه َو َم ْن َب َل َغ‬19 :‫]ال أنعام‬

“And this Qur’an has been revealed to me in order that I warn you and those whom it reaches.”

That is, those that the Qur’an reaches. Hence the Prophet is a warner unto the worlds. The Qur’an told
him to announce (7: 158),

{‫[ } ُق ْل َيا َأ ُّي َها ال َّنا ُس ِإ ِّني َر ُسو ُل ال َّل ِه ِإ َل ْي ُك ْم َج ِمي ًعا‬158 :‫]ال أعراف‬

“O people. I am indeed Allah’s Messenger unto you all.”

Ibn Zayd also said that Allah never sent a Messenger to the entire world population except Nuh, with
whom He began a new creation, and Muhammad, with whom He ended (the series) – Ibn Jarir.

Qurtubi also explains the verse in the above manner.

The Prophet said,

‫ُب ِع ْث ُت ِإ َلى ال َّنا ِس َكا َّف ًة ال َأ ْح َم ِر َوال َأ ْس َو ِد‬

“I have been sent to the entire mankind, the reds and the blacks.”

The above report of Ahmad was evaluated as Hasan by Shu`ayb al-Arna’ud (Au.).

He also said,

‫ُأ ْع ِطي ُت َخ ْم ًسا َل ْم ُي ْع َط ُه َّن َأ َح ٌد َق ْب ِلي … َو َكا َن ال َّن ِب ُّي ُي ْب َع ُث ِإ َلى َق ْو ِم ِه َخا َّص ًة َو ُب ِع ْث ُت ِإ َلى ال َّنا ِس َعا َّم ًة‬

“I enjoy specialty in five things...and a Prophet used to be raised for his own people alone while I have
been raised for the entire mankind.”

He mentioned as fifth that while others were sent to a particular nation, he was sent to all the peoples
(Ibn Kathir).

Bukhari and Muslim recorded it (Au.).

7. “The worlds” – i.e., (the worlds of) the Jinn and mankind (Zamakhshari, Razi).

Sayyid adds: “The words, ‘that he may be a warner unto the worlds’ appearing in a Makkan Surah leave
no room that from the first day of revelation this message was meant for all mankind; and not, as some
historians state, that originally the message was for the Arabs alone; later, with the subjugation of the
Arabs, the Prophet declared it for others too, as a later thought and introduction. This ayah disproves
this theory.”

___________________

Tafsir ishraq Al-ma'ani

While according to famous tafsir book of tafsir Tafhimul Qur'an by Sayyida Abul Ala maududi he
explained the verse as:

Al-Furqan: The Criterion. The Qur'an has been called Al-Furqan because it is the Criterion for judging
right and wrong virtue and vice, truth and falsehood.

The word nazzala implies revelation of the Qur'an piecemeal by degrees. The wisdom of this
introductory remark will be explained in the study of verse 32, where the objection of the disbelievers of
Makkah -as to why the Qur'an has not been sent down all at once-has been dealt with

“..... warner to all mankind": To warn all mankind of the evil consequences of their heedlessness and
deviation. The warner may be AI-Furgan or the Holy Prophet to whom it was revealed. In fact, both were
the warners because they were both sent for one and the same purpose. The message of the Qur'an and
Prophethood of Muhammad (Allah's peace and blessings be upon him) were not meant for any
particular country but for the whole world; and not for their own time, but for all times to come. This
has been stated at several places in the Qur'an; for instance: "O Muhammad, say, O mankind, I am a
Messenger to all of you from Allah ....." (Al-A`raf: 158); "..... and this Qur'an has been revealed to me so
that I should thereby warn you all and whom it may reach." (Al An'am: 19); "We have sent you only as a
bearer of good news and as a warner to all mankind." (Saba: 28); and "We have sent you as a mercy to
all mankind." (Al Anbiya': 107) The Holy Prophet himself has stated this clearly in the Hadith; for
instance, he said: "I have been sent to all men-the red and the black." "Before me a Prophet was sent
only to his own people, but 1 have been sent to all mankind." (Bukhari, Muslim). "I have been sent to all
mankind, and I am the Last of the Prophets." (Muslim).

______________

Tafhimul Qur'an by Sayyid Abul Ala maududi,

The versatility of the Holy Qur'an is beyond perception as it is the book comprises of everything and
every knowledge indulge in it by Allah the most wise the merciful. It is the book to which there is no
doubt, a guide to the seekers of truth and guidance as Allah proved:

‫ٰذ ِل َك ا ْل ِك ٰت ُب َلا َر ْي َب ۛ  ِفي ِه ۛ  ُه ًدى ِّل ْل ُم َّت ِقي َن‬

"This is the Book about which there is no doubt, a guidance for those conscious of Allah -"

(QS. Al Baqara 2: Verse 2)

Commentary

Ibn Kathir (Abridged)

Ma'arif Al-Qur'an

There is no Doubt in the Qur'an

The Book, is the Qur'an, and Rayb means doubt. As-Suddi said that Abu Malik and Abu Salih narrated
from Ibn `Abbas, and Murrah Al-Hamadani narrated from Ibn Mas`ud and several other Companions of
the Messenger of Allah ‫ ﷺ‬that,

‫ل َا َر ْي َب ِفي ِه‬

(In which there is no Rayb), means about which there is no doubt. Abu Ad-Darda', Ibn `Abbas, Mujahid,
Sa`id bin Jubayr, Abu Malik, Nafi` `Ata', Abu Al-`Aliyah, Ar-Rabi` bin Anas, Muqatil bin Hayyan, As-Suddi,
Qatadah and Isma`il bin Abi Khalid said similarly. In addition, Ibn Abi Hatim said, "I do not know of any
disagreement over this explanation." The meaning of this is that the Book, the Qur'an, is without a
doubt revealed from Allah. Similarly, Allah said in Surat As- Sajdah,

‫ َذ ِل َك ا ْل ِك َتا ُب ل َا َر ْي َب ِفي ِه ُه ًدى ِّل ْل ُم َّت ِقي َن‬- ‫الم‬

(Alif Lam Mim). The revelation of the Book (this Qur'an) in which there is no doubt, is from the Lord of all
that exists) (2:12).
Some scholars stated that this Ayah - 2:2 - contains a prohibition meaning, "Do not doubt the Qur'an."
Furthermore, some of the reciters of the Qur'an pause upon reading,

‫ل َا َر ْي َب‬

(there is no doubt) and they then continue;

‫ِفي ِه ُه ًدى ِّل ْل ُم َّت ِقي َن‬

(in which there is guidance for the Muttaqin (the pious and righteous persons)). However, it is better to
pause at,

‫ل َا َر ْي َب ِفي ِه‬

(in which there is no doubt) because in this case,

‫ُه ًدى‬

(guidance) becomes an attribute of the Qur'an and carries a better meaning than,

‫ِفي ِه ُه ًدى‬

(in which there is guidance).

Guidance is granted to Those Who have Taqwa

Hidayah - correct guidance - is only granted to those who have Taqwa - fear of Allah. Allah said,

‫ُق ْل ُه َو ِل َّل ِذي َن َءا َم ُنو ْا ُه ًدى َو ِش َف آ ٌء َوا َّل ِذي َن ل َا ُي ْؤ ِم ُنو َن ِفى َءا َذا ِن ِه ْم َو ْق ٌر َو ُه َو َع َل ْي ِه ْم َع ًمى ُأ ْو َلـ ِئ َك ُي َنا َد ْو َن ِمن َّم َكا ٍن َب ِعي ٍد‬

(Say: It is for those who believe, a guide and a healing. And as for those who disbelieve, there is
heaviness (deafness) in their ears, and it (the Qur'an) is blindness for them. They are those who are
called from a place far away (so they neither listen nor understand)) (41:44), and,

‫َو ُن َن ِّز ُل ِم َن ا ْل ُق ْر َء ا ِن َم ا ُه َو ِش َف آ ٌء َو َر ْح َم ٌة ِّل ْل ُم ْؤ ِم ِن ي َن َو ل َا َي ِز ي ُد ا ل َّظ ـ ِل ِم ي َن َإ ل َّا َخ َس ا ًر ا‬

(And We send down of the Qur'an that which is a healing and a mercy to those who believe (in Islamic
Monotheism and act on it), and it increases the Zalimin (wrongdoers) in nothing but loss) (17:82).

This is a sample of the numerous Ayat indicating that the believers, in particular, benefit from the
Qur'an. That is because the Qur'an is itself a form of guidance, but the guidance in it is only granted to
the righteous, just as Allah said,

‫َي َأ ُّي َها ال َّنا ُس َق ْد َج آ َء ْت ُك ْم َّم ْو ِع َظ ٌة َّمن َّر ِّب ُك ْم َو ِش َف آ ٌء ِّل َما ِفى ال ُّص ُدو ِر َو ُه ًدى َو َر ْح َم ٌة ِّل ْل ُم ْؤ ِم ِني َن‬

(O mankind! There has come to you a good advice from your Lord (i. e. the Qur'an, enjoining all that is
good and forbidding all that is evil), and a healing for that (disease of ignorance, doubt, hypocrisy and
differences) which is in your breasts, ‫ ـ‬a guidance and a mercy (explaining lawful and unlawful things) for
the believers) (10:57).
Ibn `Abbas and Ibn Mas`ud and other Companions of the Messenger of Allah ‫ ﷺ‬said,

‫ُه ًدى ِّل ْل ُم َّت ِقي َن‬

(guidance for the Muttaqin (the pious and righteous persons), means, a light for those who have Taqwa.

The Meaning of Al-Muttaqin

Ibn `Abbas said about,

‫ُه ًدى ِّل ْل ُم َّت ِقي َن‬

(guidance for the Muttaqin) that it means, "They are the believers who avoid Shirk with Allah and who
work in His obedience." Ibn `Abbas also said that Al-Muttaqin means, "Those who fear Allah's
punishment, which would result if they abandoned the true guidance that they recognize and know.
They also hope in Allah's mercy by believing in what He revealed." Further, Qatadah said that,

‫ِّل ْل ُم َّت ِقي َن‬

(Al-Muttakin), are those whom Allah has described in His statement;

‫ا َّل ِذي َن ُي ْؤ ِم ُنو َن ِبا ْل َغ ْي ِب َو ُي ِقي ُمو َن ال َّص ٰلو َة‬

(Who believe in the Ghayb and perform the Salah) (2:3), and the following Ayat. Ibn Jarir stated that the
Ayah (2:2) includes all of these meanings that the scholars have mentioned, and this is the correct view.
Also, At-Tirmidhi and Ibn Majah narrated that `Atiyah As-Sa`di said that the Messenger of Allah ‫ ﷺ‬said,

«‫» َلا َي ْب ُل ُغ ا ْل َع ْب ُد َأ ْن َي ُكو َن ِم َن ا ْل ُم َّت ِقي َن َح َّتى َي َد َع َما َلا َب ْأ َس ِب ِه َح َذ ًرا ِم َّما ِب ِه َب ْأس‬

(The servant will not acquire the status of the Muttaqin until he abandons what is harmless out of fear
of falling into that which is harmful.) At-Tirmidhi then said "Hasan Gharib."

There are Two Types of Hidayah (Guidance)

Huda here means the faith that resides in the heart, and only Allah is able to create it in the heart of the
servants. Allah said,

‫ِإ َّن َك ل َا َت ْه ِدى َم ْن َأ ْح َب ْب َت‬

(Verily, you (O Muhammad ) guide not whom you like) (28:56),

‫َّل ْي َس َع َل ْي َك ُه َدا ُه ْم‬

(Not upon you (Muhammad ) is their guidance) (2:272),

‫َمن ُي ْض ِل ِل ال َّل ُه َفل َا َها ِد َي َل ُه‬

(Whomsoever Allah sends astray, none can guide him) (7:186), and,

‫َمن َي ْه ِد ال َّل ُه َف ُه َو ا ْل ُم ْه َت ِد َو َمن ُي ْض ِل ْل َف َلن َت ِج َد َل ُه َو ِل ًّيا ُّم ْر ِش ًدا‬

(He whom Allah guides, he is the rightly guided; but he whom He sends astray, for him you will find no
Wali (guiding friend) to lead him (to the right path)) (18:17).

Huda also means to explain the truth, give direction and lead to it. Allah, the Exalted, said,

‫َو ِإ َّن َك َل َت ْه ِدى ِإ َلى ِص َٰر ٍط ُّم ْس َت ِقي ٍم‬

(And verily, you (O Muhammad ) are indeed guiding (mankind) to the straight path (i.e. Allah's religion of
Islamic Monotheism)) (42: 52),

‫ِإ َّن َم آ َأن َت ُمن ِذ ٌر َو ِل ُك ِّل َق ْو ٍم َها ٍد‬

(You are only a warner, and to every people there is a guide) (13:7), and,

‫َو َأ َّما َث ُمو ُد َف َه َد ْي َٰنـ ُه ْم َفا ْس َت َح ُّبو ْا ا ْل َع َمى َع َلى ا ْل ُه َدى‬

(And as for Thamud, We showed and made clear to them the path of truth (Islamic Monotheism)
through Our Messenger (i.e. showed them the way of success), but they preferred blindness to
guidance) (41:17).

testifying to this meaning.

Also, Allah said,

‫َو َه َد ْي َٰنـ ُه ال َّن ْج َدي ِن‬

(And shown him the two ways (good and evil).) (90:10)

This is the view of the scholars who said that the two ways refer to the paths of righteousness and evil,
which is also the correct explanation. And Allah knows best.

Meaning of Taqwa

The root meaning of Taqwa is to avoid what one dislikes. It was reported that `Umar bin Al-Khattab
asked Ubayy bin Ka`b about Taqwa. Ubayy said, "Have you ever walked on a path that has thorns on it"
`Umar said, "Yes." Ubayy said, "What did you do then" He said, "I rolled up my sleeves and struggled."
Ubayy said, "That is Taqwa."

______________

Tafsir ibn kathir

‫َن َّز َل َع َل ْي َك ا ْل ِك ٰت َب ِبا ْل َح ِّق ُم َص ِّد ًقا ِّل َما َب ْي َن َي َد ْي ِه َو َأ ْن َز َل ال َّت ْو ٰرى َة َوا ْل ِإ ْن ِجي َل‬

"He has sent down upon you, [O Muhammad], the Book in truth, confirming what was before it. And He
revealed the Torah and the Gospel."

(QS. Al~imran 3: Verse 3)

‫ِم ْن َق ْب ُل ُه ًدى ِّلل َّنا ِس َو َأ ْن َز َل ا ْل ُف ْر َقا َن ۗ  ِإ َّن ا َّل ِذي َن َك َف ُروا ِب َئا ٰي ِت ال َّل ِه َل ُه ْم َع َذا ٌب َش ِدي ٌد ۗ  َوال َّل ُه َع ِزي ٌز ُذو ا ْن ِت َقا ٍم‬

"Before, as guidance for the people. And He revealed the Qur'an. Indeed, those who disbelieve in the
verses of Allah will have a severe punishment, and Allah is exalted in Might, the Owner of Retribution."

(QS. Aali Imraan 3: Verse 4)

Commentary

1. The heavenly Books of prophets, one after another, are complementary of the formers:

"... Conforming that which is before it..."

2. The confirmation of the former heavenly Books and prophets is a factor to the unity of theists and
one of the manners of inviting others as the verse says: "conforming".

3. The confirmation of the Qur'an upon the Torah and Evangel is a proof for the idea that the former
Books are heavenly, and, also, it is a great right upon the former divine religions. Otherwise, regarding
the superstitions that have been added to the Torah and the Evangel, their being heavenly would be
forgotten totally.

4. The means, instructions and the stages of training are different in the former Books due to their time
and conditions of revelations. Yet they are on the same line of development and unity towards the godly
goal.

5. The revelation of the Qur'an unto the Prophet (S) is an honour from Allah unto his personality when
He preceded the phrase 'to you' to the term 'the Book':

"He has sent down to you the Book with truth..."

6. The Qur'an is secured from any falsehood, and it is coinciding to reality and truth.

7. In spite of the fact that the Torah and the Evangel were separately and wholly revealed at once, the
Quran was sent down gradually during twenty three years. This is in the condition that it was revealed to
the heart of the Prophet (S) at once by the Grand Night.

8. Considering the qualities mentioned in the previous verse, the revelation of the heavenly Books is a
manifestation of the attributes of Allah.

The first part of this verse, added to the meaning of the previous verse, addresses the Prophet of Islam
(S) and totally says:
"He has sent down to you the Book with truth, confirming that which is before it, and He sent down the
Turah and the Evangel."

"Aforetime, a guidance for people ..."

Then, pointing to the Qur'an which separates right and wrong, it follows saying:

"... and He sent down the Furqan..."

Hence, after the verses of the Qur'an being sent down from the side of Allah and completing the
argument being done, together with testification of innate disposition and intellect to the truthfulness
of prophets in their invitations, there is not any other way against rejecters but punishment.

That is why, in this verse, next to the statement about the rightfulness of the Prophet (S) and the Qur'an,
it says:

"...Verily those who disbelieve in the Signs of Allah, for them there is a severe chastisement ..."

And, to make it clear that there is no doubt about the capability of Allah in accomplishing His threats, it
says:

"... and Allah is Mighty, the Lord of Retribution.

__________________

An enlightening commentary into the light of the holy Quran.

‫َأ َف َلا َي َت َد َّب ُرو َن ا ْل ُق ْر َءا َن ۚ  َو َل ْو َكا َن ِم ْن ِع ْن ِد َغ ْي ِر ال َّل ِه َل َو َج ُدوا ِفي ِه ا ْخ ِت ٰل ًفا َك ِثي ًرا‬

"Then do they not reflect upon the Qur'an? If it had been from [any] other than Allah, they would have
found within it much contradiction."

(QS. An Nisaa 4: Verse 82)

Commentary

"What, do they not ponder, do they [not] contemplate, the Qur’ān?, and the marvellous truths
contained in it. If it had been from other than God surely they would have found therein much
inconsistency, [much] contradiction in meaning and irregularity in arrangement."

(QS. An Nisaa 4: Verse 82)

______________

Tafsir Jalalayn

‫ج ِّمۡن ُه ِل ُتن ِذ َر ِب ِهۦ َو ِذ ۡك َر ٰى ِل ۡل ُم ۡؤ ِم ِني َن‬ٞ ‫ِك َٰت ٌب ُأن ِز َل ِإ َلۡي َك َف َلا َي ُكن ِفي َص ۡد ِر َك َح َر‬
"This is, a Book that is revealed to you — addressing the Prophet (s) — so let there be no inhibition, [no]
constraint, in your breast because of it, to convey it — for fear that you might be called a liar — that you
may warn thereby (li-tundhira, ‘that you may warn’, is semantically connected to unzila, ‘is revealed’,
that is to say, ‘[it is revealed] for [the purposes of] warning’) and as a reminder for those who believe, in
it."

(QS. Al A'raaf 7: Verse 2)

‫ا ٓلر ۚ  ِت ْل َك َءا ٰي ُت ا ْل ِك ٰت ِب ا ْل َح ِكي ِم‬

"Alif, Lam, Ra. These are the verses of the wise Book"

(QS. Yunus 10: Verse 1)

‫َأ َكا َن ِلل َّنا ِس َع َج ًبا َأ ْن َأ ْو َح ْي َن ٓا ِإ ٰلى َر ُج ٍل ِّم ْن ُه ْم َأ ْن َأ ْن ِذ ِر ال َّنا َس َو َب ِّش ِر ا َّل ِذي َن َءا َم ُن ٓوا َأ َّن َل ُه ْم َق َد َم ِص ْد ٍق ِع ْن َد َر ِّب ِه ْم ۗ  َقا َل ا ْل ٰك ِف ُرو َن ِإ َّن ٰه َذا َل ٰس ِح ٌر ُّم ِبي ٌن‬

"Have the people been amazed that We revealed [revelation] to a man from among them, [saying],
Warn mankind and give good tidings to those who believe that they will have a [firm] precedence of
honor with their Lord? [But] the disbelievers say, Indeed, this is an obvious magician."

(QS. Yunus 10: Verse 2)

‫ُأ‬
‫آلرۚ  ۚ  ِكٰتٌب ْحِكَمْت َءاٰيُتُهۥ ُثَّم ُفِّصَلْت ِمْن َّلُدْن َحِكيٍم َخ ِبيٍر‬

"Alif, Lam, Ra. [This is] a Book whose verses are perfected and then presented in detail from [one who
is] Wise and Acquainted."

(QS. Hud 11: Verse 1)

‫َّو َأ ِن ا ْس َت ْغ ِف ُروا َر َّب ُك ْم ُث َّم ُتو ُب ٓوا ِإ َل ْي ِه ُي َم ِّت ْع ُك ْم َّم ٰت ًعا َح َس ًنا ِإ ٰل ٓى َأ َج ٍل ُّم َس ًّمى َّو ُي ْؤ ِت ُك َّل ِذى َف ْض ٍل َف ْض َل ُهۥ ۖ  َو ِإ ْن َت َو َّل ْوا َف ِإ ِّن ٓى َأ َخا ُف َع َل ْي ُك ْم َع َذا َب َي ْو ٍم َك ِبي ٍر‬

"And [saying], Seek forgiveness of your Lord and repent to Him, [and] He will let you enjoy a good
provision for a specified term and give every doer of favor his favor. But if you turn away, then indeed, I
fear for you the punishment of a great Day."

(QS. Hud 11: Verse 3)

‫ا ٓلر ۚ  ِت ْل َك َءا ٰي ُت ا ْل ِك ٰت ِب ا ْل ُم ِبي ِن‬

"Alif, Lam, Ra. These are the verses of the clear Book."

(QS. yusif 12: Verse 1)

‫ِإ َّن ٓا َأ ْن َز ْل ٰن ُه ُق ْر ٰء ًنا َع َر ِب ًّيا َّل َع َّل ُك ْم َت ْع ِق ُلو َن‬

"Indeed, We have sent it down as an Arabic Qur'an that you might understand."

(QS. yusif 12: Verse 2)

‫َن ْح ُن َن ُق ُّص َع َل ْي َك َأ ْح َس َن ا ْل َق َص ِص ِب َم ٓا َأ ْو َح ْي َن ٓا ِإ َل ْي َك ٰه َذا ا ْل ُق ْر َءا َن َو ِإ ْن ُك ْن َت ِم ْن َق ْب ِل ِهۦ َل ِم َن ا ْل ٰغ ِف ِلي َن‬

"We relate to you, [O Muhammad], the best of stories in what We have revealed to you of this Qur'an
although you were, before it, among the unaware."

(QS. yusif 12: Verse 3)

‫ا ٓلاـ ّٓم ٰر ۚ  ِت ْل َك َءا ٰي ُت ا ْل ِك ٰت ِب ۗ  َوا َّل ِذ ٓى ُأ ْن ِز َل ِإ َل ْي َك ِم ْن َّر ِّب َك ا ْل َح ُّق َو ٰل ِك َّن َأ ْك َث َر ال َّنا ِس َلا ُي ْؤ ِم ُنو َن‬

"Alif, Lam, Meem, Ra. These are the verses of the Book; and what has been revealed to you from your
Lord is the truth, but most of the people do not believe."

(QS. Ar Ra'd 13: Verse 1)

‫ا ٓلر ۚ  ِك ٰت ٌب َأ ْن َزْل ٰن ُه ِإ َل ْي َك ِل ُت ْخ ِر َج ال َّنا َس ِم َن ال ُّظ ُل ٰم ِت ِإ َلى ال ُّنو ِر ِب ِإ ْذ ِن َر ِّب ِه ْم ِإ ٰلى ِص ٰر ِط ا ْل َع ِزي ِز ا ْل َح ِمي ِد‬

"Alif, Lam, Ra. [This is] a Book which We have revealed to you, [O Muhammad], that you might bring
mankind out of darknesses into the light by permission of their Lord - to the path of the Exalted in
Might, the Praiseworthy -

(QS. Ibrahim 14: Verse 1)

‫ال َّل ِه ا َّل ِذى َل ُهۥ َما ِفى ال َّس ٰم ٰو ِت َو َما ِفى ا ْل َأ ْر ِض ۗ  َو َو ْي ٌل ِّل ْل ٰك ِف ِري َن ِم ْن َع َذا ٍب َش ِدي ٍد‬

"Allah, to whom belongs whatever is in the heavens and whatever is on the earth. And woe to the
disbelievers from a severe punishment"

(QS. Ibrahim 14: Verse 2)

‫ا ٓلر ۚ  ِت ْل َك َءا ٰي ُت ا ْل ِك ٰت ِب َو ُق ْر َءا ٍن ُّم ِبي ٍن‬

"Alif, Lam, Ra. These are the verses of the Book and a clear Qur'an."

(QS. Al Hijr 15: Verse 1)

‫ِذ ْك ُر َر ْح َم ِت َر ِّب َك َع ْب َد ُهۥ َز َك ِر َّي ٓا‬

"[This is] a mention of the mercy of your Lord to His servant Zechariah"

(QS. Maryam 19: Verse 2)

‫َم ٓا َأ ْن َز ْل َنا َع َل ْي َك ا ْل ُق ْر َءا َن ِل َت ْش ٰق ٓى‬

"We have not sent down to you the Qur'an that you be distressed"

(QS. Taa Haa 20: Verse 2)

‫ِإ َّلا َت ْذ ِك َر ًة ِّل َم ْن َي ْخ ٰشى‬

"But only as a reminder for those who fear [Allah] -"

(QS. Taa Haa 20: Verse 3)

‫َت ْن ِزي ًلا ِّم َّم ْن َخ َل َق ا ْل َأ ْر َض َوال َّس ٰم ٰو ِت ا ْل ُع ٰلى‬

"A revelation from He who created the earth and highest heavens,"

(QS. Taa Haa 20: Verse 4)

‫ِت ْل َك َءا ٰي ُت ا ْل ِك ٰت ِب ا ْل ُم ِبي ِن‬

"These are the verses of the clear Book."

(QS. Ash Shu'araa 26: Verse 2)

‫َل َع َّل َك ٰب ِخ ٌع َّن ْف َس َك َأ َّلا َي ُكو ُنوا ُم ْؤ ِم ِني َن‬

"Perhaps, [O Muhammad], you would kill yourself with grief that they will not be believers."

(QS. Ash Shu'araa 26: Verse 3)

‫ط ٓس ۚ  ِت ْل َك َءا ٰي ُت ا ْل ُق ْر َءا ِن َو ِك َتا ٍب ُّم ِبي ٍن‬

"Ta, Seen. These are the verses of the Qur'an and a clear Book"

(QS. An Naml 27: Verse 1)

‫ُه ًدى َو ُب ْش ٰرى ِل ْل ُم ْؤ ِم ِني َن‬

"As guidance and good tidings for the believers"

(QS. An Naml 27: Verse 2)

‫ُه ًدى َو ُب ْش ٰرى ِل ْل ُم ْؤ ِم ِني َن‬

"As guidance and good tidings for the believers"

(QS. An Naml 27: Verse 2)

‫ِت ْل َك َءا ٰي ُت ا ْل ِك ٰت ِب ا ْل ُم ِبي ِن‬

"These are the verses of the clear Book."

(QS. Al Qasas 28: Verse 2)

‫َن ْت ُلوا َع َل ْي َك ِم ْن َّن َب ِإ ُمو ٰسى َو ِف ْر َع ْو َن ِبا ْل َح ِّق ِل َق ْو ٍم ُي ْؤ ِم ُنو َن‬

"We recite to you from the news of Moses and Pharaoh in truth for a people who believe."

(QS. Al 28: Verse 3)

‫ِت ْل َك َءا ٰي ُت ا ْل ِك ٰت ِب ا ْل َح ِكي ِم‬

"These are verses of the wise Book,"

(QS. Luqman 31: Verse 2)

‫ُه ًدى َو َر ْح َم ًة ِّل ْل ُم ْح ِس ِني َن‬

"As guidance and mercy for the doers of good"

(QS. Luqman 31: Verse 3)

‫ا َّل ِذي َن ُي ِقي ُمو َن ال َّص ٰلو َة َو ُي ْؤ ُتو َن ال َّز ٰكو َة َو ُه ْم ِبا ْل َأا ِخ َر ِة ُه ْم ُيو ِق ُنو َن‬

"Who establish prayer and give zakah, and they, of the Hereafter, are certain [in faith]."

(QS. Luqman 31: Verse 4)

‫ُأو ٰٓل ِئ َك َع ٰلى ُه ًدى ِّم ْن َّر ِّب ِه ْم ۖ  َو ُأو ٰٓل ِئ َك ُه ُم ا ْل ُم ْف ِل ُحو َن‬

"Those are on [right] guidance from their Lord, and it is those who are the successful."

(QS. Luqman 31: Verse 5)

‫َت ْن ِزي ُل ا ْل ِك ٰت ِب َلا َر ْي َب ِفي ِه ِم ْن َّر ِّب ا ْل ٰع َل ِمي َن‬

"[This is] the revelation of the Book about which there is no doubt from the Lord of the worlds."

(QS. As Sajda 32: Verse 2)

‫َأ ْم َي ُقو ُلو َن ا ْف َت ٰرى ُه ۚ  َب ْل ُه َو ا ْل َح ُّق ِم ْن َّر ِّب َك ِل ُت ْن ِذ َر َق ْو ًما َّم ٓا َأ ٰتى ُه ْم ِّم ْن َّن ِذي ٍر ِّم ْن َق ْب ِل َك َل َع َّل ُه ْم َي ْه َت ُدو َن‬

"Or do they say, He invented it? Rather, it is the truth from your Lord, [O Muhammad], that you may
warn a people to whom no warner has come before you [so] perhaps they will be guided."

(QS. As Sajda 32: Verse 3)

‫َوا ْل ُق ْر َءا ِن ا ْل َح ِكي ِم‬

"By the wise Qur'an."

(QS. Yaseen 36: Verse 2)

‫ٓص ۚ  َوا ْل ُق ْر َءا ِن ِذى ال ِّذ ْك ِر‬

"Sad. By the Qur'an containing reminder..."

(QS. Saad 38: Verse 1)

‫َت ْن ِزي ُل ا ْل ِك ٰت ِب ِم َن ال َّل ِه ا ْل َع ِزي ِز ا ْل َح ِكي ِم‬

"The revelation of the Qur'an is from Allah, the Exalted in Might, the Wise."
(QS. Az Zumar 39: Verse 1)

‫ِإ َّن ٓا َأ ْن َز ْل َن ٓا ِإ َل ْي َك ا ْل ِك ٰت َب ِبا ْل َح ِّق َفا ْع ُب ِد ال َّل َه ُم ْخ ِل ًصا َّل ُه ال ِّدي َن‬

"Indeed, We have sent down to you the Book, [O Muhammad], in truth. So worship Allah, [being]
sincere to Him in religion."

(QS. Az Zumar 39: Verse 2)

‫َت ْن ِزي ُل ا ْل ِك ٰت ِب ِم َن ال َّل ِه ا ْل َع ِزي ِز ا ْل َع ِلي ِم‬

"The revelation of the Book is from Allah, the Exalted in Might, the Knowing."

(QS. Ghafir 40: Verse 2)

‫َت ْن ِزي ٌل ِّم َن ال َّر ْح ٰم ِن ال َّر ِحي ِم‬

"[This is] a revelation from the Entirely Merciful, the Especially Merciful -"

(QS. Fussilat 41: Verse 2)

‫ِك ٰت ٌب ُف ِّص َل ْت َءا ٰي ُت ُهۥ ُق ْر َءا ًنا َع َر ِب ًّيا ِّل َق ْو ٍم َي ْع َل ُمو َن‬

"A Book whose verses have been detailed, an Arabic Qur'an for a people who know,"

(QS. Fussilat 41: Verse 3)

‫َك ٰذ ِل َك ُيو ِح ٓى ِإ َل ْي َك َو ِإ َلى ا َّل ِذي َن ِم ْن َق ْب ِل َك ال َّل ُه ا ْل َع ِزي ُز ا ْل َح ِكي ُم‬

"Thus has He revealed to you, [O Muhammad], and to those before you - Allah, the Exalted in Might, the
Wise."

(QS. Ash Shura 42: Verse 3)

‫َوا ْل ِك ٰت ِب ا ْل ُم ِبي ِن‬

"By the clear Book,"

(QS. Az Zukhruf 43: Verse 2)

‫ِإ َّنا َج َع ْل ٰن ُه ُق ْر ٰء ًنا َع َر ِب ًّيا َّل َع َّل ُك ْم َت ْع ِق ُلو َن‬

"Indeed, We have made it an Arabic Qur'an that you might understand."

(QS. Az Zukhruf 43: Verse 3)

‫َوا ْل ِك ٰت ِب ا ْل ُم ِبي ِن‬

"By the clear Book,"

(QS. Ad Dukhaan 44: Verse 2)

‫ِإ َّن ٓا َأ ْن َز ْل ٰن ُه ِفى َل ْي َل ٍة ُّم ٰب َر َك ٍة ۚ  ِإ َّنا ُك َّنا ُم ْن ِذ ِري َن‬

"Indeed, We sent it down during a blessed night. Indeed, We were to warn [mankind]."

(QS. Ad 44: Verse 3)

‫َت ْن ِزي ُل ا ْل ِك ٰت ِب ِم َن ال َّل ِه ا ْل َع ِزي ِز ا ْل َح ِكي ِم‬

"The revelation of the Book is from Allah, the Exalted in Might, the Wise."

(QS. Al Jaathiya 45: Verse 2)

‫َت ْن ِزي ُل ا ْل ِك ٰت ِب ِم َن ال َّل ِه ا ْل َع ِزي ِز ا ْل َح ِكي ِم‬

"The revelation of the Book is from Allah, the Exalted in Might, the Wise."

(QS. Al Ahqaf 46: Verse 2)

‫َما َخ َل ْق َنا ال َّس ٰم ٰو ِت َوا ْل َأ ْر َض َو َما َب ْي َن ُه َم ٓا ِإ َّلا ِبا ْل َح ِّق َو َأ َج ٍل ُّم َس ًّمى ۚ  َوا َّل ِذي َن َك َف ُروا َع َّم ٓا ُأ ْن ِذ ُروا ُم ْع ِر ُضو َن‬

"We did not create the heavens and earth and what is between them except in truth and [for] a
specified term. But those who disbelieve, from that of which they are warned, are turning away."

(QS. Al Ahqaf 46: Verse 3)

‫ٓق ۚ  َوا ْل ُق ْر َءا ِن ا ْل َم ِجي ِد‬

"Qaf. By the honored Qur'an..."

(QS. Qaaf 50: Verse 1)

‫َو ِك ٰت ٍب َّم ْس ُطو ٍر‬

"And [by] a Book inscribed"

(QS. At Tur 52: Verse 2)

‫َع َّل َم ا ْل ُق ْر َءا َن‬

"Taught the Qur'an,"

(QS. Ar Rahmaan 55: Verse 2)

‫ُق ْل ُأو ِح َى ِإ َل َّى َأ َّن ُه ا ْس َت َم َع َن َف ٌر ِّم َن ا ْل ِج ِّن َف َقا ُل ٓوا ِإ َّنا َس ِم ْع َنا ُق ْر َءا ًنا َع َج ًبا‬

"Say, [O Muhammad], It has been revealed to me that a group of the jinn listened and said, 'Indeed, we
have heard an amazing Qur'an."

(QS. Al Jinn 72: Verse 1)

‫َي ْه ِد ٓى ِإ َلى ال ُّر ْش ِد َف َئا َم َّنا ِب ِهۦ ۖ  َو َل ْن ُّن ْش ِر َك ِب َر ِّب َن ٓا َأ َح ًدا‬

"It guides to the right course, and we have believed in it. And we will never associate with our Lord
anyone."

(QS. Al Jinn 72: Verse 2)

‫َو َأ َّن ُهۥ َت ٰع ٰلى َج ُّد َر ِّب َنا َما ا َّت َخ َذ ٰص ِح َب ًة َوَلا َو َل ًدا‬

"And [it teaches] that exalted is the nobleness of our Lord; He has not taken a wife or a son"

(QS. Al Jinn 72: Verse 3)

‫َأ ْو ِز ْد َع َل ْي ِه َو َر ِّت ِل ا ْل ُق ْر َءا َن َت ْر ِتي ًلا‬

"Or add to it, and recite the Qur'an with measured recitation."

(QS. Al Muzzammil 73: Verse 4)

‫َوال َّس َم ٓا ِء َذا ِت ال َّر ْج ِع‬

"By the sky which returns [rain]"

(QS. At Taariq 86: Verse 11)

‫َوا ْل َأ ْر ِض َذا ِت ال َّص ْد ِع‬

"And [by] the earth which cracks open,"

(QS. At Taariq 86: Verse 12)

‫ِإ َّن ُهۥ َل َق ْو ٌل َف ْص ٌل‬

"Indeed, the Qur'an is a decisive statement,"

(QS. At Taariq 86: Verse 13)

‫َو َما ُه َو ِبا ْل َه ْز ِل‬

"And it is not amusement."

(QS. At 86: Verse 14)

‫ُقل َّل ْو َكا َن ا ْل َب ْح ُر ِم َدا ًدا ِّل َك ِل ٰم ِت َر ِّبى َل َن ِف َد ا ْل َب ْح ُر َق ْب َل َأ ْن َت ْن َف َد َك ِل ٰم ُت َر ِّبى َو َل ْو ِج ْئ َنا ِب ِم ْث ِل ِهۦ َم َد ًدا‬

"Say, If the sea were ink for [writing] the words of my Lord, the sea would be exhausted before the
words of my Lord were exhausted, even if We brought the like of it as a supplement."

(QS. Al Kahf 18: Verse 109).

Commentary
"Say: ‘If the sea, in other words, [if] its waters, were ink (midād is what one writes with) for the Words of
my Lord, [Words] that testify to His laws and His marvels, such that these [Words] are written with it,
the sea would be spent, in recording them, before the Words of my Lord were spent (read as tanfada or
yanfada) even though We brought the like of it, namely, [the like of] the sea, as replenishment’, in order
to add it to the other [sea], it would [also] be spent, while they [God’s Words] would not be exhausted
(madadan, ‘as replenishment’, in the accusative because it is a specification)."

(QS. Al Kahf 18: Verse 109)

THE INTRODUCTION OF THE BASIC CONCEPT

‫بسم الله الرحمن الرحيم‬

In the Name of Allah, the All-Merciful, the Most Merciful!

It's vehemently and gregariously the tremendous knowledge indulged in this prestigious statement of
‫بسم الله الرحمن الرحيم‬. It actually comprises of any kind of knowledge and the whole explanation on
the concept of God, power and the whole creations.
It is related from the book of yunabi'ul mawadda from the writer of Durrul Munazzam, that: The whole
reasons against the revelation of the Devine books is in the holy Qur'an, and whatever knowledge is in
the holy Qur'an, it had been diluted in suratul Fatiha, and whatever is in Suratul Fatiha, it is there in ‫بسم‬
‫الله الرحمن الرحيم‬. And whatever is in this statement of BASMALAH it is really in the letter (‫)ب‬, of the
statement.

It is also narrated from Abdullahi bin Abbas that, One-day we went out with Imam Ali bin Abu Talib (may
Allah be pleased with him) and he asked me to read from the book of Allah, so I read ‫بسم الله الرحمن‬
‫الرحيم‬. And he asked me about the first letter (‫ )ب‬of the statement but I remained stoned, so imam
start explaining the letter until dawn.

__________________________

See Yanabi'ul mawadda of Sheikh Qanduzi, volume 1, page 64.

Madalibul Al-su'al of Imam Shafi'i, volume 1, Ummul-Qura publication.

Spiritual vindication towards the theological therapeutics of Islamic Philosophy by Aliyu Salisu
Muhammad DAN~SIDI.

According to the famous tafsir work of Imam Qurtubi, he explained the concept of BASMALAH as qouted
below:

Scholars say that "In the Name of Allah, the All-Merciful, Most Merciful" is an oath from our Lord
beginning of every sura. By it, He swears to His slaves, "What I have laid down for you, My slaves, in this
sura is true. I will fulfill for you all that I guarantee in this sura of My promise, kindness and gentleness."
"In the Name of Allah, the All-Merciful, Most Merciful" is part of what Allah revealed in His Book, and
this is special for this Community after Sulayman. Some scholars say that "In the Name of Allah, the All-
Merciful, Most Merciful" contains all the Shari'a because it indicates the Essence and the Attributes.

Sa'ld ibn Abi Sakina said that he heard that 'Ali ibn Abi Talib looked at a man who had written, "In the
Name of Allah, the All-Merciful, Most Merciful," and told him, "Do it well. If a man does it well, he will be
forgiven." Sa'id said, "I heard that a man looked at a parchment on which was written 'In the Name of
Allah, the All-Merciful, Most Merciful,' and kissed it and placed it on his eyes and was forgiven on
account of that." There is a similar story from Bishr Al-Ilafi. When he picked up a rag on which was "the
Name of Allah" and perfumed it, his own name became honoured.

Al-Qushayri mentioned that:

An-Nasa'i reports from Abu'l-Malih about a man who rode behind the Messenger of Allah that he
mentioned that, the Messenger of Allah, may Allah bless him and grant him peace, said, "When your
animal stumbles with you, do not say, 'Shaytan has made it stumble!' because that puffs him up until he
becomes like a house and says with strength, 'I have done it.' Rather say,
'In the Name of Allah, the All-Merciful, Most Merciful' and Shaytan will become small until he is like a
fly."'

'Ali ibn Al-hasan said about the words of the Almighty, "When you mention your Lord alone in the
Qur'an, they turn their backs in flight" (17:46), "That refers to when you say, 'In the Name of Allah, the
All-Merciful, Most Merciful."' It is reported that 'Abdullah ibn Mas'ud said, "Whoever wants Allah to
rescue him from the nineteen Zabiiniyya should recite, 'In the Name of Allah, the All-Merciful, Most
Merciful' so that Allah will make each let ter a shield for him against one of them. The basmala contains
nineteen letters according to the number of the angels of the Fire which He revealed at the who Allah
says are also nineteen. (74:30) They say in all that they do, 'In the Name of Allah, the All-Merciful, Most
Merciful.' That is their strength and they take their strength from the name of Allah."

Ibn 'Atiyya said, "The like of this is what they say about the Night of Power being the 27th night, taking
note of the position of the word "hiya" in the words of the sura, Al-Qadr (97:1-5). [It is the twenty-
seventh word in the sura.] That is like what they say about the number of angels who hastened to report
the words of the one who said, 'My Lord, praise is Yours, abundant, excellent and blessed,' [after rising
from ruku' when the Prophet said,

"Allah hears the one who praises Him."] It is about thirty letters. That is why the Prophet, may Allah
bless him and grant him peace, said, 'I saw about thirty angels racing to see which would be the first to
write it down."' Ibn 'Atiyya said, "This is an elegant tafsir, but not a firm tafsir."

Ash-Sha'bi and al-A'mash report that the Messenger of Allah, may Allah bless him and grant him peace,
wrote, "In Your Name, 0 Allah" until he was commanded to write "In the Name of Allah," and then he
wrote that. When it was revealed, "Say: 'Call on Allah or call on the All-Merciful" (17:110), he wrote, "In
the Name of Allah, the All-Merciful." When there was revealed, "It is from Sulayman and says, 'In the
Name of Allah, the All-Merciful, Most Merciful'" (27:30), he wrote that. In Abil Dawiid, ash-Sha'bi, Abii
Malik, Qatada and Thabit ibn 'Umara said that the Prophet did not write "In the Name of Allah, the All-
Merciful, Most Merciful" until Surat an-Naml (27) was revealed.

It is reported that Ja'far assadiq said that the basmala is the crown of the suras. This indicates that it is
not an ayat of the Fatiha or other suras. People disagree about this and have three positions regarding
it.

• It is not an ayat of the Fatihata or any other sura. This is the position of Malik.

• It is an ayat of every sura, and this is the position of 'Abdullah ibn al-Mubarak.

• Ash-Shafi 'i said that it is an ayat of the Fatiha and what he says about the other suras varies.
Sometimes he says that it is an ayat of every sura and sometimes that it is only one of the Fatiha. There
is no disagreement that it is an ayat of the Qur'an inside Surat an-Naml.

Ash-Shafi 'i' s evidence is what ad-Daraqutni related from Abii Hurayra that the Prophet, may Allah bless
him and grant him peace, said, "When you read 'Praise belongs to Allah, the Lord of all the worlds, ' then
recite 'In the Name of Allah, the All-Merciful, Most Merciful'. It is the Mother of the Qur'an, the Mother
of the Book, and the Seven Mathani. 'In the Name of Allah, the All- Merciful, Most Merciful' is one of its
ayats."

The evidence of lbn al-Mubarak and one of the positions of ash-Shafi 'i is what Muslim reports from
Anas: "One day while the Messenger of Allah, may Allah bless him and grant him peace, was among us
he nodded off and then he raised his head smiling.

We asked, 'What has made you smile, Messenger of Allah?' He replied, 'A sura was just revealed to me.
It is: "In the Name of Allah, the All-Merciful, Most Merciful. Truly We have given you the Great
Abundance. So pray to your Lord and sacrifice. It is the one who hates you who is cut off without an
heir.'" (108)"

The sound position is that of Malik because the Qur'an is not established by single reports, but by way of
definitive multiple transmission about which there is no disagreement. lbn al-'Arab1i said, "It is enough
for you that there is no disagreement between people about the Qur'an. There is no disagreement
about the Qur'an." Sound reports which cannot be attacked indicated that the basmala is not an ayat of
al-Fatiha or any other sura except for Surat an-Naml. Muslim reported that Abii Hurayra said, "I heard
the Messenger of Allah, may Allah bless him and grant him peace, say, 'Allah says, "I have divided the
prayer into two halves between Me and My slave, and My slave will have what he asks for. When My
slave says, "Praise be to Allah, the Lord of all the worlds, "Allah says, 'My slave has praised Me." He says,
"the All-Merciful, the Most Merciful," and the Lord says, "My slave has lauded Me." My slave says, "the
King of the Day of Judgement," and Allah says, "My slave has magnified Me (or entrusted to Me)." The
slave says, "You alone we worship. You alone we ask for help, " and Allah says, "This is between Me and
My slave and My slave will have what He asks for." The slave says, "Guide us on the Straight Path, the
Path of those whom You have blessed, not of those with anger on them, nor of the misguided, " and
Allah says, "Those are for My slave and My slave will have what He asks for.""'

Allah said, "I have divided the prayer", meaning the Fatiha, and he called it "prayer" because the prayer
is not valid except with it. So He designated the first three ayats for Himself, singling them out for
Himself, and the Muslims do not disagree about that. Then he made the fourth between Him and His
slave because it contains the abasement of the slave and seeking help from Him. That contains esteem
for Allah. Then three ayats conclude the seven. They are three, because He uses the plural, not the dual
in "Those are for My slave," So "those You have blessed" is an ayat. Ibn Bukayr reported that Malik said
that "those You have blessed" is an ayat.

This is confirmed by the division and by what the Prophet said to Ubayy when he asked the Prophet,
"How do you recite when you begin the prayer?" He replied, "I recited, 'Praise be to Allah, the Lord of all
the worlds' to the end." The basmala was not part of it. That was the position of the people of Madina,
the people of Syria and the people of Basra. Most reciters counted "those You have blessed" as an ayat.
This is also related from Abii Nadhra from Abii Hurayra who said that the sixth ayat is "those You have
blessed". The people of Kufa count the basmala as part of it and do not count "those You have blessed".
If it is said that it is confirmed in the copies of the Qur'an that the basmala is written and transmitted as
it is transmitted in an-Naml and that this is multiple transmission, we reply that that is sound, but is it
because it is Qur'an, or is it a divider between suras as is related from the Companions, "We did not
know the end of the sura until 'In the Name of Allah, the All-Merciful, Most Merciful' was revealed."
(Abii Dawiid) Or it may be for the bless-ing, in the same way that the Community agrees to write it at the
beginning of books and letters. All of that is possible. Al-Jurayri said, "Al-Hasan was asked about 'In the
Name of Allah, the All- Merciful, Most Merciful' and said 'At the beginning of letters."' He also said, "'In
the Name of Allah, the All-Merciful, Most Merciful' was not revealed in any of the Qur'an except an-
Naml."

The criterion is that the Qur'an is not established by logic and deduction. It is established by definitive
multiple transmission. So the statement of ash-Shafi 'i about the basmala being at the beginning of each
sura is unsound because the basmala is not an ayat of each sura. Praise belongs to Allah.

It is reported that a group related that the basmalas are part of the Qur'an. Ad-Daraqu~nl dealt with all
of that. We do not deny the transmission of that and we have indicated it, but we have firm reports
which counter it which are related by reliable imams and fuqahii '. In Sahih Muslim, 'A'isha is reported as
saying, "The Messenger of Allah, may Allah bless him and grant him peace, used to begin the prayer with
the takbir and the recitation of 'Praise be to Allah, the Lord of all the worlds."' Muslim also reported that
Anas ibn Malik said, "I prayed behind the Prophet, Abii Bakr and 'Umar, and they began with 'Praise be
to Allah.'

They did not mention 'In the Name of Allah, the All-Merciful, Most Merciful', either at the beginning or
the end of recitation." So our school prefers that, and it is logical. That is because the Mosque of the
Prophet in Madina passed through many years from the time of the Messenger of Allah, until the time of
Malik and during all that time no one recited, "In the Name of Allah, the All-Merciful, Most Merciful,"
following the Sunna. This refutes the ahadith you cite. Our people, however, prefer to recite it in the
voluntary prayers, and there are traditions (athar) about reciting it or an allowance for doing that. Malik
said, "There is no harm in reciting it in the nafila or simply reading the Qur'an."

A group of the school of Malik and his people said that it is not part of the Fatiha or any other sura, and
it is not read by the one who prays the obligatory or any other prayer, either silently or aloud. It is
permitted to recite it in nafila prayers. This is well-known in his school and with his people. There is
another transmission that it is recited at the beginning of the sura in nafila prayers but not at the
beginning of the Fatiha. It is related that Ibn Nafi' began his recitation with it in the obligatory and nafila
prayers and did not ever omit it. Some of the people of Madina say that there must be "In the Name of
Allah, the All-Merciful, Most Merciful," among them Ibn 'Umar and Ibn Shihab. Ash-Shafi 'i, Ahmad,
Ishaq, Abii Thawr and Abu 'Ubayd said that. That indicates that it is a matter of ijtihad and not definitive,
as some ignorant individuals claim.

A group of scholars believe that it is recited silently with the Fatiha They include Abii Hanifa and ath-
Thawri. That is related

from 'Umar, 'Ali, Ibn Mas'ud, 'Ammar and Ibn az-Zubayr. It is also the view of al-Hakam and Al-Hammad,
and it is stated by Ahmad ibn hanbal and Abu 'Ubayd. Something similar to that is related from al-Awza'i
The evidence is the report from Anas ibn Malik:
"The Messenger of Allah, may Allah bless him and grant him peace, led us in the prayer and we did not
hear him recite, 'In the Name of Allah, the All-Merciful, Most Merciful.'" This is a good position, and the
traditions (athar) reported from Anas agree on it and remove the disagreement about the recitation of
the basmala. It is related that Sa'Id ibn Jubayr said, "The idolaters used to come to the mosque. When
the Messenger of Allah, may Allah bless him and grant him peace, recited, 'In the Name of Allah, the All-
Merciful, Most Merciful,' they said, 'This Muhammad mentioned the Rahman of Yamama,' meaning
Musaylama. So he was commanded to recite it silently and it was revealed, 'Do not be too loud in your
prayer or too quiet in it.' (17:110)" At-Tirmidhl al-Hakim said, "That has lasted until now, even if the
cause no longer exists, as running remains in tawaf even if the cause no longer exists and silence in the
day prayers even if the cause no longer exists."

The Community agree that it is permitted to write it at the beginning of every book of knowledge and
letter. There is disagreement about books of poetry and whether or not it should be written in them.

The Shari'a recommends mentioning the basmala at the begin- ning of every action, like eating, drinking,
slaughtering, sex, purification, embarking on a ship and the like. Allah says, "Eat that over which the
name of Allah has been mentioned" ( 6: 118) and "He said, 'Embark in it. In the name of Allah be its
voyage and its landing!'" (11 :41) The Prophet, may Allah bless him and grant him peace, said, "Lock your
door and mention the Name of Allah.

Put out your lamp and mention the Name of Allah. Cover your vessel and mention the Name of Allah.
Tie up your water skin and mention the Name of Allah." He said, "If anyone of you wants to go to his
wife, he should say, 'In the Name of Allah. 0 Allah, keep Shaytan away from us and keep Shaytan away
from what You provide us with.' If a child is decreed for them, Shaytan will not harm him at all." He told
'Umar ibn Abi Salama, "Boy, say the name of Allah Almighty and eat with your right hand and eat what is
in front of you."

When 'Uthman ibn Abi'l-'As complained to him of a pain he had in his body since he had become
Muslim, the Messenger of Allah said to him, "Place your hand on that part of your body which pains you
and say 'In the Name of Allah' three times and then say seven times, 'I seek refuge in the might and
power of Allah from the evil of what I feel and am on my guard against.'"

Ibn Majah and at-Tirmidhi report that the Prophet said, "The veil between the jinn and the private parts
of the sons of Adam when he enters the privy is to say, 'In the Name of Allah."' Ad-Daraqutni reported
that 'A' isha said, "When the Prophet touched his wur)u' vessel, he said the Name of Allah and then
poured the water on his hands."

Our scholars say that this refutes the Qadarites and others who say that our actions are decreed for us.
The argument against them in that is that Allah commanded us to begin every action with the basmala.

The meaning of "In the Name of Allah" is "by Allah", and the meaning of "by Allah" is by His creation and
determination to reach what is reached. Some say that "In the Name of Allah" means "I begin with the
help, success and blessing of Allah." This is Allah teaching His slaves to mention His Name at the
beginning of actions.
There is disagreement about the derivation of ism (Name), with two basic positions. The Basrans say
that it is derived from sumu, which means height and elevation. It is said that ism means that the person
is in an elevated place. It is said that the name lifts the named from others. It is said that the name is
called that because it is higher than the other parts of speech by its strength. The noun (ism) is stronger
by agreement because it is the root. These are three statements.

The Kufans say that it is derived from sima, which means a sign, because the Name is a sign of the One
to whom it is given. So the root of ism is wasam. The first is sounder because of the form of the
diminutive and the form of plural which is asma'.

Another disagreement indicates the soundness of that, and it is the next point.

If it is true that ism is derived from height, Allah was described by it before creation existed, after it
existed and will be when it is annihilated, and creatures have no effect on the Names or Attributes. This
is the position of the people of the Sunna. Those who say that it is derived from sima, say that before
time Allah was without name or attribute. When He created creatures, they gave Him Names and
Attributes. When He annihilates them, He will again have no name or attribute. This is the position of
the Mu 'tazilites, and it is contrary to that on which the Community agree. It is worse than their error
when they say, "His Word is created." Exalted above that is Allah! It is according to this that there is a
disagreement about the Name and Named.

The people of truth believe, as Qadi Abil Bakr ibn at-Tayyib al-Baqillani mentioned, that the name is the
thing named and Ibn Farak is content with that. It is the position of Abii 'Abida and Sibuwayh. If
someone says, "Allah is Knowing" his words indicate the Essence which is described as being knowing.
So the Name is Knowing and it is what is Named. It is the same when someone says, "Allah is the
Creator." The Creator is the Lord and it is the Name itself. So their view is that the Name is the Named
itself with no distinction.

Ibn Hassar said, "Those among the innovators who deny the attributes claim that namings have no
meaning except the Essence. That is why they say that the Name is not the Named. Whoever affirms the
Attributes, affirms that the Names have meanings which are the qualities of the Essence. They are not
expressions, but they are Names in their view. More of this will come in al-Baqara and al-A 'raf Allah is
the greatest and most comprehensive of all the Names, so that one scholar said that it is the Greatest
Name of Allah and no one else has it. That is why it has no dual nor plural. That is one of two
interpretations of the words of the Almighty, "Do you know of any other with His Name?" ( 19:65), in
other words anyone named with His Name which is Allah. Allah is the Name of the True Existent who
has all the Divine Attributes and is described as Lord and alone possesses real existence. There is no god
but Him. Glory be to Him! It is said that it means the One who should be worshipped. It is said that it
means the One whose existence is necessary who always was and always will be. The meaning is the
same.

There is disagreement about whether this Name is derived or is a unique designation for the Divine
Essence. Many of the people of knowledge believe the first but then disagree on its actual derivation
and root. Sibuwayh reported from al-Khalil that its root is ilah, on the measure offi'al. The alif and lam
replace the hamza.

Sibuwayh said that it is like an-nas (people) whose root is anas. It is said that its root is lah and the alif
and lam are added to exalt it. This is what Sibuwayh preferred. Al-Kisa'i and al-Farra' said that
"bismillah" is made up of bismi - al-ilah and elision has occurred and the first lam assimilated into the
second and so becomes a double lam.

It is said that the name Allah is derived from walaha, to be bewildered. Walah means loss of intellect,
and someone who is walih is bewildered. Allah bewilders minds when they think on the realities of His
attributes and reflect on gnosis of Him. So the basis of ilah is walah and the hamza is changed from the
waw. That is also reported from al-Khalil. It is related that Al-dahhaq said, "He is called Allah because
creatures devote (ta'alah) themselves to Him in their needs and make supplication to Him in times of
hardship. It is related that al-Khali:l ibn Al).mad said something similar. It is also said that it is derived
from elevation and that the Arabs used to use "laha" for something elevated and they used the verb for
sunrise.

It is said that the name Allah is derived from the word ilah (god), which means an object of worship so
that the meaning of "Allah" is the Object of Worship. So what the unifiers say, "There is no god but
Allah" means "there is no object of worship other than Allah." Here "illa" means "other", not "except."
Some claim that the root is al-hii' which alludes to the third person. That is since they affirm Him as
existing in their natural intellects and indicate Him with the letter of allusion. Then the lam of possession
is added to it since they know that He is the Creator and Master of things, and laha then is added for
magnification.

The second position is taken by a group of scholars, including ash-Shafi'I, Abu'l-Ma'all, al-Khattabi, al-
Ghazali, al-Mufaddal and others, and is related from al-Khalll and Sibuwayh. It is that the alif and lam are
a necessary part of it and cannot be elided from it. Al-Khattabi said that the evidence that the alif and
lam are an intrinsic part of the structure of this name and not added for definition is that it is included in
the vocative, as "Ya Allah!" The vocative is not combined with the definite article alif-lam. One does not
say, "Ya Ar-Rahman"

Allah knows best.

There is also disagreement about the derivation of Ar-Rahman Some of them said that it has no
derivation because it is one of the names particular to Him and if it had been derived from mercy
(rahma), it would be connected to the one shown mercy and it would be possible to say, "Allah is
Rahman to His slaves" as one does with rahim. If it had been derived from rahma, the Arabs would not
have denied it when they heard it because they did not deny the mercy of their Lord. Allah says, "When
they are told to prostrate to the All-Merciful, they say, 'And what is the All-Merciful?'" (25:60)

At al-Hudaybiyya, when 'All wrote at the command of the Prophet "In the Name of Allah, the All-
Merciful, Most Merciful," Suhayl ibn 'Amr said, "As for 'In the Name of Allah, the All- Merciful, Most
Merciful,' we do not know 'In the Name of Allah, the All-Merciful, Most Merciful'! Rather write 'In Your
Name, 0 Allah."' Ibn al-'Arab! says that this indicates that they did not know the attribute rather than the
One Described. Evidence is found in the fact that they said, "What is the Rahman ?" not "Who is the
Rahman?" Ibn al-Amr said, "It is as if he [the one who said this] had not recited the other ayat, 'Yet they
still reject the All-Merciful.' (13:30)" One group believe that it is derived from rahma, and is intensive,
meaning the One who possesses mercy such as no one else has. It has no plural or dual whereas rahim
can be dual or plural.

Ibn al-Hassar said that part of what indicates the derivation is what at-Tirmidhi transmitted as sound
from 'Abdu'r-Rahman ibn 'Awf. He heard the Messenger of Allah say, "Allah Almighty says, 'I am the All-
Merciful. I created kinship (rahim) and split it as a name from My Name. Whoever maintains ties, I
maintain ties with him. Whoever severs it, I sever him."' This is a text for its derivation and so there is no
point in contention about it. The denial of the Arabs was simply due to their ignorance of Allah and what
is mandatory for Him.

Al-Anbari mentions that al-Mubarrad stated that ar-Rahman is a Hebrew name. Ahmad ibn Yahaya also
said that. This view is unwarranted. Abii'l-'Abbas says that the attribute is for praise. Qurtub says that it
is possible to combine the two for stress. There is disagreement about whether the two names Rahman
and Rahim have one meaning or two meanings. It is said, that they mean the same, as do nadman and
nadim. It is said that Rahman is a special name with general action and Rahim is a general name with a
particular action. This is the position of the majority.

Abii 'Ali al-Farisi said that Rahman is a general name for all types of mercy for which Allah is singled out.
Rahim can be used for how He is towards the believers, as He says, "He is merciful to the believers."
(33:43). AI-'Arazami says that Rahman is merciful to all His creatures with rain, physical and general
blessings, and Rahman is merciful to the believers in guiding them and being kind to them. Ibn al-
Mubarak said that when the Rahaman is asked He gives and when the Rahim is not asked, He is angry.
Ibn 'Abbas said that they are two fine (raqiq) names, and one is finer than the other, meaning that it has
more mercy.

Al-Khattabi said, "This is problematic because fineness has no place in any of the attributes of Allah." Al-
Husayn ibn al-Fadl al-Bajali said, "This is an error on the part of the transmitter because fineness (riqqa)
is not part of the attributes of Allah at all. Rather, they are two compassionate (rafiq) names, one more
compassionate than the other. Compassion is one of the Attributes of Allah Almighty. The Prophet, may
Allah bless him and grant him peace, said, 'Allah is Compassionate. He loves compassion and gives for
compassion what he does not for harshness."'

Most scholars agree that the name ar-Rahman is used only for Allah Almighty and it is not permitted to
call anyone else by it. Do you not see that He says, "Say: 'Call on Allah or call on the All-Merciful'" (17: 11
0)? So it is equal to the Name in which none but Him share. He says, "Ask those We sent before you as
Our Messengers: Have We ever designated any gods to be worshipped besides the All-Merciful?"
(43:445) So they reported that the Rahman deserved worship. Musaylima the Liar' - may Allah curse him
-was outrageous and called himself "the Rahman of Yamama" and so was called "the Liar". Ar-Rahim is
general and can be used in respect of creatures. As ar-Rahman is universal as we said, ar-Rahim provides
harmony and balance in the revelation. Al-Mahdawi stated that. It is said that the meaning of ar-Rahim
is: "It is by the Ra/:t!m that you reach to the Rahman." So ar-Rahim is the attribute of Muhammad, may
Allah bless him and grant him pea e, and Allah described him with that. He says, "compassionate,
merciful" when describing him.

_______________

1. Musaylima the Liar: a false Prophet of the Banii I:Ianifa in Yamana who was one of the leaders of the
Ridda.

So it is as if the meaning of saying, "In the Name of Allah, the All-Merciful, the Most Merciful" is "It is by
Muhammad, may Allah bless him and grant him peace, that you will reach Me," in other words by
following him and what he has brought, you will reach My reward, honour and the vision of My face."
Allah knows best.

It is related that 'Ali: ibn Abi 'falib said, "Bisimillah is healing from every illness and protection against
every disease. Ar-Rahman is a help for everyone who believes in Him. It is a name not used for anyone
else. Ar-Rahim is for those who repent, believe and perform righteous actions."

Some of them explained the meaning according to the letters. It is related that 'Uthman ibn 'Aflan asked
the Messenger of Allah, may Allah bless him and grant him peace, about the interpretation of "In the
Name of Allah, the All-Merciful, Most Merciful." He said, "The ba ' is the trial ( bala) of Allah, His relief,
brilliance and radiance (baha'). The sin is the splendour (sana') of Allah. The mim is the kingdom (mulk)
of Allah. As for Allah, there is no god but Him. The Rahman is kind to both the pious and impious of His
creatures. The Rahim is kind only to the believers." It is reported that Ka'b al-Al;lbar said, "The ba' is His
radiance (baha'), the sin is His splendour (sana'), and there is nothing higher than it. The mim is His
kingdom, and He has power over all things and nothing is hard for Him." It is said that every letter is the
opening of one of His Names. The ba' is the key to His Name Basirr (All-Seeing). The sin is the key to His
Name, Sami' (All-Hearing). The mim is the key to His Name, Malik (Master). The alifis the key to His
Name, Allah. The La is the key to His Name, Latif (Kind). The ha' is the key to His Name, Hadi (Guide). The
ra' is the key to His Name, Raziq (Provider). The ha' is the key to His Name, Halim (Forbearing).

The nun is the key to His Name, Nur (Light). The meaning of all this is supplication of Allah at the
beginning of everything. There is disagreement how 'ar-Rahim' is connected in recitation to 'al-
hamdulillah '. Umm Salama related that the Prophet recited 'ar-Rahim' with a sukun on the mim,
stopping there and then beginning with a fresh alif. Some of the Kufans recited it in that way. Most
people recite, 'ar-Rahimi'l-hamdu', with a kasra on the mim and connecting it to the alif in al-hamd. Al-
Kisa'I reported that some Arabs read it 'ar-Rahima'l-hamdu', with fatha on the mim and connected to
the alif, as if the mim was in fact silent, but with an elision into the alif. lbn 'Atiyya said, "This recitation is
not reported from anyone I know."

____________________
Tafsir Qurtubi by Abu-Abdullah Muhammad bin Ahmad ibn Abubakar ibn Farh, Al-Ansari Al-Khazraji Al-
Andalusi, Al-Qurtubi. Volume 1 page 78-92, Translated by Aisha Bewley.

According to one of my famous books "Spiritual Vindication towards the theological therapeutics of
Islamic philosophy" I have qouted concerning the meaning of ‫بسم الله الرحمن الرحيم‬. as fallows:

"Meaning" In the name of Allah the most beneficial the most merciful.

This above mentioned fantabulous statement can be segregated into different parts for the benefit of
one another as I have mentioned below:

The Arabic letter "B" (‫ )ب‬is brilliance and goodness of Allah.

The Arabic letter "C" (‫ )س‬is for effulgence and loftiness of Almighty Allah.

The Arabic letter "M" (‫ )م‬is for (MULK) that's kingdom of Almighty Allah. Allah is diety of everything.

The Arabic word "ARRAHMAN" is a common trait: that's the kind one and the sustainer of all creatures
in the universe.

The Arabic word "ARRAHEEM" is a special quality that is reserved only for the believers.

__________________

Bihar Anwar vol 89 page 228, Tauheed page 230, Al-Muhassin vol 1 page 238, ma'anul AKHBAR page 3,
Alkafi vol 1, page 59, Tafsir Ayyashi vol 1 page 22, Tafsir Qummi page 112, vol 1. Attabrirurruhi lil ilajatul
lahutiyyat lil falsafatul islamiyya, volume 1, page 22 by Aliyu Salisu Muhammad Dan~sidi.

So the BASMALAH here compresis of seven different parts to which I have wrote this book upon, as an
exegesis of this Noble statement. These parts are [(‫)ب‬, for brilliance and goodness of Allah, and (‫)س‬, for
effulgence and loftiness of the Almighty Allah, (‫)م‬, is for Kingdom of the Almighty, while the Word (
‫)الرحمن‬, is a common trait, that's the kind one and the sustainer of All creations, and finally (‫)الرحيم‬,
which is a special quality that is reserved only for the believers].

Light is a powerful force of nature, and has affected our lives profoundly throughout the ages.
Scientifically, light is the visible form of energy, which is produced by electro-magnetic radiation. Light is
a wave-like form of energy that is composed of photons, and is responsible for a range of phenomena,
including the behaviour of atoms, the chemical reaction of particles, and the reflection of colour.
Philosophically, light is a potent symbol in many human cultures, including those of Ancient Greece,
where it was associated with the divine.

On a scientific level, light is a wave-like form of energy that is produced in different forms, including
ultraviolet, infrared, and visible forms. It is a form of electromagnetic radiation that is composed of
particles known as photons, which travel in straight lines at the speed of light. Light provides the
photons which are essential for the production of energy, and is responsible for numerous behaviours,
from the reflection of colour to the reaction of chemicals.
Light's impact is not only seen in its scientific application. Throughout history, many cultures have
attached spiritual meaning to light, and it has been associated with divine power. In Ancient Greek
mythology, for example, light was believed to be associated with the gods, and it was seen to be the
source of all knowledge – the proverbial ‘light of wisdom’. Light was also used throughout the Middle
Ages as a symbol of virtue and purity, and has remained a powerful metaphor for truth and
understanding.

Light’s influence has been felt in many areas of human life, from science to religion. On a scientific level,
light is seen as an important source of energy, providing photons which enable the production of power
and the behaviour of atoms. Philosophically, light has come to represent knowledge, understanding, and
truth, and has been associated with the divine since the dawn of time. All in all, light is an incredibly
powerful force, and its influence on our lives continues to be felt.

Praise be to Allah Who lies inside all hidden things and towards Whom all open things are guided. He
cannot be seen by the eye of an onlooker, but the eye which does not see Him cannot deny Him while
the mind that proves His existence cannot perceive Him. He is so high in sublimity that nothing can be
more sublime than He, while in nearness, He is so near that no one can be nearer than He. But his
sublimity does not put Him at a distance from anything of His creation, nor does His nearness bring
them on equal level to Him. He has not informed (human) wit about the limits of His qualities.
Nevertheless, He has not prevented it from securing any essential knowledge of Him. He is such that all
signs of existence testify for Him till the denying mind also believes in Him. Allah is sublime beyond what
is described by those who liken Him to things or those who deny Him.

Everything submits to Him and everything exists by Him. He is the satisfaction of the poor, the dignity of
the low, the energy for the weak and shelter for the oppressed. Whoever speaks, He hears his speaking
and whoever keeps quiet, He knows his secret. On Him is the livelihood of everyone who lives and
whoever dies returns to Him.

He is One, but not by counting. He is everlasting without any limit. He is existent without any support.
Minds admit His existence without the need for the senses. Things which can be seen testify to Him
without confronting Him. Imagination cannot encompass Him. He manifests Himself to the imagination
with his help for the imagination, refusing to be conceived by the imagination. He has made imagination
the arbiter. He is not “big” in the sense that volume is vast, so His body is also “big”. Nor is He great in
the sense that His limits should extend to the utmost, so His frame should be extensive. But He is ‘big” in
status and great in authority.

Allah is the First, but His precedence is not temporal so as to be in contradiction with His being the Last.
He is the Manifest, but not in the sense of being physically visible or perceptible by the senses; His
Manifestness does not contradict His Hiddenness. In fact, His being the First is identical to His being the
Last; similarly, His being Manifest and Hidden are identical; they are not two different things:

“Praise be to Allah, for whom one condition does not precede another, so that He may he the First
before being the Last or may be Manifest before being Hidden.”(Sermon 65)
“Time is not His accomplice, nor does He need the assistance of tools and agents His Being transcends
time. His Existence transcends nothingness and His pre-eternity transcends all beginning.”(Sermon 186)

The Divine Essence`s transcendence over time, nothingness, beginning, and end is one of the most
profound concepts of al-hikma philosophy. Allah’s pre-eternity does not mean that Allah has always
existed. Certainly Allah has always existed but the Divine pre-eternity (azaliyya) is something greater in
meaning than “existence at all times” because, “existing at all times” assumes existence in time; but
Allah`s Being has not only been at all times, It precedes time itself. This is the meaning of the Divine pre-
eternity. This shows that His precedence is something other than temporal precedence. Praise be to
Allah Whose creation bears testimony to His Existence; temporality (huduth) of whose creation is the
testimony of His “pre-eternity” the similarity and likeness among whose creation proves that He is
unique. The senses do not perceive Him and nothing can conceal Him (Sermon 152).

That is to say, Allah is both Hidden and Manifest. By Himself, He is Manifest but is Hidden from,
undetectable by, the human senses. His Hiddenness from the senses is due to man`s own limitations,
not because of Him.

It needs no proof that existence is synonymous with manifestation; the more powerful the existence of
a being, the more manifest it will be. Conversely, the weaker its being is and the more intermingled with
non-being, the less manifest it is to itself and to others.

For everything, there are two modes of being: its being-in-itself (wujud fi nafsih), and its being-for-
others (wujud fi ghayrih). The being of everything for us depends upon the structure of our senses and
on certain special conditions. Accordingly, the manifestation of a thing, a being, is also of two kinds: its
manifestation-in-itself (zuhur fi nafsih) and its manifestation-for-others (wujud fi ghayrih).

(O Allah!) The eyes have not seen You, so as to be aware of You, but You were before the describers of
Your creation. You did not create the creation on account of loneliness, nor did You make them work for
gain. He whom You catches cannot go farther than You and he whom You holds cannot escape You. He
who disobeys You does not decrease Your authority and he who obeys You does not add to Your Might.
He who disagrees with Your judgement cannot turn it and he who turns away from Your command
cannot do without You. Every secret before You is open and for You every absent is present.

You are everlasting, there is no end to You. You are the highest aim, there is no escape from You, You
are the promised (point of return) from which there is no deliverance except towards You. In Your hand
is the forelock of every creature and to You is the return of every living being. Glory to You! How great is
Your creation that we see, but how small is this greatness by the side of Your Might. How awe-striking is
Your realm that we notice, but how humble is this against what is hidden from us out of Your authority.
How extensive are Your bounties in this world, but how small are they against the bounties of the next
world.

Praise be to Allah! He is such that senses cannot perceive Him, place cannot contain Him, eyes cannot
see Him and veils cannot cover Him. He proves His eternity by the bringing His creation into existence
and (also) by originating His creation. (He proves) His existence, and through their spirituality, He proves
that there is nothing similar to Him.

He is true to His promise. He is too Sublime to be unjust to His creatures. He stands by equity among His
creation and implements justice over them in His commands. He provides testimony of His own being
through the creation of things from ever, through their marks of incapability of His power and through
their powerlessness against death of His eternity.

Mosquitoes are a family, the Culicidae, of some 3,600 species of small flies. The word "mosquito"
(formed by mosca and diminutive -ito)[2] is Spanish for "little fly".[3][4] Mosquitoes have a slender
segmented body, one pair of wings, three pairs of long hair-like legs, and specialized, highly elongated,
piercing-sucking mouthparts. All mosquitoes drink nectar from flowers; females of some species have in
addition adapted to drink blood. Evolutionary biologists view mosquitoes as micropredators, small
animals that parasitise larger ones by drinking their blood without immediately killing them. Medical
parasitologists view mosquitoes instead as vectors of disease, carrying protozoan parasites or bacterial
or viral pathogens from one host to another.

Mosquito

Early Cretaceous (Barremian) – Recent

Aedes aegypti

Scientific classificationEdit this classification

Domain: Eukaryota

Kingdom: Animalia

Phylum: Arthropoda

Class: Insecta

Order: Diptera

Superfamily: Culicoidea

Family: Culicidae

Meigen, 1818[1]

Subfamilies

Anophelinae

Culicinae

Diversity
112 genera

The mosquito life cycle consists of egg, larva, pupa, and adult stages. Eggs are laid on the water surface;
they hatch into motile larvae that feed on aquatic algae and organic material. These larvae are
important food sources for many freshwater animals, such as dragonfly nymphs, many fish, and birds
such as ducks. Adult females of many species have mouthparts adapted to pierce the skin of a host and
feed on blood of a wide range of vertebrate hosts, and some invertebrates, primarily other arthropods.
Some species only produce eggs after a blood meal.

The mosquito's saliva is transferred to the host during the bite, and can cause an itchy rash. In addition,
blood-feeding species can ingest pathogens while biting, and transmit them to later hosts. Those species
include vectors of parasitic diseases such as malaria and filariasis, and arboviral diseases such as yellow
fever and dengue fever. By transmitting diseases, mosquitoes cause the deaths of over 725,000 people
each year.

Description and life cycle

Overview

Like all flies, mosquitoes go through four stages in their life cycles: egg, larva, pupa, and adult. The first
three stages—egg, larva, and pupa—are largely aquatic,[5] the eggs usually being laid in stagnant water.
[6] They hatch to become larvae, which feed, grow, and molt until they change into pupae. The adult
mosquito emerges from the mature pupa as it floats at the water surface. Mosquitoes have adult
lifespans ranging from as short as a week to around a month. Some species overwinter as adults in
diapause.[7]

Adult

Mosquitoes have one pair of wings, with distinct scales on the surface. Their wings are long and narrow;
the legs are long and thin. The body, usually grey or black, is slender, typically 3–6 mm long. At rest,
mosquitoes hold their first pair of legs outwards, whereas the somewhat similar Chironomid midges
hold these legs forwards.[8] The Anopheles mosquito can fly for up to four hours continuously at 1 to 2
km/h,[9] traveling up to 12 km in a night. Males beat their wings between 450 and 600 times per
second, driven indirectly by muscles which vibrate the thorax.[10][11]

Mosquitoes can develop from egg to adult in hot weather in as few as five days, but it may take up to a
month.[12] At dusk, within days of pupating, males assemble in swarms, mating when females fly in.[13]
In species that need blood for the eggs to develop, the female finds a host and drinks a full meal of
blood. She then rests for two or three days to digest the meal and allow her eggs to develop. She is then
ready to lay the eggs and repeat the cycle of feeding and laying.[13] Females can live for up to three
weeks in nature, depending on temperature, humidity, their ability to obtain a blood meal, and avoiding
being killed by their vertebrate hosts.[13][14]

Anatomy of an adult female mosquito

Adult yellow fever mosquito Aedes aegypti, typical of subfamily Culicinae. Male (left) has bushy
antennae and longer palps than female (right)

Eggs

The eggs of most mosquitoes are laid in stagnant water, which may be a pond, a marsh, a temporary
puddle, a water-filled hole in a tree, or the water-trapping leaf axils of a bromeliad. Some lay near the
water's edge while others attach their eggs to aquatic plants. A few, like Opifex fuscus, can breed in salt-
marshes.[6] Wyeomyia smithii breeds in the pitchers of pitcher plants, its larvae feeding on decaying
insects that have drowned there.[15]

Oviposition, egg-laying, varies between species. Anopheles females fly over the water, touching down or
dapping to place eggs on the surface one at a time; their eggs are roughly cigar-shaped and have floats
down their sides. A female can lay 100–200 eggs in her lifetime.[13] Aedes females drop their eggs
singly, on damp mud or other surfaces near water; their eggs hatch only when flooded.[16] Females in
genera such as Culex, Culiseta, and Uranotaenia lay their eggs in floating rafts.[17][18] Mansonia
females lay their eggs in arrays, attached usually to the under-surfaces of waterlily pads.[19]

Clutches of eggs of most mosquito species hatch simultaneously, but Aedes eggs in diapause hatch
irregularly over an extended period.[16]

Anopheles eggs with side floats

Electron micrograph of a culicine egg

Culex egg raft

Larva

The mosquito larva's head has prominent mouth brushes used for feeding, a large thorax with no legs,
and a segmented abdomen. It breathes air through a siphon on its abdomen, so must come to the
surface frequently. It spends most of its time feeding on algae, bacteria, and other microbes in the
water's surface layer. It dives below the surface when disturbed. It swims either by propelling itself with
its mouth brushes, or by jerkily wriggling its body. It develops through several stages, or instars, molting
each time, after which it metamorphoses into a pupa.[12] Aedes larvae, except when very young, can
withstand drying; they go into diapause for several months if their pond dries out.[16]

Anopheles larva

Anatomy of a Culex larva

Culex larvae plus one pupa

Pupa

The head and thorax of the pupa are merged into a cephalothorax, with the abdomen curving around
underneath. The pupa or "tumbler" can swim actively by flipping its abdomen. Like the larva, the pupa
of most species must come to the surface frequently to breathe, which they do through a pair of
respiratory trumpets on their cephalothoraxes. They do not feed; typically they pass their time hanging
from the surface of the water by their respiratory trumpets. If alarmed, they swim downwards by
flipping their abdomens in much the same way as the larvae. If undisturbed, they soon float up again.
The adult emerges from the pupa at the surface of the water and flies off.[12]

Mosquito pupae, shortly before the adults emerged. The head and thorax are fused into the
cephalothorax.

Mosquito pupae, shortly before the adults emerged. The head and thorax are fused into the
cephalothorax.

Feeding by adults

Diet

Further information: Anautogeny

Female Ochlerotatus notoscriptus feeding on blood from a human arm.

Both male and female mosquitoes feed on nectar, aphid honeydew, and plant juices,[14] but in many
species the females are also blood-sucking ectoparasites. In some of those species, a blood meal is
essential for egg production; in others, it just enables the female to lay more eggs.[20] Both plant
materials and blood are useful sources of energy in the form of sugars. Blood supplies more
concentrated nutrients, such as lipids, but the main function of blood meals is to obtain proteins for egg
production.[21][22] Mosquitoes like Toxorhynchites reproduce autogenously, not needing blood meals.
Disease vector mosquitoes like Anopheles and Aedes are anautogenous, requiring blood to lay eggs.
Many Culex species are partially anautogenous, needing blood only for their second and subsequent
clutches of eggs.[23]

Host animals

Blood-sucking mosquitoes favour particular host species, though they are less selective when food is
short. Different mosquito species favor amphibians, reptiles including snakes, birds, and mammals. For
example, Culiseta melanura sucks the blood of passerine birds, but as mosquito numbers rise they attack
mammals including horses and humans, causing epidemics of Eastern equine encephalitis virus in North
America.[24] Loss of blood from many bites can add up to a large volume, occasionally causing the death
of livestock as large as cattle and horses.[25] Malaria-transmitting mosquitoes seek out caterpillars and
feed on their haemolymph,[26] causing harm.[27]

Feeding on a snake

Feeding on a frog

Feeding on a bird
Finding hosts

Blood-feeding female mosquitoes find their hosts using multiple cues, including exhaled carbon dioxide,
heat, and many different odorants.

Most mosquito species are crepuscular, feeding at dawn or dusk, and resting in a cool place through the
heat of the day.[28] Some species, such as the Asian tiger mosquito, are known to fly and feed during
daytime.[29] Female mosquitoes hunt for hosts by smelling substances such as carbon dioxide (CO2) and
1-octen-3-ol (mushroom alcohol, found in exhaled breath) produced from the host, and through visual
recognition.[30] The semiochemical that most powerfully attracts Culex quinquefasciatus is nonanal.[31]
Another attractant is sulcatone.[32] A large part of the mosquito's sense of smell, or olfactory system, is
devoted to sniffing out blood sources. Of 72 types of odor receptors on its antennae, at least 27 are
tuned to detect chemicals found in perspiration.[33] In Aedes, the search for a host takes place in two
phases. First, the mosquito flies about until it detects a host's odorants; then it flies towards them, using
the concentration of odorants as its guide.[34] Mosquitoes prefer to feed on people with type O blood,
an abundance of skin bacteria, high body heat, and pregnant women.[35][36] Individuals' attractiveness
to mosquitoes has a heritable, genetically-controlled component.[37]

Mouthparts

Further information: Insect mouthparts

Female mosquito mouthparts are highly adapted to piercing skin and sucking blood. Males only drink
sugary fluids, and have less specialized mouthparts.[38]

Externally, the most obvious feeding structure of the mosquito is the proboscis, composed of the
labium, U-shaped in section like a rain gutter, which sheaths a bundle (fascicle) of six piercing
mouthparts or stylets. These are two mandibles, two maxillae, the hypopharynx, and the labrum. The
labium bends back into a bow when the mosquito begins to bite, staying in contact with the skin and
guiding the stylets downwards. The extremely sharp tips of the labrum and maxillae are moved
backwards and forwards to saw their way into the skin, with just one thousandth of the force that would
be needed to penetrate the skin with a needle, resulting in a painless insertion.[39][40][41]

Evolution of mosquito mouthparts, with grasshopper mouthparts (shown both in situ and separately)
representing a more primitive condition. All the mouthparts except the labium are stylets, formed into a
fascicle or bundle.

Evolution of mosquito mouthparts, with grasshopper mouthparts (shown both in situ and separately)
representing a more primitive condition. All the mouthparts except the labium are stylets, formed into a
fascicle or bundle.

Mouthparts of a female mosquito while feeding on blood, showing the flexible labium sheath supporting
the piercing and sucking tube which penetrates the host's skin
Mouthparts of a female mosquito while feeding on blood, showing the flexible labium sheath supporting
the piercing and sucking tube which penetrates the host's skin

Saliva

Mosquito saliva contains enzymes that aid in sugar feeding,[42] and antimicrobial agents that control
bacterial growth in the sugar meal.[43]

For a mosquito to obtain a blood meal, it must circumvent its vertebrate host's physiological responses.
Mosquito saliva blocks the host's hemostasis system, with proteins that reduce vascular constriction,
blood clotting, and platelet aggregation, to ensure the blood keeps flowing.[44] It modulates the host's
immune response via a mixture of proteins which lower angiogenesis and immunity; create
inflammation;[44][45] suppress tumor necrosis factor release from activated mast cells;[46] suppress
interleukin (IL)-2 and IFN-γ production;[47][48] suppress T cell populations;[49][50][51] decrease
expression of interferon−α/β, making virus infections more severe;[52][53] increase natural killer T cells
in the blood; and decrease cytokine production.[54]

Egg development and blood digestion

An Anopheles stephensi female is engorged with blood and beginning to pass unwanted liquid fractions
to make room in its gut for more of the solid nutrients.

Females of many blood-feeding species need a blood meal to begin the process of egg development. A
sufficiently large blood meal triggers a hormonal cascade that leads to egg development.[55] Upon
completion of feeding, the mosquito withdraws her proboscis, and as the gut fills up, the stomach lining
secretes a peritrophic membrane that surrounds the blood. This keeps the blood separate from anything
else in the stomach. Like many Hemiptera that survive on dilute liquid diets, many adult mosquitoes
excrete surplus liquid even as they feed. This permits females to accumulate a full meal of nutrient
solids. The blood meal is digested over a period of several days.[55][56] Once blood is in the stomach,
the midgut synthesizes protease enzymes, primarily trypsin assisted by aminopeptidase, that hydrolyze
the blood proteins into free amino acids. These are used in the synthesis of vitellogenin, which in turn is
made into egg yolk protein.[55][57]

Ecology

Distribution

Mosquitoes have a cosmopolitan distribution, occurring in every land region except Antarctica[55] and a
few islands with polar or subpolar climates, such as Iceland, which is essentially free of mosquitoes.[58]
This absence is probably caused by Iceland's climate. Its weather is unpredictable, freezing but often
warming suddenly in mid-winter, making mosquitoes emerge from pupae in diapause, and then freezing
again before they can complete their life cycle.[59][60]

Eggs of temperate zone mosquitoes are more tolerant of cold than the eggs of species indigenous to
warmer regions.[61][62] Many can tolerate subzero temperatures, while adults of some species can
survive winter by sheltering in microhabitats such as buildings or hollow trees.[63] In warm and humid
tropical regions, some mosquito species are active for the entire year, but in temperate and cold regions
they hibernate or enter diapause. Arctic or subarctic mosquitoes, like some other arctic midges in
families such as Simuliidae and Ceratopogonidae may be active for only a few weeks annually as melt-
water pools form on the permafrost. During that time, though, they emerge in huge numbers in some
regions and may take up to 300 ml of blood per day from each animal in a caribou herd.[64]

For a mosquito to transmit disease, there must be favorable seasonal conditions,[65] primarily humidity,
temperature, and precipitation.[66] El Niño affects the location and number of outbreaks in East Africa,
Latin America, Southeast Asia and India. Climate change impacts the seasonal factors and in turn the
dispersal of mosquitoes.[67] Climate models can use historic data to recreate past outbreaks and to
predict the risk of vector-borne disease, based on an area's forecasted climate.[68] Mosquito-borne
diseases are currently most prevalent in East Africa, Latin America, Southeast Asia, and India. An
emergence in Europe was observed early in the 21st century. It is predicted that by 2030, the climate of
southern Great Britain will be suitable for transmission of Plasmodium vivax malaria for two months of
the year, and that by 2080, the same will be true for southern Scotland.[69][70]

Predators and parasites

Mosquito larvae are among the commonest animals in ponds, and they form an important food source
for freshwater predators. Among the many aquatic insects that catch mosquito larvae are dragonfly and
damselfly nymphs, whirligig beetles, and water striders. Vertebrate predators include fish such as catfish
and the mosquitofish, amphibians including the spadefoot toad and the giant tree frog, freshwater
turtles such as the red-eared slider, and birds such as ducks.[71]

Emerging adults are consumed at the pond surface by predatory flies including Empididae and
Dolichopodidae, and by spiders. Flying adults are captured by dragonflies and damselflies, by birds such
as swifts and swallows, and by bats.[72]

Mosquitoes are parasitised by hydrachnid mites, ciliates such as Glaucoma, microsporidians such as
Thelania, and fungi including species of Saprolegniaceae and Entomophthoraceae.[72]

Pollination

A mosquito visiting a marigold flower for nectar

Several flowers including members of the Asteraceae, Rosaceae and Orchidaceae are pollinated by
mosquitoes, which visit to obtain sugar-rich nectar. They are attracted to flowers by a range of
semiochemicals such as alcohols, aldehydes, ketones, and terpenes. Mosquitoes have visited and
pollinated flowers since the Cretaceous period. It is possible that plant-sucking preadapted mosquitoes
to blood-sucking.[14]

Parasitism

Further information: Parasitism

Ecologically, blood-feeding mosquitoes are micropredators, small animals that feed on larger animals
without immediately killing them. Evolutionary biologists see this as a form of parasitism; in Edward O.
Wilson's phrase "Parasites ... are predators that eat prey in units of less than one."[73] Micropredation is
one of six major evolutionarily stable strategies within parasitism. It is distinguished by leaving the host
still able to reproduce, unlike the activity of parasitic castrators or parasitoids; and having multiple hosts,
unlike conventional parasites.[74][75] From this perspective, mosquitoes are ectoparasites, feeding on
blood from the outside of their hosts, using their piercing mouthparts, rather than entering their bodies.
Unlike some other ectoparasites such as fleas and lice, mosquitoes do not remain constantly on the
body of the host, but visit only to feed.[76]

Evolution

Fossil record

Fossilized mosquito encased in amber

Culex malariager mosquito infected with the malarial parasite Plasmodium dominicana, in Dominican
amber of Miocene age, 15–20 million years ago[77]

The oldest known mosquitoes are currently those of Libanoculex intermedius found in Lebanese amber,
dating to the Barremian stage of the Early Cretaceous, around 125 million years ago. The mouthparts of
male individuals of this species are similar to living female mosquitoes, indicating that they consumed
blood, unlike living male mosquitoes.[78] Three other species of Cretaceous mosquito are currently
known. Burmaculex antiquus and Priscoculex burmanicus are known from Burmese amber from
Myanmar, which dates to the earliest part of the Cenomanian stage of the Late Cretaceous, around 99
million years ago.[79][80] Paleoculicis minutus, is known from Canadian amber from Alberta, Canada,
which dates to the Campanian stage of the Late Cretaceous, around 79 million years ago.[81] P.
burmanicus can be definitively assigned to the Anophelinae, indicating that the split between this
subfamily and the Culicinae took place over 99 million years ago.[80] Molecular estimates suggest that
this split occurred 197.5 million years ago, during the Early Jurassic, but that major diversification did not
take place until the Cretaceous.[82]

Taxonomy

Further information: List of mosquito genera

Over 3,500 species of mosquitoes in 112 genera have been described.[83][84][85] They are traditionally
divided into two subfamilies, the Anophelinae and the Culicinae,[86] which carry different diseases.
Roughly speaking, protozoal diseases like malaria are transmitted by anophelines, while viral diseases
such as yellow fever and dengue fever are transmitted by culicines.[87]

Phylogeny

External
Mosquitoes are members of a family of true flies (Diptera): the Culicidae (from the Latin culex, genitive
culicis, meaning "midge" or "gnat").[88] The phylogenetic tree is based on the FLYTREE project.[89][90]

Diptera

Ptychopteromorpha (phantom and primitive crane-flies)

Culicomorpha

Chironomidae (non-biting midges)

Simulioidea (blackflies and biting midges)

Culicoidea

Dixidae (meniscus midges)

Corethrellidae (frog-biting midges)

Chaoboridae (phantom midges)

Culicidae

other midges and gnats

all other flies, inc. Brachycera

(true flies)

Internal

Kyanne Reidenbach and colleagues analysed mosquito phylogenetics in 2009, using both nuclear DNA
and morphology of 26 species. They note that Anophelinae is confirmed to be rather basal, but that the
deeper parts of the tree are not well resolved.[91]

Culicidae

basal spp.

Anophelinae

Culicinae

other spp.

Aedini

other spp.

Sabethini
Interactions with humans

Anopheles albimanus mosquito feeding on a human arm. As mosquitoes are the only vectors of malaria,
controlling them reduces its incidence.

Vectors of disease

Main article: Mosquito-borne disease

Mosquitoes are vectors for many disease-causing microorganisms including bacteria, viruses, and
protozoan parasites. Nearly 700 million people acquire a mosquito-borne illness each year, resulting in
over 725,000 deaths.[92] Common mosquito-borne viral diseases include yellow fever[93] and dengue
fever transmitted mostly by Aedes aegypti.[94] Parasitic diseases transmitted by mosquitoes include
malaria and lymphatic filariasis. The Plasmodium parasites that cause malaria are carried by female
Anopheles mosquitoes. Lymphatic filariasis, the main cause of elephantiasis, is spread by a wide variety
of mosquitoes.[95] A bacterial disease spread by Culex and Culiseta mosquitoes is tularemia.[96]

Control

Mosquito nets can prevent people being bitten while they sleep.

Approaches

Main article: Mosquito control

Many measures have been tried for mosquito control, including the elimination of breeding places,
exclusion via window screens and mosquito nets, biological control with parasites such as fungi[97][98]
and nematodes,[99] or predators such as fish,[100][101][102] copepods,[103] dragonfly nymphs and
adults, and some species of lizard and gecko.[104] Another approach is to introduce large numbers of
sterile males.[105] Genetic modification methods including cytoplasmic incompatibility, chromosomal
translocations, sex distortion and gene replacement, solutions seen as inexpensive and not subject to
vector resistance, have been explored.[106] Control of disease-carrying mosquitoes may one day
become possible using gene drives.[107][108]

Repellents

Main article: Insect repellent

Mosquito repellents (including a mosquito coil) in a Finnish store

Insect repellents are applied on skin and give short-term protection against mosquito bites. The
chemical DEET repels some mosquitoes and other insects.[109] Some CDC-recommended repellents are
picaridin, eucalyptus oil (PMD), and ethyl butylacetylaminopropionate (IR3535).[110] Pyrethrum (from
Chrysanthemum species, particularly C. cinerariifolium and C. coccineum) has been reviewed favorably
in research published in 2021.[111] Electronic insect repellent devices that produce ultrasounds
intended to keep away insects (and mosquitoes) are marketed. No EPA or university study has shown
that these devices prevent humans from being bitten by a mosquito.[112]

Bites

Further information: Mosquito bite allergies

Mosquito bites lead to a variety of skin reactions and more seriously to mosquito bite allergies.[113]
Such hypersensitivity to mosquito bites is an excessive reaction to mosquito saliva proteins.[114]
Numerous species of mosquito can trigger such reactions, including Aedes aegypti, A. vexans, A.
albopictus, Anopheles sinensis, Culex pipiens,[115] Aedes communis, Anopheles stephensi,[116] C.
quinquefasciatus, C. tritaeniorhynchus,[117] and Ochlerotatus triseriatus.[118] Cross-reactivity between
salivary proteins of different mosquitoes implies that allergic responses may be caused by virtually any
mosquito species.[119] Treatment can be with anti-itch medications include some taken orally, such as
diphenhydramine, or applied to the skin like antihistamines or corticosteroids such as hydrocortisone.
Aqueous ammonia (3.6%) also provides relief.[120] Both topical heat[121] and cold may be useful as
treatments.[122]

In human culture

Greek mythology

Arthur Rackham's illustration of the fable of "The Bull and the Mosquito", 1912

Ancient Greek beast fables including "The Elephant and the Mosquito" and "The Bull and the Mosquito",
with the general moral that the large beast does not even notice the small one, derive ultimately from
Mesopotamia.[123]

Origin myths

The peoples of Siberia have origin myths surrounding the mosquito. One Ostiak myth tells of a man-
eating giant, Punegusse, who is killed by a hero but will not stay dead. The hero eventually burns the
giant, but the ashes of the fire become mosquitoes that continue to plague mankind. Other myths from
the Yakuts, Goldes (Nanai people), and Samoyed have the insect arising from the ashes or fragments of
some giant creature or demon. Similar tales found in Native North American myth, with the mosquito
arising from the ashes of a man-eater, suggest a common origin. The Tatars of the Altai had a variant of
the same myth, involving the fragments of the dead giant, Andalma-Muus, becoming mosquitoes and
other insects.[124]

Lafcadio Hearn tells that in Japan, mosquitoes are seen as reincarnations of the dead, condemned by the
errors of their former lives to the condition of Jiki-ketsu-gaki, or "blood-drinking pretas".[125]

Modern era

How a Mosquito Operates (1912)

Winsor McCay's 1912 film How a Mosquito Operates was one of the earliest works of animation. It has
been described as far ahead of its time in technical quality.[126] It depicts a giant mosquito tormenting a
sleeping man.[127]

The de Havilland Mosquito was a high-speed aircraft manufactured between 1940 and 1950, and used in
many roles.[128]

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Chapter 2The mosquito

An introduction

Frances M. Hawkes and Richard J. Hopkins.

Mosquitoes are one of the most intensely studied creatures on the planet. Their role in disease
transmission and nuisance biting makes them worthy of such attention. There are over 3,500 species of
mosquito on Earth and they are found on every continent except Antarctica. Yet, from this great
diversity, only a small handful can carry the pathogens that cause disease and it is these species which
have been studied most intensely. For the purposes of public health, this substantial body of research
has helped us to understand mosquito-borne disease transmission and informed the development of
mosquito- and disease-control methods. But parallel to this, a fascinating and complex biology has been
revealed, showing the mosquito’s incredible and unusual behavioural, anatomical and physiological
traits. This chapter will explore some of the unique aspects of mosquito biology and ecology, drawn
from the ever-growing body of research on the most medically important species, and highlight the
sophisticated adaptations of mosquitoes to their ecological niche.

Mosquitoes are some of the most intensely studied creatures on the planet and their role in disease
transmission and nuisance biting makes them worthy of such attention. There are over 3,500 species of
mosquito on earth, being found everywhere except in Antarctica. Yet, from this great diversity, only a
small handful can carry the pathogens that cause human disease and it is these species which have been
studied most thoroughly. For the purposes of public health, this substantial body of research has helped
us to understand mosquito-borne disease transmission and informed the development of mosquito and
disease control methods. A fascinating spinoff of that body of research has been to reveal a complex
biology, showing the mosquito’s incredible and unusual behavioural, anatomical and physiological traits.

Animal behaviours are linked to the intricate displays of brightly coloured tropical birds, the long-range
migrations of grazing mammals, the semaphore flashing of fireflies in a darkened landscape, together
with an infinite variety of other activities and colourful patterns across the animal kingdom. Mosquitoes,
like all animals, are driven by a fundamental set of needs. The behavioural organization of an individual
animal species is at the core of understanding the ecology of that species. In Dutch Biologist Nikolaas
Tinbergen’s classic 1963 paper, “On aims and methods of ethology,” he defines four categories of
explanations of animal behaviour: causation, evolution, development and function (Tinbergen, 1963). In
pragmatic terms, these categories can then be regarded as either “proximate” or “ultimate” causes of a
behaviour. The patterns of behaviour exhibited by mosquitoes are complex and driven by sensory
systems that are adapted to the environments they inhabit and the ecological niches they exploit. Their
life cycle, anatomy, physiology and behaviour make these creatures both an extraordinary object of
study and crucial to human culture.

Mosquitoes are holometabolous insects. This means that, just as caterpillars develop into butterflies,
mosquitoes undergo a complete metamorphosis, hatching from eggs into larvae, then into a pupal
phase (where the juvenile form liquifies and reforms into the adult body), finally hatching as fully adult
flies (Figure 2.1). All mosquitoes therefore start life as an egg with a gravid female mosquito laying as
many as 250 of them in each clutch. Since the mosquito’s juvenile stages are aquatic, eggs must be
deposited in or near water, or somewhere where water may return after flood or rainfall. The female’s
choice of location for laying eggs is a critical factor for determining the offspring’s survival in the
immature stages (eggs, larvae and pupae). Once an egg has been laid then the immature must develop
in the site selected by its mother. Broadly speaking, mosquito eggs are of two types: rapid-hatch eggs
which are laid directly on or adjacent to the water surface, which hatch within a couple of days; and
delayed-hatch eggs typically laid adjacent to water or on moist soil or vegetation some metres distant
from water. Delayed-hatch eggs can survive for long periods—months or years—being resistant to
desiccation and undergoing extremes of temperature that may include freezing winters. Egg laying
varies widely by mosquito species, from single eggs to small rafts of eggs cemented to each other and
placed directly on the water or else laid nearby, always ready to hatch as soon as the water level rises.

Figure 2.1. The mosquito life cycle.

The mosquito life cycle.

A gravid female mosquito aims to hatch her eggs in a water resource that is sufficiently rich in nutrients
and long-lasting to allow the larvae to grow, develop and produce pupae from which adult mosquitoes
can successfully emerge. Her selection of a laying site is dependent upon various chemical, visual,
olfactory and tactile cues that influence the behaviour of the female before an egg is laid (Ignell and Hill,
2020). For some Anopheles mosquitoes, water vapour itself is attractive to gravid females (Okal et al.,
2013), while tall riparian vegetation has been found to be a deterrent to laying eggs (Low et al., 2016).
Whatever the egg-laying strategy of the species, the female’s search for a location to lay her eggs may
last several days. Mosquito flight is generally favoured by warm humid conditions, and if the wind is too
strong then mosquitoes will not attempt to fly. Whilst generally associated with flights relatively close to
the ground, some gravid females have been captured hundreds of metres in the air with little being
known about these long-range movements.

Dictated by the availability of aquatic habitats, mosquitoes are found in a wide range of environments
from the tropics to the Arctic circle. Their larvae can be found wriggling in vast marshlands, flood plains
and wherever else water collects continuously or periodically, such as in small tree holes or human-
made containers, which have become ideal reproductive niches for many species. Even the water-filled
leaf axils of such plants as bromeliads can be mosquito larvae habitat. After hatching, the young
mosquito larva harvests nutrients from the water. Larvae are essentially detritivores, filter feeding on
decomposing organic matter, bacteria and algae for several weeks, spending much of their time at the
water’s surface to take in air, much like a snorkeler. As they build food reserves necessary to tide them
through the intense process of metamorphosis, larvae can be aggressively predated on by fish,
amphibians and other aquatic invertebrates. Moreover, once emerged as adults, mosquitoes transfer
huge volumes of biomass into the terrestrial food web, in turn contributing to the diets of insectivorous
mammals, birds and other invertebrates. So fundamental are mosquitoes in the food chain that when
their numbers were controlled in the Camargue region of southern France, breeding success of the
house martin, Delichon urbicum, was reduced by some 25% compared to untreated areas (Poulin et al.,
2010).

After progressing through four larval stages, or instars, the mosquito is ready to pupate. Pupae
physically resemble so many commas swimming in the water, and though they do not feed, they are
highly mobile and respond to the slightest threat by tumbling down through the water to escape
potential predation. When the mosquito pupa is fully developed, it will rise to the surface of the water
one last time. Here the adult mosquito is ready to emerge, and so straightens its body, splits open its
exoskeleton and emerges upright into the air (Figure 2.2)

Figure 2.2. An emerging male Culex pipiens.

An emerging male Culex pipiens.

After resting briefly on the surface of the water, the adult mosquito must take a short shaky flight to find
a refuge, typically in surrounding vegetation, where it rests to allow its newly pumped-up wings to dry
and properly harden.
As a consequence of the wide range of their larval habitats, there is a correspondingly large variation in
emergence patterns of different mosquito species. Whilst the number of mosquitoes that are able to
grow in and emerge from the water in a tin can will always be small, the numbers emerging from larger
bodies of water can be massive and dramatic. This is most apparent for species such as the inland
floodwater mosquito, Aedes vexans, a highly cosmopolitan mosquito found in many countries with a
range covering every continent except Antarctica and South America. Although capable of transmitting a
range of human pathogens, Aedes vexans is best known for its role as a nuisance mosquito, hence its
binomial name: the Latin vexāre means to torment or harass. This mosquito does not lay its eggs directly
in the water but in the moist soil above the waterline. After a period of drying, these eggs can survive for
years waiting for the water level to rise and be sufficiently warm. If the water is too cold or clear, the
eggs will not hatch. When suitable waters do come to inundate meadows or river flood plains, Aedes
vexans eggs can hatch across vast areas; hundreds of larvae can be found in a litre of floodwater,
equating to over 100 million larvae per hectare (Becker et al., 2010), the subsequent emergence of adult
mosquitoes reaching biblical proportions. Such population levels of this, and other floodwater species,
can become so extreme as to deter all normal human activities. Vexāre indeed.

Once emerged as adults, both male and female mosquitoes feed extensively on sources of plant-based
sugars, such as those found in nectar and fruit juices. Only the adult female mosquito seeks a blood
meal, which has the requisite nutrients to support egg development. When the female does feed on
blood it does not necessarily select humans as the source of this blood, and indeed humans are rarely a
mosquito’s main blood source. Other mammals, birds or reptiles will also satisfy the palate of some
mosquito species. It is when the mosquito selects a human for its blood meal that the insect earns its
reputation as an annoyance and harbinger of disease.

It might also be noted that while the majority of mosquito species follow this general dietary pattern,
there is a notable exception in the genus Toxorhynchites. This group includes the world’s largest
mosquito, Toxorhynchites speciosus, whose wingspan may be four times larger than most other species.
The idea of a giant mosquito may seem somewhat terrifying; however, the 90 or so species in this genus
are particularly noteworthy because the adults do not take any blood meals at all and feed exclusively
on plant sugars. This is on account of the predatory nature of the larvae, which kill live prey—including
other mosquito larvae—in order to acquire sufficient reserves of protein for egg development when
adult. So aggressive are the predatory (and even cannibalistic) appetites of Toxorhynchites amboinensis
larvae that this species has been cultivated and distributed as an effective form of mosquito biocontrol,
especially for the control of species of mosquitoes that typically breed on shipping containers and which
are associated with the transmission of dengue and Zika virus (Collins and Blackwell, 2000).

The secret life of a mosquito

Whilst for most people, mosquitoes are defined by an incessant whining that disturbs their sleep at
night or by an intense swollen itch on their ankle, there is far more to mosquito biology than their
interactions with humans. These are but brief moments in their admittedly rather short lives—in the
tropics most individuals will seldom survive longer than a month as adults. Throughout the course of
their adult life, mosquitoes must perform a series of complex searches for resources at different times
and in different places across the landscape. They interact not only with their blood-meal host, but with
plants that are also sources of food, with a mate, and with places and structures for resting and laying
their eggs. The resources that each individual insect searches for vary in their spatial and temporal
availability and are sometimes to be found in widely differing environments. The behaviour of a
searching adult mosquito is driven by external cues in the form of chemical and visual signals that the
insect must process to guide its searches.

The organs that the mosquito uses to inform the searches are all remarkable in their own way. Stimuli
that are visual, chemical and aural all play important roles at some point in the life of the adult
mosquito. The compound eyes of night-active mosquitoes are amongst the most sensitive to low light
levels in the animal kingdom and the structure of the nocturnal mosquito eye is uniquely adapted to
maximize this sensitivity. Such eye sensitivity enables visually guided flight in light conditions equivalent
to moonless, starless nights, although such sensitivity comes at the expense of visual resolution, with
the world appearing as a heavily pixelated image of light and dark patches. The mosquito antenna is
another remarkable organ. It can detect odours that distinguish potential hosts, sugar meal sources and
egg-laying (oviposition) sites. This antenna not only has humidity and thermal receptors, it is also
endowed with one of the most sensitive sound detectors in the insect kingdom, the Johnston's organ.
Described by and named after Christopher Johnston (1855), these are the most complex
mechanosensitive organs yet found in insects. This doughnut-shaped organ at the base of the antenna
detects minute vibrations from sound waves. The Johnston’s organ’s extreme sensitivity to sound allows
a flying mosquito to detect the tones produced by the wingbeats of other flying mosquitoes and
distinguish these from tones produced by its own wings. Fascinatingly, conspecific male and female
mosquitoes not only detect the sound of each other’s wingbeats, they also adjust their own wingbeat
frequency to match that of their potential partner during flight, producing a harmonious duet in a
prelude to mating (Gibson and Russell, 2006).

Often, all the resources that a mosquito needs can be found in a relatively small area, with conventional
wisdom being that mosquitoes, on the whole, travel relatively short distances, generally less than a few
hundred metres, with exceptions of up to several tens of kilometres being documented. With the high
habitat diversity and host availability found in some environments, there may be little need to travel far.
However, research carried out in Africa’s Sahel has demonstrated that malaria mosquitoes fly hundreds
of metres up, where they may be carried on the wind for distances of around 300 kilometres in a single
night (Huestis et al., 2019). The majority of these insects were found to be blood-fed females, so that
their travel over long distances may have significant repercussions for transmitting such diseases as
malaria and for the ecological aspects of their searching for egg-laying sites during prolonged periods of
drought.

In any case, flying between resources and in search of shelter requires a great deal of energy and adult
mosquitoes need to feed in order to gather the energy that flight requires. Whilst people generally think
of mosquitoes as exclusively blood-feeding, sugar feeding is a cornerstone of adult mosquito life. Newly
emerged mosquitoes cannot survive for long without taking in a sugar meal and it is the consumption of
sugar that facilitates females in their search for blood. Although female adult mosquitoes in all but a few
species require a blood meal to produce eggs, both male and females feed on nectar and other sources
of plant sugars to provide energy. Given the choice, the majority of female mosquitoes will consume a
sugar meal before they take blood, with floral nectar being by far the most important of the sugar
sources for mosquitoes. Although the phrase “sugar feeding” is commonly used, it is perhaps a
misnomer for the process of mosquitoes feeding on plants, since mosquitoes gain greater value from a
plant’s other nutrients than simple carbohydrates. “Phytophagy” may therefore be the more
appropriate way to describe mosquito feeding (Peach and Gries, 2020). It has even been further
demonstrated that compounds present in nectar can differentially affect the development of malaria
parasites within the mosquito to such an extent that their feeding on certain plants can suppress the
malaria cycle in the insect phase (Hien et al., 2016).

Since floral meals are the most significant contributor to mosquito phytophagy, then how do
mosquitoes find this key food source? In general, it is common amongst phytophagous insects that their
attraction to the plant is governed by a gestalt of signals available to guide the insect to the correct
plant. It is almost certainly the case for mosquitoes seeking nectar to fuel their flights that their search is
based on a range of signals coming from flowers. The visual cues associated with flowers and insects
have been extensively explored for day-flying pollinators, such as bees, but less is known about floral
visitation by nectar-foraging mosquitoes. Broadly speaking, nectar-foraging mosquitoes often visit
flowers that are white or pale yellow to the human eye. For nocturnal species of mosquito, this can be
linked to the fact that many flowers that make nectar available at night are often pale flowers, which
have a strong contrast against the dark landscape, thereby offering stronger visual cues for the
pollinators. In addition to visual cues, floral odours, metabolic heat and the nocturnal respiration of
carbon dioxide are all associated with floral location by mosquitoes (Peach et al., 2019). Floral feeding by
mosquitoes has been relatively understudied compared to other aspects of mosquito biology. It can be
concluded that mosquitoes do feed on extrafloral nectaries and on fruit juices, and that mosquitoes will
utilize a broader range of plant material than just flowers when flowers are scarce.

Such nectar-feeding habits mean that mosquitoes may have a role in pollination. Mosquitoes appear
able to pick up small clusters of pollen during nectar feeding, thereby facilitating plant pollination, even
if there are very few documented cases of obligate pollination by mosquitoes. The blunt-leaved orchid,
Platanthera obtusata, which can be found in the bogs, swamps and wooded fens of northern North
America, is one such mosquito-pollinated plant, with Aedes communis attracted to the orchid’s chemical
compounds (Lahondère et al., 2020).

While locating plant-based sugars is one occupation of male mosquitoes, their other chief activity is
mate-seeking. The growing importance of research about male mate-seeking stems from an interest in
mating disruption or using modified males to spread characteristics which disrupt a mosquito’s ability to
act as a vector of disease (Takken et al., 2006). Like most insect species, male adult mosquitoes typically
emerge slightly ahead of females by a day or so, requiring that extra time to become sexually mature.
When females emerge, they are often ready to mate almost immediately. It is common for many
mosquito species to have mating “swarms” formed predominantly by the males, usually around dusk.
These swarms often form close to visible structures or other conspicuous landmarks, although the exact
mechanisms that guide the positioning of these mating swarms are poorly understood. Most female
mosquitoes will mate just once during their lifetime, storing the sperm to fertilize all subsequent eggs
they produce. Alongside the genetic material that allows sexual reproduction to take place, various
proteins are also transferred from the male to the female during mating. This transmitted chemical
concoction triggers changes in female behaviour, switching from mate-seeking to searching for blood to
nourish her eggs.

Commonplace misconceptions regarding the seemingly indomitable ability of mosquitoes to locate and
bite us—and us specifically, rather than anybody else—arise from an awareness that our body odour or
something of our scent is somehow detectable and traceable by mosquitoes. Olfaction is indeed the
critical mechanism that females use to locate a suitable blood meal. The mosquito’s primary scent-
sensing organs are the antennae. These paired appendages are covered with hundreds of tiny hairs
called sensilla, each capable of detecting airborne molecules, including various chemical odours
emanating from animal skin (Sutcliffe, 1994). Breath also releases important telltale chemicals and
mosquitoes carry a pair of sensory palpi next to the antennae crucial for detecting those, too.

Like other blood-sucking insects, adult female mosquitoes are highly attracted to carbon dioxide.
Produced during respiration by every vertebrate animal and exhaled in the breath, carbon dioxide is an
extremely reliable indicator of the presence of living animals and, thus, potential blood meals. Of course,
carbon dioxide is also a natural component of the atmosphere, but malaria mosquitoes can detect
changes in concentration of as little as 0.01%. Such minute changes in concentration are sufficient to
trigger flight in mosquitoes at rest (Healy and Copland, 1995). Trails of the molecule can also be
identified by mosquitoes from some distance, with Anopheles melas able to detect plumes of carbon
dioxide at 18 metres (Gillies and Wilkes, 1969). This first detection of carbon dioxide initiates a complex
sequence of behaviours that ultimately lead the insect to its prey.

For some species of mosquito, carbon dioxide appears to be all that they require to locate the source of
a blood meal; the stronger the signal, the better. In the case of Culex tarsalis, the greater the volume of
carbon dioxide released, the more mosquitoes are attracted to the source (Reeves, 1953; Allan, Bernier
and Kline, 2006). For Anopheles species that can carry malaria, their attraction to carbon dioxide can
have important implications for the spread of mosquito-borne disease. For example, women in the later
stages of pregnancy exhale about 21% more carbon dioxide per breath, meaning that, other factors
remaining equal, the number of Anopheles gambiae attracted to pregnant women can double (Lindsay
et al., 2000). Since such women are at greater risk of complications from malaria, this particular aspect
of mosquito behaviour can produce dangerous consequences to those most vulnerable.

While the carbon dioxide in the air is a generic clue that a host may be nearby, it provides no definitive
information about which kind of animal is producing the gas. For opportunist species, such as the
Caribbean tree hole mosquito, Aedes mediovittatus, any blood-carrying animal is targeted, be it sheep,
rat, pig, horse, cow, goat, cat, dog, chicken or human (Barrera et al., 2012). Other mosquitoes are even
more catholic in their diet, with Culex erraticus, for example, also feeding on reptiles, amphibians, as
well as birds, large-hooved mammals and humans (Clements, 1999).

But several medically important mosquito species are extremely specific in their preferred host. Indeed,
a key factor in making Anopheles gambiae a highly efficient malaria vector is its faithful choice of
humans for a blood meal, a trait known as anthropophily. Once infected with human malaria parasites
after the first feeding, a female mosquito can feed several more times, potentially transmitting parasites
to a new human host each time (a cow, to consider a different vertebrate, cannot become infected with
human malaria and so suffers no ill-effects if fed upon by mosquitoes carrying human malaria; from the
human point of view, every blood meal taken from non-human vertebrates is one less chance to spread
infection). Anopheles gambiae evolved its extreme specialization to humans through ongoing
association with agricultural communities, which provided it with reliable sources of food and niches for
resting and oviposition (Besansky et al., 2004). How, then, can this species that is so discerning in its
host, distinguish between all animal sources of carbon dioxide and pinpoint a human? And is every
person’s scent equally appetizing to a hungry mosquito?

An animal’s emanations of sweat, breath and bacteria are composed of a complicated array of volatile
compounds. Around 350 different chemicals have been identified in human skin odours alone (Bernier
et al., 2000). Anopheles gambiae’s sensitive antennae detect those chemicals that are tied most closely
to humans alone. Chemical analysis and direct recording of electrical signals from live Anopheles
gambiae antennae revealed the identity of these compounds (Cork and Park, 1996). Yet even if a
mosquito can detect an odour compound, it does not always respond to it, making behavioural research
into their responses to chemical stimuli a delicate operation, with results that may vary according to an
odour’s concentration, volume or the presence of other odours at the same time. Key, behaviour-
influencing compounds include carboxylic acids, lactic acid and ammonia (related to sweat production
and its incubation, respectively), and octenol, which is more abundant in cattle, for example, than
humans and may therefore aid Anopheles gambiae in discriminating between them. All of these odours
stem from communities of bacteria that live harmlessly on our skin but vary from person to person, and
it is these individual differences that affect attractiveness to mosquitoes, regardless of the gender or age
of the person (Verhulst et al., 2011); thus, certain unfortunate people are genuinely more attractive to
malaria mosquitoes than others, simply by virtue of their unique skin microbiota (Qiu et al., 2006).
Moreover, recent evidence suggests that an infection of Plasmodium parasites enhances one’s
production of volatile chemicals attractive to malaria mosquitoes (Robinson et al., 2018). In this way,
Plasmodium seems to be luring mosquitoes in for a blood meal that will serve to propagate the
plasmodium.

How valid are claims that consuming garlic or spicy foods, or vitamin B supplements, might disguise a
person’s odour fingerprint, and make it less appealing to mosquitoes? There is limited evidence that a
person’s diet can alter his or her attractiveness to a mosquito—although the results of a study about
beer consumption offer some intriguing news: controlling for individual variation and baseline
attractiveness, researchers found that those drinking a litre of beer caused more malaria mosquitoes to
fly towards them than those drinking a litre of water (Lefèvre et al., 2010). Researchers speculate that
ingesting alcohol causes changes in breath and volatile odours that are more attractive to Anopheles
gambiae—with implications for beverage choice when needing to confront this particular mosquito.

Beyond the dangers presented by a handful of disease vectors and nuisance biters, most other mosquito
species will rarely, if ever, bite a person. Much less is known of the details of life histories and ecological
interactions of these other species, as they have not been deemed so worthy of research. However,
some investigations are beginning to reveal the stunning complexity in other mosquito–host
interactions.

Cold-blooded hosts, such as toads, frogs, salamanders, lizards and even mudskippers (amphibious fish),
are important blood sources for many mosquitoes in the genera Mimomyia, Uranotaenia and
Deinocerites. Since body heat has been shown to be an important signal for other mosquitoes locating
hosts over short distances, these colder hosts present a challenge. It turns out that several mosquito
species, including Japanese Uranotaenia yaeyamana, American pale-footed Uranotaenia lowii and
European Uranotaenia unguiculata, are apparently sensitive to the sounds of their hosts, since
recordings of certain frog calls have been shown to attract these mosquitoes (Borkent and Belton, 2006;
Tamashiro et al., 2011; Camp et al., 2018).

While the idea of mosquitoes feeding on frogs and toads may seem peculiar, there are very few groups
of animals off the menu to at least one or two species of mosquito. Researchers have recently identified
the mystery host animals of Uranotaenia sapphirina, a mosquito from eastern North America
characterized by attractive stripes of iridescent blue scales, tracing it to various annelid worms and
leeches (Reeves et al., 2018). Even fish blood has been extracted and identified from the gut of
engorged Aedes baisasi mosquitoes (Tamashiro et al., 2011). DNA sequencing of blood meals has been
used to verify that these mosquitoes also feed on various species of eel, goby, mudskipper, rockskipper
(or blenny) and triggerfish common to mangrove lagoons and rocky reefs (Miyake et al., 2019). Many of
these fish are amphibious or air-breathing and some eels will wriggle out of the water and across muddy
ground, while others inadvertently expose their upperparts to air when feeding in shallow waters. Aedes
baisasi is able to exploit these brief moments of vulnerability.

Whatever a mosquito’s choice of host animal, the insect must undertake the risky business of landing on
and puncturing the host’s skin. The mouthparts of all insects are derived from common structures, with
adaptations to suit their particular diets, be they nectar, grain or, in the case of mosquitoes, blood. A
mosquito’s long and slender mouthparts, collectively known as a proboscis, are supremely adapted to
the task of blood feeding and give the mosquito a reputation as a flying syringe—yet the underlying
anatomy is far more sophisticated than a simple needle. Once the female has landed on a suitable host,
she begins to probe, repeatedly driving her mouthparts into the host. Using a pair of blade-like
mandibles, which in other insects may grasp or slice food, the mosquito pins the host skin firmly in
place. A second pair of serrated maxillae proceed to saw through the surface of the skin. These
appendages are so sharp that the host is often quite unaware of their action. Once the skin is pierced,
the mosquito inserts two hollow tubular structures through the skin to search for a blood vessel from
which to draw the vital fluid. The first tube, called the labrum, moves freely, bending and curving as it
probes the tissue until it detects a suitable capillary (Choumet et al., 2012). When a vessel is located, the
labrum pierces it and begins drawing up the host’s protein-rich blood. Meanwhile, the second tubular
structure, called the hypopharynx, injects the mosquito’s saliva into the surrounding tissue. This saliva
contains over 100 proteins that keep the blood flowing while slowing the immune and defensive
responses of the host (Vogt et al., 2018). The proteins include also anesthetics that numb the area
surrounding the bite, anti-inflammatories to maintain blood pressure, vasodilators to keep blood vessels
wide and anti-clotting agents to keep the blood flowing near the feeding site.
It is the host’s inflammatory immune response to mosquito saliva that results in the painful, itchy welts
associated with the insect’s bite. Yet, the impressive array of chemicals within mosquito saliva may also
include compounds of significant value in developing the next generation of pharmaceutical medicines.
Researchers have found that anophelin, the salivary protein produced by Anopheles gambiae as an anti-
coagulant, can be modified for helping to dissolve human blood clots, thereby opening up new
possibilities for developing novel drugs that can prevent stroke and deep vein thrombosis (Watson et al.,
2018).

A mosquito’s reliance on blood feeding is the characteristic that allows mosquitoes to transmit
pathogens from infected to healthy hosts. Pathogens are taken up by mosquitoes incidentally when the
latter are imbibing a blood meal. These viruses and parasites, which accumulate in the mosquito’s
salivary glands, are then injected into new hosts along with saliva during the probing phase. However,
this is not the full story, since only a small fraction of the 3,500 mosquito species can transmit human
pathogens, mostly limited to those of just three genera: Anopheles, Aedes and Culex. Only mosquitoes
of the genus Anopheles can transmit human malaria parasites, and only three species of over 500
described anophelines are responsible for the majority of malaria transmission. It is worth remembering
that most species of mosquitoes rarely, or never, bite humans, having specialized instead to take blood
meals from other mammalian, avian, reptilian or amphibian hosts.

Moreover, the simple exchange of blood and saliva is insufficient for making a mosquito into a vector, at
least not instantaneously. Most mosquito-borne pathogens must undergo a process of replication or
development within the body of the mosquito, which can take up to 23 days for Plasmodium vivax
(Thomas et al., 2018), but can be as quick as two days in the case of dengue viruses, depending on
environmental conditions (Chan and Johansson, 2012). Should the insect feed on another host before
this process is complete, she will not yet be able to pass on viable infective agents. Yet proof that the
mosquito can successfully imbibe blood from infected hosts, and then inject pathogen-laden saliva into
uninfected hosts, is shown by the nearly 700 million cases of mosquito-borne illnesses occurring each
year (World Mosquito Program, 2020).

It would be a mistake to assume that the presence of the pathogen is unproblematic to the mosquito,
for even the immune system of the vector will attempt to tackle the invading pathogen (Rodrigues et al.,
2010). A female mosquito consuming a meal from a malaria-infected human can ingest thousands of
gametocytes (the stage of malaria parasites found in infected human blood). By digesting these
gametocytes and exposing them to other toxic processes, her immune system can reduce their numbers
from thousands to the tens (Smith et al., 2014). Moreover, white blood cells belonging to the host
animal and ingested by the mosquito during feeding will also continue to target the malaria parasite for
hours after entering the mosquito gut (Lensen et al., 1997). But even this two-pronged attack can fail to
halt the malaria parasite’s growth. Despite these gametocyte elimination processes, a small number
may still survive to form oocysts (the parasite’s next developmental stage) on the outside of the
mosquito’s gut lining. When mature, these burst open, each releasing thousands of sporozoites, which
migrate to the mosquito’s salivary glands ready to infect the next suitable host. The successful
development of just one oocyst is, therefore, all it takes for the mosquito to become infectious.
Vectors, too, can be stricken by the effects of the pathogens which infect them following a blood meal.
Dengue virus, for instance, detrimentally affects the fecundity and fitness of its principal vector, Aedes
aegypti. Dengue-infected mosquitoes lay fewer or no eggs, with adult longevity being halved (Sylvestre
et al., 2013). Malaria parasites can also upset the normal reproductive processes in anophelines. By
promoting cell death in the lining of the insect midgut, the rodent malaria, Plasmodium yoelii, is found
to cause Anopheles stephensi to reabsorb ovarian follicles, essentially destroying the next clutch of eggs
the female would have produced (Hopwood et al., 2001).

In other considerations, certain parasites have been shown to manipulate their hosts for enhancing their
own chances of survival. Malaria parasites have highly complex associations with their mosquito and
vertebrate hosts, and it has been suggested that these single-celled protozoa may also influence
mosquito physiology and behaviour to increase the probability of their survival and transmission. The
introduction of a malaria parasite into a mosquito vector undoubtedly changes the mosquito, although
whether such changes are truly an adaptive manipulation or simply a side effect of infection can be
difficult to determine (Hurd, 2003). Nonetheless, several studies suggest that parasites produce
physiological and behavioural changes in mosquitoes that favour their onward transmission.

It has been determined that wild Anopheles gambiae mosquitoes infected with the most deadly human
malaria parasite, Plasmodium falciparum, take blood meals from more than one person in a single night,
whereas uninfected mosquitoes are more likely to feed on a single individual. Such multiple feeding
behaviour increases the number of contacts between the infected mosquito vector and human hosts,
thereby increasing the transmission potential of the parasite (Koella et al., 1998). Laboratory studies also
emphasize the complexity of the vector–host relationship. Research has shown that Anopheles gambiae
mosquitoes infected with Plasmodium berghei (a strain of rodent malaria) were willing to probe a
mouse’s skin more often than uninfected mosquitoes (Choumet et al., 2012). Although the source of this
behaviour is unknown, it may nonetheless confer advantage to the parasite by increasing the likelihood
that a mosquito successfully feeds on blood and passes the parasites to the next host.

Ultimately, we realize that the unflattering reputation of mosquitoes may be well-earned. Their visceral
parasitic strategy for acquiring nutrition can certainly appear more gruesome, and less honourable, than
other predators going about their lives in the animal kingdom. That they stalk us and frequently attack in
stealth, often under the cover of darkness, and target specific individuals may be perceived at some
level as both devious and personal. That their feeding aggravates us, disturbs our rest, creates an itch
that can stay for days and may even infect us with debilitating and potentially deadly disease elevates
mosquitoes to the level of the positively dastardly. It may therefore seem unsurprising for those of us
involved in the study of this insect to be asked, “What is the point of a mosquito?” Although scientists
aim to avoid teleological explanations for natural phenomena, some of the less commonly studied
aspects of mosquito biology already demonstrate biotic interactions that are much more complicated
than those of a simple blood-sucking pest and carrier of disease. Although we are just beginning to
uncover some of their more cryptic behaviours, much work remains to be done. Thousands of mosquito
species have evolved marvelous and intricate biological adaptations for generating diverse behavioural
and ecological traits that are still unknown to science. The activities of mosquitoes in the ecosystem are
as sophisticated and specialized as that of any other creature, and indeed more complex than many. The
study of mosquito biology may reveal biochemical, anatomical and behavioural secrets that may not
only enrich our understanding of nature but also become a source of bioinspiration in future sciences
and technologies, from the design of pain-free microneedles (Gurera et al., 2018) to algorithms for flying
drones (Nakata et al., 2020). Critically, the overwhelming majority of insects that fall within the Culicidae
do not pose a threat to human health or comfort, so caution must be exercised when discussing
“mosquitoes”, generically, as carriers of disease. For the mosquitoes that are vectors of disease, it is the
pathogens and parasites they harbour which cause us morbidity and mortality, and in some cases not
without cost to the infected mosquito itself. This is key. While malaria cannot persist without
mosquitoes, mosquitoes can persist without malaria, or dengue, or Zika. When driven to distraction by
the whining of a mosquito or the itch from their bite, many will not realize that “that wretched
mosquito” is but one of myriad species each occupying a unique niche in the environment. Appreciating
these subtleties in how we frame debates about “mosquito eradication” can inform a more nuanced
discussion, where these key differences call for differences in our response.

Go to:

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E4218. doi:10.1073/pnas.1721610115. [PMC free article] [PubMed]

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53–70. doi:10.7601/mez.62.53.

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of the extrinsic incubation period of plasmodium vivax and plasmodium falciparum: A study from a
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World Mosquito Program. 2020. Mosquito-Borne Diseases. Available at: https://www

.worldmosquitoprogram.org/en/learn/mosquito-borne-diseases.

Frances M. Hawkes is a Senior Research Fellow at the Natural Resources Institute, University of
Greenwich. Incorporating laboratory and field-based studies on mosquito behaviour and ecology, her
research spans temperate and tropical species, identifying traits that can be exploited in supporting
vector control. Her work developing new mosquito surveillance techniques was recognized in a 2019
Queen’s Anniversary Prize for smart and sustainable pest management.

Richard J. Hopkins is Professor of Behavioural Entomology at the Natural Resources Institute, University
of Greenwich. His research involves the study of the searching behaviour of crop pests and vectors of
diseases in temperate and tropical regions. A particular interest in piecing together the different aspects
of how mosquitoes locate breeding and resting sites reflects the importance of understanding the
ecology of vectors in the wider sense, and contributed to the award of the 2019 Queen’s Anniversary
Prize for smart and sustainable pest management to NRI.

ANOPHOLES MOSQUITO
Anopheles or Marsh Mosquitoes(/əˈnɒfɪliːz/[1]) is a genus of mosquito first described and named by J.
W. Meigen in 1818.[2] About 460 species are recognized. Many are vectors of malaria in birds, reptiles,
and mammals. Over 100 can transmit human malaria, but only 30–40 of those commonly transmit
parasites of the genus Plasmodium, which cause the disease in endemic areas. Anopheles gambiae is
one of the best known, because of its predominant role in the transmission of the most dangerous
malaria parasite species (to humans) – Plasmodium falciparum.

Anopheles

Scientific classification

Domain: Eukaryota

Kingdom: Animalia

Phylum: Arthropoda

Class: Insecta

Order: Diptera

Family: Culicidae

Subfamily: Anophelinae

Genus: Anopheles

The name comes from the Ancient Greek word ἀνωφελής anōphelḗs 'useless', derived from ἀν- an-,
'not', 'un-' and ὄφελος óphelos 'profit'.[3][1]

Mosquitoes in other genera (Aedes, Culex, Culiseta, Haemagogus, and Ochlerotatus) can also serve as
vectors of disease agents, but not human malaria.

Evolution

The ancestors of Drosophila and the mosquitoes diverged 260 million years ago.[4] The culicine and
Anopheles clades of mosquitoes diverged between 120 million years ago and 150 million years ago.[4]
[5] The Old and New World Anopheles species subsequently diverged between 80 million years ago and
95 million years ago.[4][5] Anopheles darlingi diverged from the African and Asian malaria vectors ~100
million years ago.[6] The Anopheles gambiae and Anopheles funestus clades diverged between 80
million years ago and 36 million years ago. A molecular study of several genes in seven species has
provided additional support for an expansion of this genus during the Cretaceous period.[7]
The Anopheles genome, at 230–284 million base pairs (Mbp), is comparable in size to that of Drosophila,
but considerably smaller than those found in other culicine genomes (528 Mbp–1.9 Gbp). Like most
culicine species, the genome is diploid with six chromosomes.[[]]

The only known fossils of this genus are those of Anopheles (Nyssorhynchus) dominicanus Zavortink &
Poinar contained in Dominican amber from the Late Eocene (40.4 million years ago to 33.9 million years
ago) and Anopheles rottensis Statz contained in German amber from the Late Oligocene (28.4 million
years ago to 23 million years ago).[[]]

Systematics

Main article: Taxonomy of Anopheles

The genus Anopheles Meigen (nearly worldwide distribution) belongs to the subfamily Anophelinae
together with another two genera: Bironella Theobald (Australia only) and Chagasia Cruz (Neotropics).
The taxonomy remains incompletely settled. Classification into species is based on morphological
characteristics – wing spots, head anatomy, larval and pupal anatomy, chromosome structure, and more
recently, on DNA sequences. In the taxonomy published by Harbach et al in 2016, it was shown that
three species of Bironella: confusa, gracilis, and hollandi are phylogenetically similar Anopheles
kyondawensis than other Bironella species. The same phylogeny also argues that, based on genetic
similarity, Anopheles implexus is actually divergent from the common ancestor to the Anopheles genus,
raising new questions regarding taxonomy and classification.[8]

The genus has been subdivided into seven subgenera based primarily on the number and positions of
specialized setae on the gonocoxites of the male genitalia. The system of subgenera originated with the
work of Christophers, who in 1915 described three subgenera: Anopheles (widely distributed),
Myzomyia (later renamed Cellia) (Old World) and Nyssorhynchus (Neotropical). Nyssorhynchus was first
described as Lavernia by Frederick Vincent Theobald. Frederick Wallace Edwards in 1932 added the
subgenus Stethomyia (Neotropical distribution). Kerteszia was also described by Edwards in 1932, but
then recognised as a subgrouping of Nyssorhynchus. It was elevated to subgenus status by Komp in
1937, and it is also found in the Neotropics. Two additional subgenera have since been recognised:
Baimaia (Southeast Asia only) by Harbach et al. in 2005 and Lophopodomyia (Neotropical) by Antunes in
1937.[[]]

Two main groupings within the genus Anopheles are used: one formed by the Celia and Anopheles
subgenera and a second by Kerteszia, Lophopodomyia and Nyssorhynchus. Subgenus Stethomyia is an
outlier with respect to these two taxa. Within the second group, Kerteszia and Nyssorhynchus appear to
be sister taxa.[[]]

The number of species currently recognised within the subgenera is given here in parentheses:
Anopheles (206 species), Baimaia (1), Cellia (216), Kerteszia (12), Lophopodomyia (6), Nyssorhynchus
(34) and Stethomyia (5).
Taxonomic units between subgenus and species are not currently recognised as official zoological
names. In practice, a number of taxonomic levels have been introduced. The larger subgenera
(Anopheles, Cellia and Nyssorhynchus) have been subdivided into sections and series which in turn have
been divided into groups and subgroups. Below subgroup but above species level is the species complex.
Taxonomic levels above species complex can be distinguished on morphological grounds. Species within
a species complex are either morphologically identical or extremely similar and can only be reliably
separated by microscopic examination of the chromosomes or DNA sequencing. The classification
continues to be revised.

Subgenus Nyssorhynchus has been divided in three sections: Albimanus (19 species), Argyritarsis (11
species) and Myzorhynchella (4 species). The Argyritarsis section has been subdivided into Albitarsis and
Argyritarsis groups.

The Anopheles group was divided by Edwards into four series: Anopheles (worldwide), Myzorhynchus
(Palearctic, Oriental, Australasian and Afrotropical), Cycloleppteron (Neotropical) and Lophoscelomyia
(Oriental); and two groups, Arribalzagia (Neotropical) and Christya (Afrotropical). Reid and Knight (1961)
modified this classification and consequently subdivided the subgenus Anopheles into two sections,
Angusticorn and Laticorn and six series. The Arribalzagia and Christya Groups were considered to be
series. The Laticorn Section includes the Arribalzagia (24 species), Christya and Myzorhynchus series. The
Angusticorn section includes members of the Anopheles, Cycloleppteron and Lophoscelomyia series.[[]]

All species known to carry human malaria lie within either the Myzorhynchus or the Anopheles series.
[[]]

Life stages

Like all mosquitoes, anophelines go through four stages in their life cycles: egg, larva, pupa, and imago.
The first three stages are aquatic and together last 5–14 days, depending on the species and the
ambient temperature. The adult stage is when the female Anopheles mosquito acts as malaria vector.
The adult females can live up to a month (or more in captivity), but most probably do not live more than
two weeks in nature.[9]

Eggs

Anopheles eggs with their distinctive side floats

Adult females lay 50–200 eggs per oviposition. The eggs are quite small (about 0.5 × 0.2 mm). Eggs are
laid singly and directly on water. They are unique in that they have floats on either side. Eggs are not
resistant to drying and hatch within 2–3 days, although hatching may take up to 2–3 weeks in colder
climates.[9]

Larvae

Anopheles larva
Feeding position of an Anopheles larva (A), compared to that of a culicine mosquito (B)

The mosquito larva has a well-developed head with mouth brushes used for feeding, a large thorax and
a nine-segment abdomen. It has no legs. In contrast to other mosquitoes, the Anopheles larva lacks a
respiratory siphon, so it positions itself so that its body is parallel to the surface of the water.[9] In
contrast, the feeding larva of a nonanopheline mosquito species attaches itself to the water surface with
its posterior siphon, with its body pointing downwards.

Larvae breathe through spiracles located on the eighth abdominal segment, so must come to the surface
frequently. The larvae spend most of their time feeding on algae, bacteria, and other microorganisms in
the surface microlayer. They dive below the surface only when disturbed. Larvae swim either by jerky
movements of the entire body or through propulsion with the mouth brushes.[9]

Larvae develop through four stages, or instars, after which they metamorphose into pupae. At the end
of each instar, the larvae molt, shedding their exoskeletons, or skin, to allow for further growth.[9] First-
stage larvae are about 1 mm in length; fourth-stage larvae are normally 5–8 mm in length.

The process from egg-laying to emergence of the adult is temperature dependent, with a minimum time
of seven days.

The larvae occur in a wide range of habitats, but most species prefer clean, unpolluted water. Larvae of
Anopheles mosquitoes have been found in freshwater or saltwater marshes, mangrove swamps, rice
fields, grassy ditches, the edges of streams and rivers, and small, temporary rain pools. Many species
prefer habitats with vegetation. Others prefer habitats with none. Some breed in open, sun-lit pools,
while others are found only in shaded breeding sites in forests. A few species breed in tree holes or the
leaf axils of some plants.[9]

Pupae

The pupa (also known as the tumbler) is comma-shaped when viewed from the side. The head and
thorax are merged into a cephalothorax with the abdomen curving around underneath. As with the
larvae, pupae must come to the surface frequently to breathe, which they do through a pair of
respiratory trumpets on their cephalothoraces. After a few days as a pupa, the dorsal surface of the
cephalothorax splits and the adult mosquito emerges.[9] The pupal stage lasts around 2–3 days in
temperate areas.

Adults

Resting positions of adult Anopheles (A, B), compared to a culicine mosquito (C)

The duration from egg to adult varies considerably among species, and is strongly influenced by ambient
temperature. Mosquitoes can develop from egg to adult in as little as five days, but it can take 10–14
days in tropical conditions.
Like all mosquitoes, adult Anopheles species have slender bodies with three sections: head, thorax and
abdomen.[9]

The head is specialized for acquiring sensory information and for feeding. It contains the eyes and a pair
of long, many-segmented antennae. The antennae are important for detecting host odors, as well as
odors of breeding sites where females lay eggs. The head also has an elongated, forward-projecting
proboscis used for feeding, and two maxillary palps.[9] These palps also carry the receptors for carbon
dioxide, a major attractant for the location of the mosquito's host.

The thorax is specialized for locomotion. Three pairs of legs and a pair of wings are attached to the
thorax.[9]

The abdomen is specialized for food digestion and egg development. This segmented body part expands
considerably when a female takes a blood meal. The blood is digested over time, serving as a source of
protein for the production of eggs, which gradually fill the abdomen.[9]

Anopheles mosquitoes can be distinguished from other mosquitoes by the palps, which are as long as
the proboscis, and by the presence of discrete blocks of black and white scales on the wings. Adults can
also be identified by their typical resting position: males and females rest with their abdomens sticking
up in the air rather than parallel to the surface on which they are resting.[9]

Adult mosquitoes usually mate within a few days after emerging from the pupal stage. In most species,
the males form large swarms, usually around dusk, and the females fly into the swarms to mate.[9]

Key to the morphology of female Anopheles

Males live for about a week, feeding on nectar and other sources of sugar. Males cannot feed on blood,
as it appears to produce toxic effects and kills them within a few days, around the same lifespan as a
water-only diet.[10] Females will also feed on sugar sources for energy, but usually require a blood meal
for the development of eggs. After obtaining a full blood meal, the female will rest for a few days while
the blood is digested and eggs are developed. This process depends on the temperature, but usually
takes 2–3 days in tropical conditions. Once the eggs are fully developed, the female lays them and
resumes host-seeking.[9]

The cycle repeats itself until the female dies. While females can live longer than a month in captivity,
most do not live longer than one to two weeks in nature. Their lifespans depend on temperature,
humidity, and their ability to successfully obtain a blood meal while avoiding host defenses.[9]

In a study by the London School of Hygiene & Tropical Medicine researchers found that female
mosquitoes carrying malaria parasites are significantly more attracted to human breath and odours than
uninfected mosquitoes.[11] The research team infected laboratory-raised Anopheles gambiae
mosquitoes with Plasmodium parasites, leaving a control group uninfected. Then tests were run on the
two groups to record their attraction to human smells. Female mosquitoes are particularly drawn to foot
odours, and one of the tests showed infected mosquitoes landing and biting a prospective host
repeatedly. The team speculates that the parasite improves the mosquitoes' sense of smell.[12] It may
also reduce its risk aversion.

Ecology

Distribution and habitat

World distribution of Anopheles species

Anopheles species live both in tropical areas known for malaria such as sub-Saharan Africa, and in colder
latitudes. Indeed, malaria outbreaks have, in the past, occurred in colder climates, for example during
the construction of the Rideau Canal in Canada during the 1820s.[13] Anopheles species that can
transmit malaria are found not only in malaria-endemic areas, but also in areas where malaria has been
eliminated. These are thus constantly at risk of reintroduction of the disease.[14]

Anopheles mosquitoes require bodies of water, possibly small and seasonal, from ponds to water tanks,
swamps, ditches and puddles for their aquatic larvae and pupae.[15] The adults can however live in dry
regions such as Africa's savanna and Sahel. They can travel far from water, and are sometimes blown
hundreds of kilometres by suitable winds. Adults can aestivate for months at a time, becoming dormant
in hot dry weather, allowing them to persist through the African dry season.[16] Further, Anopheles
mosquitoes have been documented travelling in baggage, such as on aircraft.[17]

Parasites

A number of parasites of this genus are known to exist, including microsporidia of the genera
Amblyospora, Crepidulospora, Senoma and Parathelohania.[18]

Microsporidia infecting the aquatic stages of insects, a group that includes mosquitoes and black flies,
and copepods appear to form a distinct clade from those infecting terrestrial insects and fish. Two
distinct life cycles are found in this group. In the first type, the parasite is transmitted by the oral route
and is relatively species nonspecific. In the second, while again the oral route is the usual route of
infection, the parasite is ingested within an already infected intermediate host. Infection of the insect
larval form is frequently tissue-specific, and commonly involves the fat body. Vertical (transovarial)
transmission is also known to occur.[[]]

Few phylogenetic studies of these parasites have been done, and their relationship to their mosquito
hosts is still being determined. One study suggested Parathelohania is an early diverging genus within
this group.[19]

The parasite Wolbachia bacteria have also been studied for use as control agents.[20]

Predators

The jumping spider Evarcha culicivora indirectly feeds on vertebrate blood by preying on female
Anopheles mosquitos.[21] Interestingly, juvenile spiders choose the Anopheles over all other prey
regardless of whether it actually is carrying blood.[22] Juvenile spiders have adopted an Anopheles-
specific prey-capture behavior, using the posture of Anopheles mosquitoes as a primary cue to identify
them.[21] Anopheles has a distinctive resting posture with its abdomen angled up. In this case, the
spider takes a detour and approaches from behind the mosquito and under its abdomen, and then
attacks from below.[23]

Malaria vectors

Some species are poor vectors of malaria, as the parasites do not develop well (or at all) within them.
There is also variation within species. In the laboratory, it is possible to select strains of A. gambiae that
are refractory to infection by malaria parasites. These refractory strains have an immune response that
encapsulates and kills the parasites after they have invaded the mosquito's stomach wall. Scientists are
studying the genetic mechanism for this response. Genetically modified mosquitoes refractory to
malaria possibly could replace wild mosquitoes, thereby limiting or eliminating malaria transmission.[[]]

Preferred sources for blood meals

One important behavioral factor is the degree to which an Anopheles species prefers to feed on humans
(anthropophily) or animals such as cattle or birds (zoophily). Anthropophilic Anopheles are more likely to
transmit the malaria parasites from one person to another. Most Anopheles mosquitoes are not
exclusively anthropophilic or zoophilic, including the primary malaria vector in the western United
States, A. freeborni.[24][25] However, the primary malaria vectors in Africa, A. gambiae and A. funestus,
are strongly anthropophilic and, consequently, are two of the most efficient malaria vectors in the
world.[9]

Once ingested by a mosquito, malaria parasites must undergo development within the mosquito before
they are infectious to humans. The time required for development in the mosquito (the extrinsic
incubation period) ranges from 10–21 days, depending on the parasite species and the temperature. If a
mosquito does not survive longer than the extrinsic incubation period, then she will not be able to
transmit any malaria parasites.[9]

It is not possible to measure directly the lifespans of mosquitoes in nature, but indirect estimates of
daily survivorship have been made for several Anopheles species. Estimates of daily survivorship of A.
gambiae in Tanzania ranged from 0.77 to 0.84, meaning at the end of one day, between 77% and 84%
will have survived.[26][9]

Assuming this survivorship is constant through the adult life of a mosquito, less than 10% of female A.
gambiae would survive longer than a 14-day extrinsic incubation period. If daily survivorship increased
to 0.9, over 20% of mosquitoes would survive longer than the same period. Control measures that rely
on insecticides (e.g. indoor residual spraying) may actually impact malaria transmission more through
their effect on adult longevity than through their effect on the population of adult mosquitoes.[9]

Patterns of feeding and resting

Most Anopheles mosquitoes are crepuscular (active at dusk or dawn) or nocturnal (active at night).
Some feed indoors (endophagic), while others feed outdoors (exophagic). After feeding, some blood
mosquitoes prefer to rest indoors (endophilic), while others prefer to rest outdoors (exophilic),[9]
though this can differ regionally based on local vector ecotype, and vector chromosomal makeup, as
well as housing type and local microclimatic conditions.[[]] Biting by nocturnal, endophagic Anopheles
mosquitoes can be markedly reduced through the use of insecticide-treated bed nets or through
improved housing construction to prevent mosquito entry (e.g. window screens). Endophilic mosquitoes
are readily controlled by indoor spraying of residual insecticides. In contrast, exophagic/exophilic vectors
are best controlled through source reduction (destruction of the breeding sites).[9]

Gut flora

Because transmission of disease by the mosquito requires ingestion of blood, the gut flora may have a
bearing on the success of infection of the mosquito host. This aspect of disease transmission has not
been investigated until recently.[27] The larval and pupal gut is largely colonized by photosynthetic
cyanobacteria, while in the adult, Pseudomonadota and Bacteroidota predominate. Blood meals
drastically reduce the diversity of organisms and favor enteric bacteria.[27]

Interaction with himans

Insecticide control and resistance

Insecticide-based control measures (e.g. indoor spraying with insecticides, bed nets) are the principal
ways to kill mosquitoes that bite indoors. However, after prolonged exposure to an insecticide over
several generations, mosquito populations, like those of other insects, may evolve resistance, a capacity
to survive contact with an insecticide. Since mosquitoes can have many generations per year, high levels
of resistance can evolve very quickly. Resistance of mosquitoes to some insecticides has been
documented with just within a few years after the insecticides were introduced. Over 125 mosquito
species have documented resistance to one or more insecticides. The evolution of resistance to
insecticides used for indoor residual spraying was a major impediment during the Global Malaria
Eradication Campaign. Judicious use of insecticides for mosquito control can limit the evolution and
spread of resistance. However, use of insecticides in agriculture has often been implicated as
contributing to resistance in mosquito populations. Detection of evolving resistance in mosquito
populations is possible, so control programs are well advised to conduct surveillance for this potential
problem.[9] In Malawi and other places, a shrub known as mpungabwi (Ocimum americanum) is used to
repel mosquitoes.[28]

Eradication

Currently, there are proposals to eradicate Anopheles gambiae, the main vector for malaria, with a
CRISPR-Cas9 gene drive system.[29] This system aims to eradicate the species through introducing a
gene that would cause female sterility, thus causing the gene to be unable to replicate. It has been
demonstrated in a study by Kyrou et al that such a gene drive system can suppress an entire caged An.
gambiae population through targeting and deleting the dsx gene, which is vital for female fertility.[30]
By utilizing the conservation tendencies of selfish genes, Kyrou et al demonstrated full suppression of
the population within 7–11 generations, typically less than a year. Of course, this has raised concerns
with both the efficiency of a gene drive system as well as the ethical and ecological impact of such an
eradication program.[31] Therefore, there have been efforts to use the gene drive system to more
efficiently introduce genes of Plasmodium resistance into the species, such as targeting and knocking
out the FREP1 gene in Anopheles gambiae.[32] Researchers in Burkina Faso have created a strain of the
fungus metarhizium pinghaense genetically engineered to produce the venom of an Australian funnel-
web spider; exposure to the fungus caused populations of Anopheles mosquitoes to crash by 99% in a
controlled trial.[33]

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Aedes aegypti, the yellow fever mosquito, is a mosquito that can spread dengue fever, chikungunya,
Zika fever, Mayaro and yellow fever viruses, and other disease agents. The mosquito can be recognized
by black and white markings on its legs and a marking in the form of a lyre on the upper surface of its
thorax. This mosquito originated in Africa, but is now found in tropical, subtropical and temperate
regions throughout the world.

SPECIES OF ANOPHOLES MOSQUITO

Some important species are:

A. atroparvus

A. albimanus

A. arabiensis

A. barberi

A. bellator

A. crucians

A. cruzii

A. culicifacies

A. darlingi

A. dirus

A. earlei

A. freeborni

A. funestus

A. gambiae (Giles 1902)

A. introlatus

A. latens
A. maculipennis

A. moucheti

A. nili

A. punctipennis

A. quadrimaculatus

A. stephensi

A. subpictus

A. sundaicus

A. walkeri

ANOPHOLES ATROPARVUS

Anopheles atroparvus

Anopheles atroparvus is a European species of mosquito, which was first identified in 1927. It is one of
the most abundant palearctic Anopheles species belonging to the family Culicidae, commonly called
mosquitoes. Although research interest in A. atroparvus has been low in the past several decades,
recent concern for an increase in vector-borne disease has encouraged new research into this species.

Anopheles atroparvus

Scientific classification

Domain: Eukaryota

Kingdom: Animalia

Phylum: Arthropoda

Class: Insecta

Order: Diptera

Family: Culicidae

Genus: Anopheles

Species: A. atroparvus

Binomial name
Anopheles atroparvus

A. atroparvus underwent population reduction in many areas where it was once abundant due to
pollution of natural habitats, vector control measures, and the reduction of suitable habitat through
drying or flooding.[citation needed] It is the natural prey to many fish and insect species, and preys upon
mainly humans, birds and livestock associated with farmland and farm dwellings.

Taxonomy

A. atroparvus belongs to the subgenre Anopheles among a group defined by the maculipennis complex
[1][2][3] Differentiation between Anopheles species is often not possible at a phenotypic level but
instead is observed in behavior, such as differences in habitats or hibernation habits,[3] or variation at a
genetic level.[4][3][2]

Distribution and habitat

Vastly distributed throughout most of Europe, specifically northern regions, with little or no known
populations in Southern and South eastern Europe.[5][6] A. atroparvus is currently assumed not to live
in Italy, Greece or Turkey.[6][5][7] Distribution of A. atroparvus is affected in some areas due to
competition with related species.[6] Other constraints on distribution depends on habitat suitability; A.
atroparvus is limited by the need for water and vegetation to reproduce and minimize the risk of
predation.[8] Because A. atroparvus tends to like warmer climate,[9] global climate change could result
in changing habitat distribution for A. atroparvus and it is generally considered to result in an increase in
population density of the species.[6][10]

Habitat

A. atroparvus is common to both coastal and inland areas where short lived still water bodies are
plentiful.[5][9] Adults live in close association with the animal and human blood hosts that it feeds on
and tend to live in farm homes and other dwellings.[5][10][8] Suitable habitat for larval stages is mainly
in wide, shallow non-permanent water bodies of brackish water, commonly in association with irrigated
fields especially crops such as rice.[5][8] Unlike many other species, A. atroparvus remains active in the
winter months, at which time it will still take blood meals, but will not lay eggs.[3][9] Their activity in the
winter may result in an annual reduction in population as competition for food and shelter rises.[2]A.
atroparvus displays genetic diversity for protection against common insecticides used on crops, making
it easier for them to live on farmland.[11][10]

Life cycle

Adult A. atroparvus can be active feeders all year round, but cannot lay eggs in the winter and tend to
dwell inside through the winter months.[8][9] They are most active during the summer months (June-
July).[8] During the warmer months, eggs are laid in suitable water bodies, where they will hatch and
larva will remain in the water until they develop into adults.[8][5][12] Females have been recorded to
have a short life span.[13]
Larval stages require water for development and it is not until they become adults that they can survive
a non-aquatic environment.[5] This species shows some adaptability when recent observations have
recorded larvae present in small water bodies within towns such as overgrown roadsides, which were
not considered suitable in the past.[2]

Parasitology

A. atroparvus is a well known historical vector for malaria (Plasmodium spp) particularly throughout
Europe and the UK,[5][2][12][6] and is considered the main vector for malaria in many countries such as
Romania, Portugal, France and the Netherlands.[12][13][6][10] In Spain, A. atroparvus has been
identified as the main vector for two different malaria parasites (Plasmodium vivax & Plasmodium
falciparum).[5] Although malaria has been considered absent of natural strains in Europe, changing
climate and human caused land conditioning via agriculture or environmental protection measures
could cause an increase in population of A. atroparvus in areas where it used to be a main vector for the
disease.[6][5][4][2] Because A. atroparvus is still active in the winter and tend to live in dwellings with
their blood meals, this can result in multiple infections in the same household.[9]

Control

In the past, measures such as introducing fish and insect species that prey on A. atroparvus to water
bodies has been used to decrease the presence of the malaria vector.[6][8][2][14] Household measures
were also introduced and effectively reduced malaria contraction via reduction in A. atroparvus
populations. this included indoor DDT insecticide sprays and bug nets for windows and doors.[4][2]
Through both intentional control of A. atroparvus and other factors such as pollution to A. atroparvus
breeding grounds in the Netherlands, the decrease of this vector species led to the elimination of
malaria from Europe.[6][4] although primarily research suggests that climate change would serve to
increase A. atroparvus populations, there are also implications that it may instead adversely affect
populations by reducing water bodies and rainfall that are required for reproduction.[2] There is also
concern over vector control as A. atroparvus could become infected with Plasmodium species that have
been imported from areas where malaria is more abundant.[6]

References

Harbach, R. E. (2013). The phylogeny and classification of Anopheles. In Anopheles mosquitoes-new

insights into malaria vectors. IntechOpen.

Takken, W., Geene, R., Adam, W., Jetten, T. H., & van der Velden, J. A. (2002). Distribution and dynamics
of larval populations of Anopheles messeae and A. atroparvus in the delta of the rivers Rhine and
Meuse, The Netherlands. AMBIO: A Journal of the Human Environment, 31(3), 212-218

Talbalaghi, A., & Shaikevich, E. (2011). Molecular approach for identification of mosquito species
(Diptera: Culicidae) in Province of Alessandria, Piedmont, Italy. European Journal of Entomology, 108(1),
35.
 Vicente, J. L., Sousa, C. A., Alten, B., Caglar, S. S., Falcutá, E., Latorre, J. M., ... & Toma, L. (2011). Genetic
and phenotypic variation of the malaria vector Anopheles atroparvus in southern Europe. Malaria
journal, 10(1), 5

Birnberg, L., Aranda, C., Talavera, S., Núñez, A. I., Escosa, R., & Busquets, N. (2020). Laboratory
colonization and maintenance of Anopheles atroparvus from the Ebro Delta, Spain. Parasites & Vectors,
13(1), 1-5

5. Capinha, C., Gomes, E., Reis, E., Rocha, J., Sousa, C. A., Do Rosario, V. E., & Almeida, A. P. (2009).
Present habitat suitability for Anopheles atroparvus (Diptera, Culicidae) and its coincidence with former
malaria areas in mainland Portugal. Geospatial health, 177-187.

> "Anopheles atroparvus- Factsheet for experts". European Centre for Disease Prevention and Control.
2014.

Roiz, D., Ruiz, S., Soriguer, R., & Figuerola, J. (2015). Landscape effects on the presence, abundance and
diversity of mosquitoes in Mediterranean wetlands. PLoS One, 10(6), e0128112.

Gowland, R. L., & Western, A. G. (2012). Morbidity in the marshes: Using spatial epidemiology to
investigate skeletal evidence for malaria in Anglo ‐Saxon England (AD 410–1050). American Journal of
Physical Anthropology, 147(2), 301-311.

Ponçon, N., Toty, C., L'Ambert, G., Le Goff, G., Brengues, C., Schaffner, F., & Fontenille, D. (2007).
Biology and dynamics of potential malaria vectors in Southern France. Malaria Journal, 6(1), 18

Hemingway, J. (1982). Genetics of organophosphate and carbamate resistance in Anopheles atroparvus

(Diptera: Culicidae). Journal of economic entomology, 75(6), 1055-1058.

Cailly, P., Balenghien, T., Ezanno, P., Fontenille, D., Toty, C., & Tran, A. (2011). Role of the repartition of
wetland breeding sites on the spatial distribution of Anopheles and Culex, human disease vectors in
Southern France. Parasites & vectors, 4(1), 65

Falcuta, E., Prioteasa, F. L., & Nicolescu, G. (2008). Investigations of the anopheline (Diptera: Culicidae)
fauna from three areas belonging to the Danube Delta Biosphere Reserve in order to evaluate the risk of
malaria re-emergence. Danube Delta National Institute Scientific Annals, 14, 15-20.

Brugman, V. A., Hernández-Triana, L. M., England, M. E., Medlock, J. M., Mertens, P. P., Logan, J. G., ... &
Carpenter, S. (2017). Blood-feeding patterns of native mosquitoes and insights into their potential role
as pathogen vectors in the Thames estuary region of the United Kingdom. Parasites & vectors, 10(1),
163.

ANOPHOLES ALBIMANUS

Anopheles albimanus is a species of mosquito in the order Diptera. It is found in coastal Central and
South America, the Caribbean, and Mexico.[3][4] It is a generalist species and capable of wide
dispersion.[5] A. albimanus is a common malaria vector.[3]
Female Anopheles albimanus

Scientific classification

Domain: Eukaryota

Kingdom: Animalia

Phylum: Arthropoda

Class: Insecta

Order: Diptera

Family: Culicidae

Genus: Anopheles

Species: A. albimanus

Binomial name

Anopheles albimanus

Anatomy

Mosquitoes have long legs and one pair of wings.[2] Females have piercing, sucking mouthparts to
penetrate hosts. Males have a proboscis as well, but are unable to pierce.[2] The vibration of
membranes on the thorax creates the distinguishable whining sound.[2]

Taxonomy and evolution

Anopheles albimanus appears to be a single species that shows polymorphism.[6] It is believed that this
species originated in the Caribbean, moved first to Central America, then South America. Studies suggest
that the occurrence of the founder effect is the cause of the current population structure.[6] Research
using mitochondrial DNA found some modest differences between Central and South American
populations.[7] It has been hypothesized that this could be due to recent pressures, such as the distance
between populations, high insecticide use, or mountain ranges that act as gene flow barriers.[7] Overall,
differences in populations were found to be relatively low. Studies suggest that this is due to these
mosquitoes showing excellent dispersal abilities, opportunistic feeding, and niche plasticity.[7] As well,
other species in the genus Anopheles are specialists with small populations, specific habitat
requirements, and limited dispersal and distribution methods. Alternately, A. albimanus is a generalist
species. Females can fly up to 32 kilometres,[5] and specimens have been found at almost 2000 meters
altitude, meaning that mountain ranges would not separate these populations as easily as other
Anopheles mosquitoes.[7] The conclusions drawn from this research was that the differences found
between A. albimanus populations were not due to recent events, but rather, Pleistocene divergence
followed by re-colonization and expansion.[7]
Distribution and behavior

These mosquitoes are currently found in Mexico, and several countries in the Caribbean, Central
America, and South America.[3][4] They are usually present in coastal areas, at 500 meters altitude or
lower;[3] however, some have been found above 1000 meters.[5] This species occurs in a variety of
habitats in these coastal areas, with regional differences in precipitation, temperature, and vegetation.
This is the most common species found in countries like Colombia.[8]

A. albimanus are seasonally abundant mosquitoes that are primarily active at dawn and dusk
(crepuscular), prefer to feed outdoors (exophagic), and tend to live outside (exophilic).[7] Seasonal
abundance is dependent on the availability of larval habitats.[9] In areas that have great differences in
rainfall depending on the time of year, these mosquitoes are most abundant in the rainy season. In
areas that have wetlands that stay flooded permanently, they are found year-round.[9] Although these
are the typical traits seen, factors such as host availability and control methods can change the
population characteristics.[7]

Feeding by adults

A. albimanus mosquitoes feed on nectar; only females take blood meals.[5][10] Hosts include mammals,
birds, reptiles, and amphibians.[10] A wide range of hosts allows these mosquitoes to maintain their
populations quite easily.[9] The proteins and nutrients gained from blood go toward egg production,
and the volume of the blood meals affects the fecundity of female mosquitoes.[10] The volume taken
during blood feeding is affected by several factors, including temperature, age of the mosquito, mating
status, number of feeds, variety of hosts, and others.[10] There are often interruptions to blood feeding,
resulting in the mosquito taking several meals, either on the same or different hosts. This, in turn,
creates more opportunities for infection or the spread of pathogens.[10]

Life cycle

These mosquitoes undergo holometabolous development, meaning that their life cycle includes egg,
larva, pupa, and adult stages. There are 4 larval instars in A. albimanus.[11] The number and maturation
success of eggs oviposited depends on the quality and quantity of blood taken by the female.[10] Eggs
can be laid one by one, or glued together in a mass.[2] The habitats that females choose to oviposit in
will affect the distribution of larvae. If a habitat is seeing an absence of larvae, this may be due to
females being selective for locations with richer food sources or protection from predation.[9] It could
also be because of environmental conditions that would kill the eggs or larvae.[9] A. albimanus larvae
are able to develop in a wide range of habitats and conditions (i.e. salinity, pollution).[5] During the wet
season in Central America, larvae are found in rivers, marshes, irrigation canals, and several other water
habitats, both permanent and temporary.[3] During the dry season, they are found only in permanent
water habitats.[3] There is a relationship between the presence of A. albimanus larvae and
cyanobacteria mats floating on the water. This is because the presence of blue-green algae results in an
elevated water temperature and less dissolved oxygen. Volatile compounds released from the algae
attract females, increasing the likelihood of oviposition.[3] There is less known about South American
populations, but larvae have been found in rice fields and drains. In Colombia, larvae are found in sunny
habitats, where temperatures are 27 to 30 degrees Celsius.[3] Understanding larval habitats and
distribution are important in developing control methods for these mosquitoes.[9]

Disease

A. albimanus is a medically important species, as it is a key Plasmodium vivax malaria vector; this means
that they carry and transmit an organism that causes a disease in another host organism, and usually are
not harmed themselves.[4][9][10] In countries such as Belize, they are considered a secondary vector
because they prefer not to enter human homes, and rather, feed on domestic animals.[10] However, in
other areas where domestic animals are not abundant, and especially regions where indigenous people
live, malaria is endemic.[10] This is due to large populations of the mosquitoes, as well as poorly built
houses.[10] Females acquire P. vivax during a blood meal from an infected host.[4] The parasite moves
to the midgut and reproduces sexually.[4] The parasite and the blood itself interact with the lining of the
midgut, eliciting stress and digestion responses in the mosquito, as well as immune reactions. The
parasite must survive these responses in order for the egg to fertilize and form an ookinete. The
ookinetes then move into the epithelial lining of the midgut to multiply further, resulting in sporozoites.
[4][12] These sporozoites travel to the salivary glands, and can then be spread to a new host when a
blood meal is taken.[12] These changes in the midgut evolved to cope with the invasion of
microorganisms, free radical formation, and digestion of the blood meal.[4]

Control

Colonies of Anopheles mosquitoes are maintained for several areas of research, such as malaria vector
ability, transmission of pathogens, the effectiveness of insecticides, resistance to insecticides, and
vaccine research.[10] By studying these topics, more insight may be gained to increase the
understanding of malaria, as well as the effects that control measures would have on the reduction of
both the vectors and the spread of the disease.[10]

References

Eduar E. Bejarano (2003). "Occurrence of the malaria vector Anopheles albimanus Wiedemman
(Diptera: Culicidae) in Isla Fuerte, Colombia". Neotropical Entomology. 32 (3): 517–518.
doi:10.1590/S1519-566X2003000300025.

"Mosquito". Columbia Electronic Encyclopedia, 6th Edition. Columbia University Press: 1.

Pinault, L. L.; Hunter, F. F. (June 2012). "Characterization of larval habitats of Anopheles albimanus,
Anopheles pseudopunctipennis, Anopheles punctimacula, and Anopheles oswaldoi s.l. populations in
lowland and highland Ecuador". Journal of Vector Ecology. 37 (1): 124–136. doi:10.1111/j.1948-
7134.2012.00209.x. PMID 22548546.

Cázares-Raga, F. E.; Chávez-Munguía, B.; González-Calixto, C.; Ochoa-Franco, A. P.; Gawinowicz, M. A.;
Rodríguez, M. H.; Hernández-Hernández, F. C. (August 2014). "Morphological and proteomic
characterization of midgut of the malaria vector Anopheles albimanus at early time after a blood
feeding". Journal of Proteomics. 111: 100–112. doi:10.1016/j.jprot.2014.07.037. PMID 25132141.
 Gómez, G. F.; Márquez, E. J.; Gutiérrez, L. A.; Conn, J. E.; Correa, M. M. (April 2, 2014). "Geometric
morphometric analysis of Colombian Anopheles albimanus (Diptera: Culicidae) reveals significant effect
of environmental factors on wing traits and presence of a metapopulation". Acta Tropica. 135: 75–85.
doi:10.1016/j.actatropica.2014.03.020. PMC 4464773. PMID 24704285.

Loaiza, J. R.; Scott, M. E.; Bermingham, E.; Sanjur, O. I.; Wilkerson, R.; Rovira, J.; Gutiérrez, L. A.; Correa,
M. M.; Grijalva, M. J.; Birnberg, L.; Bickersmith, S.; Conn, J. E. (September 2010). "Late Pleistocene
environmental changes lead to unstable demography and population divergence of Anopheles
albimanus in the northern Neotropics". Molecular Phylogenetics and Evolution. 57 (3): 1341–1346.
doi:10.1016/j.ympev.2010.09.016. PMC 3229172. PMID 20888924.

Loaiza, J. R.; Scott, M. E.; Bermingham, E.; Rovira, J.; Conn, J. E. (January 2010). "Evidence for
Pleistocene population divergence and expansion of Anopheles albimanus in Southern Central America".
The American Journal of Tropical Medicine and Hygiene. 82 (1): 156–164. doi:10.4269/ajtmh.2010.09-
0423. PMC 2803528. PMID 20065014.

Gutiérrez, L. A.; Naranjo, N. J.; Cienfuegos, A. V.; Muskus, C. E.; Luckhart, S.; Conn, J. E.; Correa, M. M.
(November 2009). "Population structure analyses and demographic history of the malaria vector
Anopheles albimanus from the Caribbean and the Pacific regions of Colombia". Malaria Journal. 8: 259.
doi:10.1186/1475-2875-8-259. PMC 2789746. PMID 19922672.

Rejmánkova, E.; Roberts, D. R.; Manguin, S.; Pope, K. O.; Komárek, J.; Post, R. A. (October 1996).
"Anopheles albimanus (Diptera: Culicidae) and cyanobacteria: An example of larval habitat selection".
Population Ecology. 25 (5): 1058–1067. doi:10.1093/ee/25.5.1058. PMID 11540613.

Phasomkusolsil, S.; Pantuwattana, K.; Tawong, J.; Khongtak, W.; Kertmanee, Y.; Monkanna, N.; Klein, T.
A.; Kim, H. C.; McCardle, P. W. (September 2015). "The relationship between wing length, blood meal
volume, and fecundity for seven colonies of Anopheles species housed at the Armed Forces Research
Institute of Medical Sciences, Bangkok, Thailand". Acta Tropica. 152: 220–227.
doi:10.1016/j.actatropica.2015.09.021. PMID 26433074.

Komínkova, D.; Rejmánkova, E.; Grieco, J.; Achee, N. (December 2012). "Fatty acids in anopheline
mosquito larvae and their habitats". Journal of Vector Ecology. 37 (2): 382–395. doi:10.1111/j.1948-
7134.2012.00242.x. PMID 23181863.

Serrano-Pinto, V.; Acosta-Pérez, M.; Luviano-Bazán, D.; Hurtado-Sil, G.; Batista, C. V. F.; Martínez-
Barnetche, J.; Lánz-Mendoza, H. (June 2010). "Differential expression of proteins in the midgut of
Anopheles albimanus infected with Plasmodium berghei". Insect Biochemistry and Molecular Biology. 40
(10): 752–758. doi:10.1016/j.ibmb.2010.07.011. PMID 20692341.

Anopheles arabiensis[1] is a zoophilic species of mosquito and a vector of disease endemic to Africa.

Anopheles arabiensis

Domain: Eukaryota
Kingdom: Animalia

Phylum: Arthropoda

Class: Insecta

Order: Diptera

Family: Culicidae

Genus: Anopheles

Species: A. arabiensis

Binomial name

Anopheles arabiensis

Polytene chromosomes have a high degree of gene polymorphism due to paracentric inversions. This is
also unusually high for the genus. (See the chapter by Kitzmiller 1976.)[2] There is a well studied
adaptive inversion. Kirkpatrick and Barrett 2015 and Sharakhov et al. 2006 find an inversion providing A.
arabiensis with some of its adaptation to arid environments. They also find this inversion has been
introgressed across more widely in the genus, providing similar adaptive benefit.[3]

Symbionts

Not thought to naturally serve as a host of Wolbachia[4] until Baldini et al. 2018 showed to the contrary.
[5][6]

Hosts

Hosts include Bos taurus.[7] A. arabiensis is especially known as a zoophilic haematophage.[8]

Parasites

Not a vector of Plasmodium berghei.[9]

Range

The distribution is Afrotropical. There was a brief invasion into Brazil in 1930 but this was quickly
eradicated. (Note that this was long misidentified as an invasion by A. gambiae. Only with genetic tools
and a great deal of time did Parmekalis et al. 2008 find it to really have been A. arabiensis.)[10] The
investigations regarding the ecology of A. arabiensis by Gwitira et al. 2018, Ageep et al. 2009 and Fuller
et al. 2012a help to model the distribution of various avian malaria pathogens.[11]

Locally A. arabiensis' is especially known as an exophage and exophile.[8] Its movements through the
local ecology are not sufficiently studied – Debebe et al. 2018 is one of the few investigations in this
question.[12]
Control

Cyhalothrins (including λ-cyhalothrin) and DDT are commonly used. Mnzava et al. 1995 finds differential
repellent effects between λc and DDT in the protection of cattle, partly due to DDT's excitorepellency.
(Some of the difference is also due to differences in keeping cattle outside or inside. A. arabiensis'
proclivity to enter or not enter, and exit or not exit barns treated with λc or DDT makes a difference.)[7]

Sterile insect technique shows promise in A. arabiensis. Irradiation in SIT is not simple however and
dosage is a touchy variable. Sterile males are also injured more generally by the process and thus are
less competitive. Helinski and Knols 2008 provide dosage information from their experiments with A.
arabiensis which is needed to perform SIT successfully.[13]

This process requires separation of the sexes which historically has been done manually, greatly limiting
throughput. Mashatola et al. 2018[14] reviews progress in automation, selective insecticide feeding, and
genetic sexing strains.[13]

SIT may also be achieved by genetic modification, disabling the reproductive process. Catteruccia et al.
2005 produced such an A. arabiensis strain and demonstrates more generally that genetic SIT is
tractable in this species.[4]

As of 2015 it has only recently been found that adult mosquitoes are vulnerable to entomopathogenic
fungi. This has provoked interest in studying this kind of control, especially Kikankie et al. 2010's success
with Beauveria bassiana.[15]

Understanding of A. arabiensis' movements through the landscape will need to improve to aid control
efforts. Debebe et al. 2018 is one of very few contributions to this area.[12]

Insecticide resistance

Some resistant A. arabiensis populations are known. Ismail et al. 2018 find a high degree of pyrethroid
resistance in Sudan and Opondo et al. 2019 find the same in The Gambia.[16] Hargreaves et al. 2003
finds DDT resistance in South Africa severe enough to impact efficacy.[17] Agricultural runoff encourages
DDT resistance: A. arabiensis larvae grow in waste water pools nearby and are encouraged toward
resistance by the insecticides applied to the crops. Oliver and Brooke 2013 find this to be especially
problematic adjacent to maize cultivation.[18]

References

"Anopheles arabiensis". Catalogue of Life. Species 2000: Leiden, the Netherlands.

Krzywinski, Jaroslaw; Besansky, Nora J. (2003). "Molecular Systematics of Anopheles: From Subgenera
to Subpopulations". Annual Review of Entomology. 48 (1). Annual Reviews: 111–139.
doi:10.1146/annurev.ento.48.091801.112647. ISSN 0066-4170. PMID 12208816.
 Angert, Amy L.; Bontrager, Megan G.; Ågren, Jon (2020-11-02). "What Do We Really Know About
Adaptation at Range Edges?". Annual Review of Ecology, Evolution, and Systematics. 51 (1). Annual
Reviews: 341–361. doi:10.1146/annurev-ecolsys-012120-091002. ISSN 1543-592X. S2CID 228833318.

McGraw, Elizabeth A.; O'Neill, Scott L. (2013-02-15). "Beyond insecticides: new thinking on an ancient
problem". Nature Reviews Microbiology. 11 (3). Nature Portfolio: 181–193. doi:10.1038/nrmicro2968.
ISSN 1740-1526. PMID 23411863. S2CID 37101176.

Caragata, Eric P.; Tikhe, Chinmay V.; Dimopoulos, George (2021-06-02). "Curious Entanglements:
Interactions between Mosquitoes, their Microbiota, and Arboviruses". Current Opinion in Virology. 37.
Elsevier: 26–36. doi:10.1016/j.coviro.2019.05.005. PMC 6768729. PMID 31176069. NIHMS 1531069.

Sicard, Mathieu; Bonneau, Manon; Weill, Mylène (2019). "Wolbachia prevalence, diversity, and ability
to induce cytoplasmic incompatibility in mosquitoes" (PDF). Current Opinion in Insect Science. 34.
Elsevier: 12–20. doi:10.1016/j.cois.2019.02.005. ISSN 2214-5745. PMID 31247412. S2CID 91796037.

Pates, Helen; Curtis, Christopher (2005-01-01). "Mosquito Behavior and Vector Control". Annual Review
of Entomology. 50 (1). Annual Reviews: 53–70. doi:10.1146/annurev.ento.50.071803.130439. ISSN
0066-4170. PMID 15355233.

"Anopheles (Cellia) arabiensis Patton, 1905". Malaria Atlas. 2018-02-15. Retrieved 2022-01-23.

Sinden, Robert E.; Butcher, Geoff A.; Beetsma, A. L. (2002). "Maintenance of the Plasmodium berghei
Life Cycle". Malaria Methods and Protocols. Methods in Molecular Medicine. Vol. 72. New Jersey:
Humana Press. pp. 25–40. doi:10.1385/1-59259-271-6:25. ISBN 1-59259-271-6. PMID 12125122.

Molina-Cruz, Alvaro; Zilversmit, Martine M.; Neafsey, Daniel E.; Hartl, Daniel L.; Barillas-Mury, Carolina
(2016-11-23). "Mosquito". Annual Review of Genetics. 50 (1). Annual Reviews: 447–465.
doi:10.1146/annurev-genet-120215-035211. ISSN 0066-4197. PMID 27732796.

Santiago-Alarcon, Diego; Alfonso, Marzal, eds. (2020). Avian Malaria and Related Parasites in the
Tropics : Ecology, Evolution and Systematics. Cham, Switzerland: Springer. pp. xiv+575. ISBN 978-3-030-
51632-1. OCLC 1204140762. ISBN 978-3-030-51633-8.

Ignell, Rickard; Hill, Sharon Rose (2020). "Malaria mosquito chemical ecology". Current Opinion in Insect
Science. 40. Elsevier: 6–10. doi:10.1016/j.cois.2020.03.008. ISSN 2214-5745. PMID 32422588. S2CID
216194693.

Caragata, E.P.; Dong, S.; Dong, Y.; Simões, M.L.; Tikhe, C.V.; Dimopoulos, G. (2020-09-08). "Prospects
and Pitfalls: Next-Generation Tools to Control Mosquito-Transmitted Disease". Annual Review of
Microbiology. 74 (1). Annual Reviews: 455–475. doi:10.1146/annurev-micro-011320-025557. ISSN 0066-
4227. PMID 32905752. S2CID 221625690.

Mashatola, Thabo; Ndo, Cyrille; Koekemoer, Lizette L.; Dandalo, Leonard C.; Wood, Oliver R.;
Malakoane, Lerato; Poumachu, Yacouba; Lobb, Leanne N.; Kaiser, Maria; Bourtzis, Kostas; Munhenga,
Givemore (2018). "A review on the progress of sex-separation techniques for sterile insect technique
applications against Anopheles arabiensis ". Parasites & Vectors. 11 (S2). BioMed Central: 127–171.
doi:10.1186/s13071-018-3219-4. ISSN 1756-3305. PMC 6304763. PMID 30583746.

Lacey, L.A.; Grzywacz, D.; Shapiro-Ilan, D.I.; Frutos, R.; Brownbridge, M.; Goettel, M.S. (2015). "Insect
pathogens as biological control agents: Back to the future" (PDF). Journal of Invertebrate Pathology. 132.
Society for Invertebrate Pathology (AP): 1–41. doi:10.1016/j.jip.2015.07.009. ISSN 0022-2011. PMID
26225455.

Jeran, Nina; Grdiša, Martina; Varga, Filip; Šatović, Zlatko; Liber, Zlatko; Dabić, Dario; Biošić, Martina
(2020-10-06). "Pyrethrin from Dalmatian pyrethrum (Tanacetum cinerariifolium/Trevir./Sch. Bip.):
biosynthesis, biological activity, methods of extraction and determination". Phytochemistry Reviews. 20
(5). Phytochemical Society of Europe + Phytochemical Society of North America (Springer): 875–905.
doi:10.1007/s11101-020-09724-2. ISSN 1568-7767. S2CID 225152789. (MG ORCID: 0000-0002-4584-
4851).

Thomas, Matthew B.; Read, Andrew F. (2007-04-11). "Can fungal biopesticides control malaria?".
Nature Reviews Microbiology. 5 (5). Nature Portfolio: 377–383. doi:10.1038/nrmicro1638.
hdl:1842/2089. ISSN 1740-1526. PMID 17426726. S2CID 14460348.

Williams, Adrian C; Hill, Lisa J (2019). "Nicotinamide and Demographic and Disease transitions:
Moderation is Best". International Journal of Tryptophan Research. 12. Sage: 117864691985594.
doi:10.1177/1178646919855940. ISSN 1178-6469. PMC 6610439. PMID 31320805.

ANOPHOLES BARBERI

Anopheles barberi is a species of tree hole–breeding mosquito from eastern North America.[1] The
larvae are predators of other mosquito larvae, such as those of Aedes triseriatus.[2] It has been shown
to be a vector of malaria in the laboratory, but it is not thought to be an important malaria vector in the
wild.[3][additional citation(s) needed]

Anopheles barberi

Domain: Eukaryota

Kingdom: Animalia

Phylum: Arthropoda

Class: Insecta

Order: Diptera

Family: Culicidae

Genus: Anopheles
Species: A. barberi

Binomial name

Anopheles barberi

References

DeVries, Peter J. (16 July 2011). "Anopheles barberi Coquillett 1903". GeoSpecies Knowledge Base.
University of Wisconsin - Madison Department of Entomology. Archived from the original on 26 July
2011.

Miner, Angela (2014). Martina, Leila Siciliano (ed.). "Aedes triseriatus". Animal Diversity Web. Retrieved
24 January 2024.

"Anopheles barberi". Medically Important Mosquitoes. Suitland, Maryland: The Walter Reed
Biosystematics Unit. Archived from the original on 20 November 2008.

ANOPHOLES BELLATOR

Anopheles bellator is a species of mosquito mainly located in southeast of State of São Paulo, Brazil, is a
main vector of malaria.[1][2][3]

References

edit

"Anopheles bellator Dyar & Knab, 1906 | Walter Reed Biosystematics Unit (WRBU)". wrbu.si.edu.
Retrieved 11 March 2022.

Forattini, Oswaldo Paulo; Kakitani, Iná; Santos, Roseli La Corte dos; Ueno, Helene Mariko; Kobayashi,
Keilla Miki (1999–2011). "Role of Anopheles (Kerteszia) bellator as malaria vector in Southeastern Brazil
(Diptera: Culicidae)". Memórias do Instituto Oswaldo Cruz. 94: 715–718. doi:10.1590/S0074-
02761999000600001. ISSN 0074-0276.

Pittendrigh, Colin S. (1950). "The Ecoclimatic Divergence of Anopheles bellator and A. homunculus".
Evolution. 4 (1): 43–63. doi:10.2307/2405533. ISSN 0014-3820.

Anopheles bellator

Scientific classificationEdit this classification

Domain:

Eukaryota

Kingdom:
Animalia

Phylum:

Arthropoda

Class:

Insecta

Order:

Diptera

Family:

Culicidae

Genus:

Anopheles

Species:

A. bellator

Binomial name

Anopheles bellator

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Anopheles crucians

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Anopheles crucians is a mosquito that exists in aquatic environments under areas with little light
presence. The preferred environment for A. crucians is areas with acidic water such as that found in
cypress swamps.[1] The mosquito breeds in semipermanent and permanent pools, ponds, lakes and
swamps.

Anopheles crucians
Scientific classificationEdit this classification

Domain:

Eukaryota

Kingdom:

Animalia

Phylum:

Arthropoda

Class:

Insecta

Order:

Diptera

Family:

Culicidae

Genus:

Anopheles

Species:

A. crucians

Binomial name

Anopheles crucians

Wiedemann, C.R., 1828

It may be a vector for malaria.

Anatomy

edit

The anatomy of A. crucians is very similar to that of Anopheles bradleyi. The proboscis is dark colored
and black like that of other mosquitoes. The pedipalps, composed of six segments, are different colors
based on the segment. The basal part is black with raised scales, segment 3 has white scales, the 4th
segment has white-ringed basally and apically, and the last segment is white. The abdomen ranges from
dark brown to black with hundreds of yellow to dark-brown hairs. The legs are dark with white patched
in the femora and tibiae. The wings, on average, are 4.0 millimeters in length and will have white-yellow
scales with contrasting spots.[1]

Malaria vector

edit

In research conducted by W.V. King in 1916, it was discovered that Anopheles female mosquitoes are
carriers of malaria. In this research, 75% of A. crucians individuals were infected with oocysts,
sporozoites, or both of the malarial parasite Plasmodium falciparum.[2]

References

edit

Carpenter, Stanley J.; LaCasse, Walter J. (1974). Mosquitoes of North America (North of Mexico).
University of California Press. ISBN 0520026381.

King, W.V (1916). "Experiments on the development of malaria parasites in three American species of
Anopheles". Journal of Experimental Medicine. 23 (6): 703–716. doi:10.1084/jem.23.6.703. PMC
2125450. PMID 19868018.

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Anopheles cruzii

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This article relies largely or entirely on a single source. (March 2022)

Anopheles cruzii, is the species of mosquito that mainly located in southern coast of Brazil, is main
vector of malaria, Plasmodium vivax.[1]

Anopheles cruzii

Scientific classificationEdit this classification

Domain:
Eukaryota

Kingdom:

Animalia

Phylum:

Arthropoda

Class:

Insecta

Order:

Diptera

Family:

Culicidae

Genus:

Anopheles

Species:

A. cruzii

Binomial name

Anopheles cruzii

Dyar and Knab, 1908

It has been going through microevolution, which appears in its wing-shape and is correlated with
urbanization.[2]

References

"ITIS standard report - Anopheles cruzii - Dyar and Knab, 1908". Integrated Taxonomic Information
System. Retrieved 11 March 2022.

Multini, Laura Cristina; Wilke, André Barretto Bruno; Marrelli, Mauro Toledo (2019-02-01).
"Urbanization as a driver for temporal wing-shape variation in Anopheles cruzii (Diptera: Culicidae)".
Acta Tropica. 190: 30–36. doi:10.1016/j.actatropica.2018.10.009. ISSN 0001-706X.

.
Wikipedia

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Anopheles culicifacies

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Anopheles culicifacies is a mosquito species complex and one of the primary vectors of malaria on the
Indian subcontinent.[1] It consists of five sibling species, provisionally designated as species A, B, C, D,
and E.[2] It prefers to rest indoors in cattle sheds, where it feeds on cattle. The control of A. culicifacies
has become difficult due to the development of insecticide resistance against all commonly used
insecticides, including new-generation insecticides such as synthetic pyrethroids.[3]

Anopheles culicifacies

Scientific classificationEdit this classification

Domain:

Eukaryota

Kingdom:

Animalia

Phylum:

Arthropoda

Class:

Insecta

Order:

Diptera

Family:

Culicidae

Genus:
Anopheles

Species:

A. culicifacies

Binomial name

Anopheles culicifacies

Giles, 1901

Phylogeny

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The Anopheles genus, commonly known as mosquitos comprises almost 500 species, including several
species complexes. Taxonomic classification of these mosquitos relies upon traditional morphology,
chromosome analysis, and DNA markers.[4] Anopheles is one of the most studied genus of the Culicidae
family. As a major vector of diseases such as malaria and filariasis, species belonging to this genus have
affected humans more than any other insects.[5]

Species Complex

edit

Species complex is defined as groups of closely related species. Anopheles culicifacies complex consists
of five sibling species, designated from A to E. Were identified on the basis of paracentric inversions on
polytene chromosome X and 2.[6] These polytene chromosome can be read through analysis of ovarian
cells.[7] Through the use of biochemical techniques based on lactate dehydrogenase allozyme (Idh)
differentiations can be made for the sibling species.[6] The five sibling species are reproductively
isolated and have premating barriers but also have postmating barriers. Crosses between species A and
B and A and C produce fertile females and infertile male offspring. The mating of species B and C can
produce fertile progeny.[7] All five of the sibling species have different distribution patterns across India,
with some regions where they overlap.[citation needed]

Life stages

edit

The species breed near bodies of water during monsoon season[6] Females lay between 50-200 eggs on
the surface of water bodies per oviposition. These eggs hatch within 2–3 days.[8]

Larvae are born with no legs but well-developed head and mouth for feeding. They also don't have a
respiratory siphon, so they must keep their body positioned parallel to the water.[8] They breathe
through spiracles located on the 8th abdominal segment.[8]
The pupae are the transitional stage between larvae and adults; at this stage, there is no feeding, only
radical morphogenesis. Anopheles species can develop from egg to adult in as little as seven days but
usually take 10-14, depending on the environmental conditions.[8]

Distribution Patterns

edit

Anopheles culicifacies is found throughout the Indian subcontinent. The species complex is also found to
a much smaller extent to the east in Yemen, Iran, Afghanistan, Pakistan, Burma, Thailand, Laos, and
Vietnam. also, to the north of India, the species can be found in Nepal and southern China.[7] The
species complex is widely distributed in rural and peri-urban India. An. culicifacies is the most widely
distributed species of mosquito in India and occurs in all mainland areas, including Kashmir and high-
elevation regions of northern India.[9] The An.culicifacies species complex thrives in flat plains that
receive fair amounts of rainfall. Populations have been observed at up to 3000m in altitude.[7] The
emergence of heavy rainfall, droughts, floods, and varying cold and hot weather in many areas of India,
An. culicifacies population has begun to see a change in distribution.[9]

Species B of the An. culicifacies species complex is the most predominant and widespread throughout
India. Species A and B are found to be sympartic in north and south India, with predominance of species
A in the north and species B in the south.[9] In eastern Uttar Pradesh and northeastern states, species B
is seen to be the only species present. In the western and eastern regions, species B and C are
predominant. Species D is observed to be sympatric with species A and B in the northwestern region and
with species A, B, and C in the central region and a few areas in southern India.[9] Species E is sympatric
with species B in southern Tamil Nadu and Sri Lanka. The distribution and proportion of sibling species is
subject to change through varying seasons. For example, in Delhi, where species A and B are sympatric,
species A is predominant throughout the year, but proportions of species B increased during post-
monsoon months.[9]

Vector of Disease

edit

The An. culicifacies species are responsible for 65-75% of malaria cases in India alone.[10] In Madhya
Pradesh 8% of the population contribute to 30% of the total malaria cases in central India, due to the
large concentration of An. culicifacies species in central India.[11] A study conducted compared two
species of mosquitoes An. subpictus and An. culicifacies and found that the An. culicifices species had a
higher sporozoite rate of 1.8% as 13 of 716 collected insects were positive for the malaria sporozoite.
[12] An. culicifacies are generally zoophilic, meaning they feed on animals, and this species has an
affinity for cattle. The species also has a low affinity for human flesh, but due to their high density, they
are a major vector for malaria.[6] The species prefers to breed near streams, rice fields, irrigation
channels, and rainwater collections. Because of this vast breeding range occupied by this species during
monsoon season, it is difficult to employ anti-larval control methods.[6] The sibling species differ in their
susceptibility to malaria sporogony; species A is known to be highly susceptible, and it is followed by
species C and B. Species B has a unique ability that allows it to kill early sporogony in its midgut through
cellular encapsulation. The control of An. culicifacies has been a major concern for vectorial control
programs in India due to the evolutions of resistance to commonly used insecticides such as DDT.[6]

Resistance

edit

During the early use of Dichlorodiphenyltrichloroethane (DDT), it was very effective against populations
of An. culicifacies when applied as residual treatment. Due to the over and improper use of DDT An.
culicifacies resistance began to emerge in several regions of India.[7] Resistance to DDT led to the use of
Hexachlorocyclohexane (HCH) and dieldrin which similarly saw resistance in An. culicifacies. This again
led to using another insecticide called malathion, and similarly, resistance evolved. As resistance
merged, large outbreaks began to occur. Today, in most parts of India An. culicifacies is resistant to DDT
and HCH, and in some states, like Gujarat and Orissa, they are also resistant to malathion and most
commonly used insecticides.[7]

References

edit

"Anopheles culicifacies". Walter Reed Biosystematics Unit (WRBU). Smithsonian Institution. 2008-11-18.
Archived from the original on 18 November 2008. Retrieved 2022-06-14.

Barik TK, Sahu B, Swain V (2009). "A review on Anopheles culicifacies: from bionomics to control with
special reference to Indian subcontinent". Acta Tropica. 109 (2): 87–97.
doi:10.1016/j.actatropica.2008.09.017. PMID 19000647.

Kareemi TI, Nirankar JK, Mishra AK, Chand SK, Chand G, Vishwakarma AK, et al. (2021). "Population
Dynamics and Insecticide Susceptibility of Anopheles culicifacies in Malaria Endemic Districts of
Chhattisgarh, India". Insects. 12 (4): 284. doi:10.3390/insects12040284. PMC 8064500. PMID 33806071.

Trent, R.J. (2005). Molecular Medicine (3rd ed.). Academic press. pp. 193–220. doi:10.1016/B978-0-12-
699057-7.X5000-0. ISBN 978-0-12-699057-7.

Harbach, R.E. (2013). Anopheles Mosquitoes New Insights into Malaria Vectors (1st ed.). Intech. pp. 3–
55. doi:10.5772/54695. ISBN 978-953-51-1188-7.

Dash, A. P.; Adak, T.; Raghavendra, K.; Singh, O.P. (2007). "The Biology and Control of Malaria Vectors in
India". Current Science. 92 (11): 1571–1578.

Subbarao, S.K. (1988). "The Anopheles culicifacies Complex and Control of Malaria". Parasitology Today.
4 (3): 72–75.

CDC (2020). "About Malaria". Centres for Disease Control and Prevention. Retrieved December 3, 2023.
 Sharma, V.P.; Dev, V. (2015). "Biology & Control of Anopheles culicifacies Giles 1901". The Indian
Journal of Medical Research. 141 (5): 525–536.

Goswami, G.; Singh, O.P.; Nanda, N.; Raghavendra, K.; Gakhar, S.K.; Subbarao, S.K. (2006).
"Identification of all members of the Anopheles culicifacies complex using allele-specific polymerase
chain reaction assays". The American Journal of Tropical Medicine and Hygiene. 75 (3): 454–460.

Sharma, A.K.; Tyagi, V.; Singh, S.; Veer, V.; Agrawal, P.O.; Sukumaran, D. (2014). "Distribution of
Anopheles culicifacies and Detection of its Sibling Species E from Madhya Pradesh: Central India".
Journal of Arthropod-Borne Diseases. 8 (2): 186–196.

Kumari, S.; Parida, S.K.; Marai, N.; Tripathy, A.; Hazra, R.K.; Kar, S.K.; Mahapatra, N. (2009). "Vectorial
role of Anopheles subpictus Grassi and anopheles culicifacies Giles in Angul District, Orissa, India".
Southeast Asian Journal of Tropical Medicine and Public Health. 40 (4): 713–719.

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Anopheles darlingi

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Anopheles darlingi, the American malaria mosquito, is a species of mosquito in the family Culicidae.[1]
[2] A. darlingi is one of the major species of mosquito known to be responsible for malaria in the
Amazonian regions.[3][4][5] It has a wide range of geographic distribution that stretches from Mexico
and Argentina but it has also been found to populate in areas affected by deforestation and
environment changes due to humans.[6]

Anopheles darlingi

Anopheles darlingi close up

Scientific classificationEdit this classification

Domain:

Eukaryota

Kingdom:

Animalia
Phylum:

Arthropoda

Class:

Insecta

Order:

Diptera

Family:

Culicidae

Genus:

Anopheles

Species:

A. darlingi

Binomial name

Anopheles darlingi

Root, 1926

Breeding

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The breeding patterns of A. darlingi are affected heavily by the precipitation seen in their environment.
Due to the fact that mosquitoes rely on the surfaces of stagnant water or areas with regular flooding as
breeding grounds for their eggs and larvae,[5] the annual density of the A. darlingi population is
dependent upon the availability of larval habitats.[3] During the wet season, riverine areas are often
abundant with mosquitoes and densities are higher in these after the peak of rainfall.[7] Oppositely, in
drier inland areas that are usually further from rivers and coexisting with the affects human disturbance,
a peak of mosquito density may occasionally occur in the dry season in areas restricted to man-made
dams and stagnant puddles.[7]

Habitat

edit
In its native range A. darlingi larvae are found by Manguin et al 1996 and Rozendaal 1992 to inhabit
littoral, riverine, and backwater bodies. In its invasive Iquitos range Lounibos and Conn 2000 also find
them in aquacultural ponds.[8]

Range

edit

A. darlingi was absent from Iquitos, Peru until shortly after 1991.[8]

Insecticide resistance

edit

Marinotti et al 2013 provides the first whole genome sequence. This has greatly accelerated
understanding of insecticide resistance in A. darlingi. Among other discoveries, Marinotti finds 20
esterases, 30 glutathione S-transferases, and 89 P450s.[9]

References

edit

"Anopheles darlingi Report". Integrated Taxonomic Information System. Retrieved 2020-01-22.

"Anopheles darlingi". GBIF. Retrieved 2020-01-22.

Hiwat, Hélène; Bretas, Gustavo (2011-09-16). "Ecology of Anopheles darlingi Root with respect to vector
importance: a review". Parasites & Vectors. 4 (1). doi:10.1186/1756-3305-4-177. ISSN 1756-3305. PMC
3183005.

World Health Organization. Global Malaria Programme. (2013). World malaria report 2013. World
Health Organization. ISBN 978-92-4-156469-4. OCLC 880833659.

Angêlla, Aline F; Salgueiro, Patrícia; Gil, Luiz HS; Vicente, José L; Pinto, João; Ribolla, Paulo EM (2014).
"Seasonal genetic partitioning in the neotropical malaria vector, Anopheles darlingi". Malaria Journal. 13
(1): 203. doi:10.1186/1475-2875-13-203. ISSN 1475-2875. PMC 4059831.

VITTOR, AMY YOMIKO; GILMAN, ROBERT H.; TIELSCH, JAMES; GLASS, GREGORY; SHIELDS, TIM; LOZANO,
WAGNER SÁNCHEZ; PINEDO-CANCINO, VIVIANA; PATZ, JONATHAN A. (2006-01-01). "THE EFFECT OF
DEFORESTATION ON THE HUMAN-BITING RATE OF ANOPHELES DARLINGI, THE PRIMARY VECTOR OF
FALCIPARUM MALARIA IN THE PERUVIAN AMAZON". The American Journal of Tropical Medicine and
Hygiene. 74 (1): 3–11. doi:10.4269/ajtmh.2006.74.3. ISSN 0002-9637.

Gil, Luís H. Soares; Alves, Fabiana P.; Zieler, Helge; Salcedo, Juan M. V.; Durlacher, Rui R.; Cunha,
Roberto P. A.; Tada, Mauro S.; Camargo, Luiz M. A.; Camargo, Erney P.; Pereira-Da-Silva, Luiz H. (2003-
09-01). "Seasonal Malaria Transmission and Variation of Anopheline Density in Two Distinct Endemic
Areas in Brazilian Amazônia". Journal of Medical Entomology. 40 (5): 636–641. doi:10.1603/0022-2585-
40.5.636. ISSN 0022-2585.

Lounibos, L. Philip (2002). "Insect". Annual Review of Entomology. 47. Annual Reviews: 233–66.
doi:10.1146/ANNUREV.ENTO.47.091201.145206. PMID 11729075. S2CID 28336540.

Liu, Nannan (2015). "Insecticide Resistance in Mosquitoes: Impact, Mechanisms, and Research
Directions". Annual Review of Entomology. 60. Annual Reviews: 537–559. doi:10.1146/annurev-ento-
010814-020828.

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Anopheles dirus

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Anopheles dirus is a vector of malaria in Asian forested zones.[2]

Anopheles dirus

Scientific classificationEdit this classification

Domain:

Eukaryota

Kingdom:

Animalia

Phylum:

Arthropoda

Class:

Insecta

Order:

Diptera
Family:

Culicidae

Genus:

Anopheles

Species subgroup:

Leucosphyrus subgroup

Species complex:

Dirus species complex

Peyton & Harrison, 1979

Species[1]

Anopheles dirus s.s. (species A)

Anopheles crascens (species B)

Anopheles scanloni (species C)

Anopheles baimaii (species D)

Anopheles elegans (species E)

Anopheles nemophilous (species F)

Anopheles takasagoensis

range map for Anopheles spp.. Anopheles dirus is in cyan in Southeast Asia

Taxonomy

edit

It is often seen as a species complex including at least seven[3][4] closely related and efficient forest-
based malaria vectors in Asia. Hence, its geographical distribution is overlapping with areas of high
malaria prevalence rates and the occurrence of drug resistant Plasmodium falciparum.[5] P. falciparum
is one of the four main protozoan parasites that cause malaria and is one of the leading causes of
malaria deaths. This species complex is of high medical importance for malaria control, in view of the
biological specificities of the members of this complex. Sympatric sibling species of the complex differ in
types of larval habitat, seasonality and behaviour. These differences also exist within the species
suggesting the role of environmental factors in determining these.[2]
Distribution

edit

The complex has been reported mainly from Northeast India, Bangladesh, Myanmar and Thailand.[3] It
has also been reported from other areas in India such as Jammu & Kashmir[6] and Andaman Islands.[7]
One of the species in the complex has also been reported from Shimoga in South India although its
vectorial status is unknown.[3] GIS-based predictive habitat modelling has revealed that over half of
several Northeast Indian states, whole of Thailand and nearly a third of large areas in South Indian states
like Kerala and Goa could harbour this complex.[8]

Hosts

edit

Hosts include cattle.[9]

Control

edit

Insecticide resistance is an increasing concern in southeast Asia.[9] Although ivermectin is thought of as

solely an endoparasiticide, it has proven effective as an endectocide against An. dirus.[9] Cramer et al
2021 use zooprophylaxis-aided ivermectin-based vector elimination (ZAIVE), injecting cattle with the
antiparasitic and getting at least 30 days of increased mortality in this species and An. epiroticus.[9]

References

edit

Sallum MA, Peyton EL,& Wilkerson RC. 2005. Six new species of the Anopheles leucosphyrus group,
reinterpretation of An. elegans and vector implications. Med. Vet. Entomol.,19, 158–199.

Obsomer, Valerie; Defourny, Pierre; Coosemans, Marc (2007). "The Anopheles dirus complex: spatial
distribution and environmental drivers". Malaria Journal. 6 (26): 1474–2875. doi:10.1186/1475-2875-6-
26. PMC 1838916. PMID 17341297.

Dash AP, Adak, T., Raghavendra, K., Singh, OP. (2007) The biology and control of malaria vectors in
India. Current Science 92(11). p.1574 PDF.

Peyton EL. 1989. A new classification for the Leucosphyrus group of Anopheles (Cellia). Mosq. Syst.
21:197–205.

Yang TH (1983) A review of literature on Anopheles balabacensis balabacensis. World Health

Organization. WHO/MAL/83.999, WHO/MAL/83.873M.

Nair, C. P., J. Commun. Dis., 1973, 5, 22–46.

Covell, G., Report of an inquiry into malaria condition in the Andamans, Govt. Press, Delhi, 1927.

Srivastava, Aruna; Nagpal, B. N.; Saxena, Rekha; Subbarao, S. K. (10 May 2001). "Predictive habitat
modelling for forest malaria vector species An. dirus in India – A GIS-based approach" (PDF). Current
Science. 80 (9): 1129–34. Retrieved 16 June 2009.

Ahmad, Sundus Shafat; Rahi, Manju; Saroha, Poonam; Sharma, Amit (10 January 2022). "Ivermectin".
Parasites & Vectors. 15 (1). BioMed Central: 20. doi:10.1186/s13071-021-05124-3. ISSN 1756-3305. PMC
8744265. PMID 35012612.

Anopheles earlei is a small insect found throughout North America. The Anopheles earlei larvae are
found in cold, clear water in ponds and other small bodies of water that contain plant life or vegetation.
[1]

Anopheles earlei

Scientific classificationEdit this classification

Domain:

Eukaryota

Kingdom:

Animalia

Phylum:

Arthropoda

Class:

Insecta

Order:

Diptera

Family:

Culicidae

Genus:

Anopheles

Species:

A. earlei
Binomial name

Anopheles earlei

Vargas, 1943

References

"WRBU: Anopheles earlei". Archived from the original on 2009-04-12. Retrieved 2009-01-27.

Walter Reed Biosystematic Unit Characteristics, Bionomics, Medical Importance

GeoSpecies Knowledge Base University of Wisconsin

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Anopheles freeborni

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Anopheles freeborni, commonly known as the western malaria mosquito, is a species of mosquito in the
family Culicidae.[1][2][3] It is typically found in the western United States and Canada. Adults are brown
to black, with yellow-brown hairs and gray-brown stripes on the thorax. Their scaly wings have four dark
spots, which are less distinct in the male.[4][5]

Anopheles freeborni

Anopheles freeborni feeding on blood from a human finger

Scientific classificationEdit this classification

Domain:

Eukaryota

Kingdom:

Animalia

Phylum:
Arthropoda

Class:

Insecta

Order:

Diptera

Family:

Culicidae

Genus:

Anopheles

Species:

A. freeborni

Binomial name

Anopheles freeborni

Aitken

Male Anopheles freeborni aggregate into swarms to attract potential mates.[6] Females are able to
overwinter, allowing for seasonal development of eggs in the spring.[4][7]

The western malaria mosquito feeds on bloodmeals.[8] Within the United States’ regions of semiarid or
arid climate, it has been historically identified as the primary transmission vector for malaria.[4][9] It
was most notably involved in the malaria outbreaks of northern California during the turn of the 20th
century.[7]

Description

edit

Adult A. freeborni are medium-sized with overall brown to black coloration. They possess a dark
proboscis as well as palpi of similar length to the proboscis on their heads. The head is covered in erect
scales, dark-colored at the posterior, yellow-white at the center, and light at the vertex. They also have a
frontal tuft composed of several light-colored setae.[4]

Adults have a gray-brown prescutellar space and brown to black scutum, which possesses hairs of light
yellow to golden-brown coloration. The central area of the scutum has gray-brown stripes and the
highest density of hairs. The sickle-shaped scutellum also has similarly-colored hairs and setae. The
thorax has setae on the pre-spiracular area, but not on the post-spiracular area.[10] The adult abdomen
is brown to black, also with hairs of yellow-brown coloration.[4]

Their legs have primarily dark scales, with lighter-colored scales at their tips. They possess wings of
about 4.5 mm in length, covered in dark scales, with four spots formed by even darker-colored scales.
These spots are less noticeable on males’ wings.[4] The media and radial sector of their wings' backsides
have tapered scales, which is unique to the species.[10]

Distribution

edit

The western malaria mosquito is present throughout North America, predominantly in the western
United States and British Columbia, Canada.[4] While most common in the United States and Canada, it
has also been observed in Mexico.[4] It typically remains west of the Rocky Mountains, but the mosquito
has been observed slightly eastward in Texas, New Mexico, and southern Colorado.[4] They are thought
to be most populous in California, particularly the valleys of San Joaquin and Sacramento.[4] An.
freeborni have been collected in southern California.[11]

Habitat

edit

Larvae are found in stagnant water; noted larval habitats include rice fields, pools along roads, and
groundwater, with particularly significant larval production in algal masses surrounding rice fields.[4]
Vegetative cover is highly sought after in selection of larval habitats.[8]

Adults typically reside in cool, shaded areas in the daytime, such as drainage tunnels, dark corners of
homes, or beneath bridges. One study in California found hibernating A. freeborni in the nests of wood
rats.[4] Females typically overwinter in human structures, such as basements, houses, garages, and
barns. They do not remain in one location during overwintering, instead traveling to different shelters
throughout the winter.[4][8]

Phylogeny

edit

A. freeborni is part of the genus Anopheles.[1][2][3] It is most closely phylogenetically related to the
species Anopheles hermsi.[12] It belongs to the Angusticorn section, the Maculipennis group, and the
Freeborni subgroup, which also includes the species A. hermsi, A. occidentalis, and A. earlei.[citation
needed]

Flight patterns

edit
Flight patterns of A. freeborni remain in close proximity to breeding sites in the summertime.[13]
Prehibernation flights, during which individuals disperse from breeding sites, take place in September.
These flights are commonly five to ten miles (8 to 16 km) in distance,[4] although flights as long as 17.5
miles (28 km) have been recorded.[13] When the temperature sufficiently rises during hibernation,
flights are restricted to movement between local shelters. In February, females begin their return to
habitats favoring larval development for oviposition.[4][13]

Life history

edit

A. freeborni undergo four stages of development, as do all other Anopheles.[14] The entire
development process takes about 20 days.[7][15]

Anopheles sp. larva

Egg

edit

During the egg, larva, and pupa stages, the mosquito is aquatic. Eggs are deposited directly into still
bodies of water, where they float. Hatching may occur as soon as two to three days after oviposition,
although it may take several weeks depending on environmental temperature.[14][15]

Larva

edit

In contrast with other mosquitoes, Anopheles larvae are horizontally situated directly underneath the
water surface.[7][15] As larvae, all Anopheles mosquitoes develop a head, thorax, and abdomen, but no
legs. They have spiracles on their abdomen, which allows them to breathe at the water surface.[4] After
4 larval instars, each succeeded by molting, they transform into pupae.[14] This process takes
approximately 15 days.[15]

Pupa

edit

Anopheles pupae transform dramatically, forming a cephalothorax from the thorax and head.
Respiratory openings in the cephalothorax again facilitate breathing at the water surface, but no feeding
occurs.[14]

Adult

edit
After several days, the cephalothorax backside ruptures and the adult arises.[14] The adult A. freeborni
gathers its strength on the water surface until it is able to fly away in search of a bloodmeal.[7]

Food resources

edit

Algae is a common food source for A. freeborni

Larvae feed on microorganisms on top of the water, such as algae.[14] Feeding is facilitated by oral
brushes, which generate spiraling currents that direct microorganisms into the mouths of larval A.
freeborni.[15]

The adult western malaria mosquito relies on bloodmeals for food. It has been suggested as an
"opportunistic feeder", with food choice influenced by available hosts.[8] They commonly feed on
humans and sizable domesticated animals, like cattle, horses, and dogs.[16] Deer and sheep are also
notable hosts.[15] A. freeborni have also been observed frequently feeding on rabbits.[16] Studies have
not indicated a general preference for human or animal hosts,[17] although host selection may vary
with availability, human activity, and housing quality.[16] Feeding occurs primarily during and after dusk,
although daytime attacks may occur in when the weather is overcast. Females are also known to
facilitate human bloodmeals by entering their homes.[4]

Mating

edit

Mating of A. freeborni occurs following the aggregation of individuals into swarms. It typically occurs on
evenings during the late summer season, from about July through September. Swarms of males
assemble minutes after sunset, with more individuals joining throughout the first 15 minutes, after
which there is a gradual decrease.[6] Swarming is initiated at light intensities of about 350 lux and ended
at about 0.5 lux. Circadian rhythms also play a major role, as swarming does not occur during the day.
These swarms can reach numbers of 500 to 3000 mosquitoes and last as long as 35 minutes.[18][6]
Smaller-sized males will initiate a swarm to increase their chances of mating by decreasing competition,
but also experience prolonged vulnerability to predators. Smaller-sized males mate with significantly
less frequency than larger-sized males. Females join the swarm during its highest density, and
copulation activity occurs from about 10 to 20 minutes after swarm formation.[19]

Parental care

edit

The gonotrophic cycle indicates the time spent seeking out a host, feeding on a bloodmeal, egg
development, and oviposition.[20] For A. freeborni, this cycle varies from 4 to 6 days, depending on
consumption status. Unfed females have a longer gonotrophic cycle due to the mating and maturation
period required by newly-molted females.[16] Sites selected for oviposition are synonymous with
favored larval habitats. Female oviposition favors shallow, still bodies of water, which are at least
partially sunlit during the day and which preferably contain some algae or other vegetative matter.
Larvae are infrequently developed in bodies of water covered in shadow.[4]

Enemies

edit

E. collocata, a common predator

Known predators include Erythemis collocata and Pantala hymenaea, species belonging to the
Libellulidae family of dragonflies. Predation partially overlaps with mating activity, as western malaria
mosquitoes are most frequently attacked during swarming. Predatory activity begins with swarm
initiation and remains intense for the next 15 minutes. Dragonfly predation is reliant on visual detection,
so attacks decrease as it gets darker. Attacks occur more frequently over open areas, reflecting dragonfly
foraging preferences.[6] Larval dragonflies may also prey upon the mosquito larvae, along with other
predators such as frogs, spiders, and flatworms; these larval predators may be used to control the
population of Anopheles mosquitos in order to decrease malarial transmission.[21]

Interactions with humans

edit

Historically, A. freeborni has been considered the primary transmission vector for malaria in the western
regions of the United States.[4][9] It was primarily involved in the malaria outbreaks of northern
California in the late 19th century and early 20th century.[7] However, the frequency of human
bloodmeals and the mosquito’s potential as a transmission vector may be limited by human behavior
and relative accessibility in certain areas.[6] Recent studies have also questioned the significance of A.
freeborni as a transmission vector due to their morphological similarity with A. hermsi, whose presence
has recently been identified throughout Colorado and Arizona.[9]

Control

edit

Insecticides have commonly been used for mosquito control in the past. Such pesticidal agents as
Bacillus thuringiensis, as well as some piperidine compounds and CIC-4, a lactone, are known to be
effective at controlling A. freeborni larvae.[22][23] In an effort to combat insecticide dependence, recent
studies have investigated mosquitofish (G. affinis), a predator of the mosquito larvae, as a means of
control. Their presence has been found to significantly reduce A. freeborni larval densities in rice fields.
[23]

References

edit
"Anopheles freeborni Report". Integrated Taxonomic Information System. Retrieved 2019-09-23.

"Anopheles freeborni". GBIF. Retrieved 2019-09-23.

"Anopheles freeborni species Information". BugGuide.net. Retrieved 2019-09-23.

Carpenter, S. J.; LaCasse, W. J. (1955). Mosquitoes of North America (North of Mexico). Berkeley, Los
Angeles, London: University of California Press. pp. 39–42. ISBN 0-520-02638-1.

Sandhu TS, Williams GA, Haynes BW, Dhillon MS. Evaluation of arboviral activity at Northwest Mosquito
and Vector Control District, Riverside County, California during 2008. Proc and Papers of the Mosq and
Vector Control Assoc of Calif, vol 77, 2009. p. 108-15.

Yuval, B.; Bouskila, A. (1993-03-01). "Temporal dynamics of mating and predation in mosquito swarms".
Oecologia. 95 (1): 65–69. Bibcode:1993Oecol..95...65Y. doi:10.1007/BF00649508. ISSN 1432-1939. PMID
28313313. S2CID 22921039.

"The Overwintering Mosquito Anopheles Freeborni" (PDF). shastamosquito.org. Shasta Mosquito and
Vector Control District. Retrieved 2019-12-02.

Sinka, Marianne E; Rubio-Palis, Yasmin; Manguin, Sylvie; Patil, Anand P; Temperley, Will H; Gething,
Peter W; Van Boeckel, Thomas; Kabaria, Caroline W; Harbach, Ralph E; Hay, Simon I (2010-08-16). "The
dominant Anopheles vectors of human malaria in the Americas: occurrence data, distribution maps and
bionomic précis". Parasites & Vectors. 3: 72. doi:10.1186/1756-3305-3-72. ISSN 1756-3305. PMC
2936890. PMID 20712879.

Hayden, Craig W.; Fink, T. Michael; Ramberg, Frank B.; Maré, C. John; Mead, Daniel G. (1 March 2001).
"Occurrence of Anopheles hermsi (Diptera: Culicidae) in Arizona and Colorado". Journal of Medical
Entomology. 38 (2): 341–343. doi:10.1603/0022-2585-38.2.341. ISSN 0022-2585. PMID 11296846. S2CID
11532179.

"WRBU: Anopheles freeborni". www.wrbu.org. Retrieved 2019-10-02.

Sandhu TS, Williams GW, Haynes BW, Dhillon MS. Population dynamics of blood-fed female mosquitoes
and comparative efficacy of resting boxes in collecting them from the northwestern part of Riverside
County, California. J Global Infect Dis [serial online] 2013 [cited 2013 Oct 22];5:15-8.

"rDNA-ITS2 sequence generated the phylogenetic tree, showing the..." ResearchGate. Retrieved 2019-
10-02.

Bailey, S. F.; Baerg, D. C. (1967). "The flight habits of Anopheles freeborni Aitken". Proceedings. And
Papers. Amer. Mosq. Control Assoc. 35: 55–69. PMID 5629692 – via CAB Direct.

Prevention, CDC-Centers for Disease Control and (2019-01-28). "CDC - Malaria - About Malaria -
Biology". www.cdc.gov. Retrieved 2019-10-01.
 "The Biology and Control of Mosquitoes in California" (PDF). California Department of Public Health.
Retrieved 2019-12-02.

McHugh, Chad P. (1989-08-01). "Ecology of a Semi-Isolated Population of Adult Anopheles Freeborni:

Abundance, Trophic Status, Parity, Survivorship, Gonotrophic Cycle Length, and Host Selection". The
American Journal of Tropical Medicine and Hygiene. 41 (2): 169–176. doi:10.4269/ajtmh.1989.41.169.
ISSN 0002-9637. PMID 2774063.

"Anopheles (Anopheles) freeborni Aitken, 1939". MAP. Retrieved 2019-10-02.

Howell, Paul I.; Knols, Bart GJ (2009-11-16). "Male mating biology". Malaria Journal. 8 (2): S8.
doi:10.1186/1475-2875-8-S2-S8. ISSN 1475-2875. PMC 2777330. PMID 19917078.

Yuval, B.; Wekesa, J. W.; Washino, R. K. (1993-05-01). "Effect of body size on swarming behavior and
mating success of maleAnopheles freeborni (Diptera: Culicidae)". Journal of Insect Behavior. 6 (3): 333–
342. doi:10.1007/BF01048114. ISSN 1572-8889. S2CID 44027286.

Barker, Christopher M.; Reisen, William K. (2019-01-01), Mullen, Gary R.; Durden, Lance A. (eds.),
"Chapter 4 - Epidemiology of Vector-Borne Diseases", Medical and Veterinary Entomology (Third
Edition), Academic Press, pp. 33–49, ISBN 978-0-12-814043-7, retrieved 2019-12-03

Collins, C. M.; Bonds, J. A. S.; Quinlan, M. M.; Mumford, J. D. (March 2019). "Effects of the removal or
reduction in density of the malaria mosquito, Anopheles gambiae s.l., on interacting predators and
competitors in local ecosystems". Medical and Veterinary Entomology. 33 (1): 1–15.
doi:10.1111/mve.12327. ISSN 0269-283X. PMC 6378608. PMID 30044507.

Coleman, Russell E.; Robert, Leon L.; Roberts, Lyman W.; Glass, Jacqulin A.; Seeley, Doug C.;
Laughinghouse, Andre; Perkins, Peter V.; Wirtz, Robert A. (1993-05-01). "Laboratory Evaluation of
Repellents Against Four Anopheline Mosquitoes (Diptera: Culicidae) and Two Phlebotomine Sand Flies
(Diptera: Psychodidae)". Journal of Medical Entomology. 30 (3): 499–502.
doi:10.1093/jmedent/30.3.499. ISSN 0022-2585. PMID 8510108.

Chandra, G.; Bhattacharjee, I.; Chatterjee, S. N.; Ghosh, A. (8 June 2006). "Mosquito control by
larvivorous fish" (PDF). Indian J Med Res. 127 (1): 13–27. PMID 18316849.

Wikipedia

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Anopheles funestus

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Edit

Anopheles funestus is a species of mosquito in the Culicidae family. This species was first described in
1900 by Giles.[1] The female is attracted to houses where it seeks out humans in order to feed on their
blood, mostly during the night. This mosquito is a major vector of malaria in sub-Saharan Africa.[2]

Anopheles funestus

Scientific classificationEdit this classification

Domain:

Eukaryota

Kingdom:

Animalia

Phylum:

Arthropoda

Class:

Insecta

Order:

Diptera

Family:

Culicidae

Genus:

Anopheles

Species:

A. funestus

Binomial name

Anopheles funestus

Giles, 1900[1]

Distribution and habitat

edit

Anopheles funestus is found in tropical sub-Saharan Africa, its range extending from Senegal to Ethiopia,
Angola, South Africa and Madagascar.[3] Breeding takes place in water, any permanent or semi-
permanent body of fresh water with some emergent vegetation being suitable, including swamps, lake
verges, ponds and rice paddies. The larvae inhabit both sunlit and shaded locations, the vegetation
probably being effective in reducing predation.[2] In the Sahel, increased aridity has moved the northern
limit of its range southward by about 100 km (62 mi). Although it is considered to be a single species,
part of a species complex, it shows some anomalies of behaviour across its range. It is present in the
paddy fields of Madagascar but not in those of West Africa; it used to breed in fast-moving streams in
South Africa before being largely eliminated by the use of insecticides, but when it became re-
established there via Mozambique, it bred in swamps.[3] It is present in mountainous areas of East
Africa at altitudes of up to 2,000 m (6,600 ft) but is largely absent from forests.[3]

Behaviour

edit

The female mosquito lays a raft of eggs on the surface of water. The larval and pupal stages of the life
cycle take place under water, but after metamorphosis, adults of both sexes leave the water and visit
flowers to feed on nectar. Before it starts to breed, the female mosquito needs a meal of vertebrate
blood to provide the protein it needs for egg production; the male does not bite.[4] The adult female
Anopheles funestus is "anthropophilic", being attracted to people rather than to other animals; however
this is not invariably the case, as in Senegal, the populations of this mosquito in the west of the country
feed on human blood while those in the east favour that of other mammals (zoophilic). It is also
"endophilic" in its behaviour; this means it is attracted to the inside of human habitations, both when
feeding and when resting. It feeds at night, typically after 10 p.m. and usually between midnight and
dawn, which gives it access to widely dispersed hosts in a non-alert state.[2]

Vector

edit

Anopheles funestus is an efficient vector of the Plasmodium parasites that cause malaria in humans. This
is because of its endophilic and anthropophilic characteristics, and because the adult insect is relatively
long-lived.[citation needed]

Insecticide resistance

edit

This is a highly adaptable species and many populations have developed resistance to pyrethroid
insecticides, resulting in an upsurge of malarial infections in sub-Saharan Africa in the 1990s.[2] A.
funestus has widespread resistance to DDT and pyrethroids in Southern and West Africa, and in the
Tororo District of Uganda in the east of the continent.[5] The Tororo population was, however, entirely
susceptible to bendiocarb, malathion, and dieldrin.[5]

References

edit

Wikispecies has information related to Anopheles funestus.

Harbach, Ralph (1 June 2019). "Valid Species". Mosquito Taxonomic Inventory. Archived from the
original on 11 June 2019. Retrieved 14 June 2019.

"Anopheles (Cellia) funestus Giles, 1900". Malaria Atlas Project. Retrieved 14 June 2019.[permanent
dead link]

Mouchet, Jean; Carnevale, Pierre; Manguin, Sylvie (2008). Biodiversity of Malaria in the World. John
Libbey Eurotext. pp. 72–74.

"The life cycle of the mosquito". The Anti-Mosquito Site. Archived from the original on 10 October 2018.
Retrieved 15 June 2019.

Morgan, John C.; Irving, Helen; Okedi, Loyce M.; Steven, Andrew; Wondji, Charles S. (2010-07-29).
Rénia, Laurent (ed.). "Pyrethroid Resistance in an Anopheles funestus Population from Uganda". PLoS
ONE. 5 (7). Public Library of Science (PLoS): e11872. Bibcode:2010PLoSO...511872M.
doi:10.1371/journal.pone.0011872. ISSN 1932-6203. PMC 2912372. PMID 20686697.

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Anopheles gambiae

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The Anopheles gambiae complex consists of at least seven morphologically indistinguishable species of
mosquitoes in the genus Anopheles. The complex was recognised in the 1960s and includes the most
important vectors of malaria in sub-Saharan Africa, particularly of the most dangerous malaria parasite,
Plasmodium falciparum.[2] It is one of the most efficient malaria vectors known. The An. gambiae
mosquito additionally transmits Wuchereria bancrofti which causes lymphatic filariasis, a symptom of
which is elephantiasis.[3]

Anopheles gambiae
Scientific classification

Kingdom:

Animalia

Phylum:

Arthropoda

Class:

Insecta

Order:

Diptera

Family:

Culicidae

Subfamily:

Anophelinae

Genus:

Anopheles

Species complex:

A. gambiae sensu lato

Species:

A. arabiensis

A. bwambae

A. melas

A. merus

A. quadriannulatus

Anopheles gambiae sensu stricto

A. coluzzii
A. amharicus

Binomial name

Anopheles gambiae

Giles 1902[1]

The tube-like heart (green) extends along the body, interlinked with the diamond-shaped alary muscles
(also green) and surrounded by pericardial cells (red). Blue depicts cell nuclei.

Discovery and elements

edit

The Anopheles gambiae complex or Anopheles gambiae sensu lato was recognized as a species complex
only in the 1960s. The A. gambiae complex consists of:

Anopheles arabiensis

Anopheles bwambae

Anopheles melas

Anopheles merus

Anopheles quadriannulatus[4][5]

Anopheles gambiae sensu stricto (s.s.)[6]

Anopheles coluzzii

Anopheles amharicus

The individual species of the complex are morphologically difficult to distinguish from each other,
although it is possible for larvae and adult females. The species exhibit different behavioural traits. For
example, Anopheles quadriannulatus is both a saltwater and mineralwater species. A. melas and A.
merus are saltwater species, while the remainder are freshwater species.[7]Anopheles quadriannulatus
generally takes its blood meal from animals (zoophilic), whereas Anopheles gambiae sensu stricto
generally feeds on humans, i.e. is considered anthropophilic.[citation needed] Identification to the
individual species level using the molecular methods of Scott et al. (1993)[8] can have important
implications in subsequent control measures.

Anopheles gambiae in the strict sense

edit
An. gambiae sensu stricto (s.s.) has been discovered to be currently in a state of diverging into two
different species—the Mopti (M) and Savannah (S) strains—though as of 2007, the two strains are still
considered to be a single species.[9][10]

The mechanism of species recognition appears to be sounds emitted by the wings and identified by
Johnston's organ.[11]

Genome

edit

An. gambiae s.s. genomes have been sequenced three times, once for the M strain, once for the S strain,
and once for a hybrid strain.[9][10] Currently, ~90 miRNA have been predicted in the literature (38
miRNA officially listed in miRBase) for An. gambiae s.s. based upon conserved sequences to miRNA
found in Drosophila.[citation needed] Holt et al., 2002 and Neafsey et al., 2016 find transposable
elements to be ~13% of the genome, similar to Drosophila melanogaster (also in Diptera).[12] However
they find the proportion of TE types to be very different from D. melanogaster with approximately the
same composition of long terminal repeat retrotransposons, non-long terminal repeat retrotransposons
and DNA transposons.[12] These proportions are believed to be representative of the genus.[12]

The genetics and genomics of sex chromosomes have been discovered and studied by Windbichler et al.,
2007 and Galizi et al., 2014 (a Physarum polycephalum homing endonuclease which destroys X
chromosomes), Windbichler et al., 2008 and Hammond et al., 2016 (methods to reduce the female
population), Windbichler et al., 2011 (trans from yeast), Bernardini et al., 2014 (a method to increase
the male population), Kyrou et al., 2018 (a female necessary exon and a homing endonuclease to drive
it), Taxiarchi et al., 2019 (sex chromosome dynamics in general) and Simoni et al., 2020 (an X
chromosome destroying site specific nuclease).[13] See § Gene drive below for their applications.

An. gambiae has a high degree of polymorphism. This is especially true in the cytochrome P450s, Wilding
et al., 2009 finding 1 single nucleotide polymorphism (SNP)/26 base pairs. This species has the highest
amount of polymorphism in the CYPs of any insect known, much tending to be found in "scaffolds" that
are found only in particular subpopulations. These are termed "dual haplotype regions" by Holt et al.,
2002 who sequenced the PEST strain.[14]: 241

In common with many chromosomes, An. gambiae codes for spindle and kinetochore-associated
proteins. Hanisch et al., 2006 locate AgSka1, the spindle and kinetochore-associated protein 1 gene, at
EAL39257.[15]

The entire Culicidae family may or may not conserve epigenetic mechanisms – as of 2012 this remains
unresolved.[16] Toward answering this question, Marhold et al., 2004 compare their own previous work
in Drosophila melanogaster against new sequences of D. pseudoobscura and An. gambiae.[16] They find
all three do share the DNA methylation enzyme DNMT2 (DmDNMT2, DpDNMT2, and AgDNMT2).[16]
This suggests all Diptera may conserve an epigenetic system employing Dnmt2.[16]

Hosts
edit

Hosts include Bos taurus, Capra hircus, Ovis aries and Sus scrofa.[17]

Parasites

edit

Parasites include Plasmodium berghei (for which it also serves as a vector),[18][19][20] and the
bioinsecticides/entomopathogenic fungi Metarhizium robertsii[18] and Beauveria bassiana.[18] All three
of these parasites combine with insecticides to reduce fitness – see § Insecticides below.[18]
CRISPR/Cas9 and U6-gRNA are increasingly (as of 2020) being used together for knockout experiments in
mosquitoes.[20] Dong et al., 2018 develops and presents a new U6-gRNA+Cas9 technique in An.
gambiae, and utilizes it to knock out fibrinogen related protein 1 (FREP1), thereby severely reducing
infection of the mosquito by P. berghei and P. falciparum.[20] However this also demonstrates the
centrality of FREP1 to the insect's success, impairing all measured activities across all life stages.[20]
Yang et al., 2020 uses the Dong method to do the same with mosGILT, also severely reducing
Plasmodium infection of the mosquito but also finding a vital life process is impaired, in mosGILT's case
ovary development.[20]

Control

edit

Insecticides

edit

Parasites/bioinsecticides and chemical insecticides synergistically reduce fitness. Saddler et al., 2015
finds even An. gambiae with knockdown resistance (kdr) are more susceptible to DDT if they are first
infected with Plasmodium berghei[18][19] and Farenhorst et al., 2009 the same for Metarhizium
robertsii or Beauveria bassiana.[18] This is probably due to an effect found by Félix et al., 2010 and
Stevenson et al., 2011: An. gambiae alters various activities – especially CYP6M2 – in response to P.
berghei invasion. CYP6M2 is known to somehow produce pyrethroid resistance, and pyrethroids and
DDT share a mechanism of action.[19]

Gene drive

edit

Research relevant to the development of gene drive controls of An. gambiae have been performed by
Windbichler et al., 2007, Windbichler et al., 2008, Windbichler et al., 2011, Bernardini et al., 2014, Galizi
et al., 2014, Hammond et al., 2016, Kyrou et al., 2018, Taxiarchi et al., 2019 and Simoni et al., 2020.[13]
For specific genes involved see § Genome above. These can all be used in pest control because they
induce infertility.[13]
Fecundity

edit

Fecundity of An. gambiae depends on the detoxification of reactive oxygen species (ROS) by catalase.
[21] Reduction in catalase activity significantly reduces reproductive output of female mosquitoes,
indicating that catalase plays a central role in protecting oocytes and early embryos from ROS damage.
[21]

Historical note

edit

An. gambiae invaded northeastern Brazil in 1930, which led to a malaria epidemic in 1938/1939.[22] The
Brazilian government assisted by the Rockefeller Foundation in a programme spearheaded by Fred
Soper eradicated these mosquitoes from this area. This effort was modeled on the earlier success in
eradication of Aedes aegypti as part of the yellow fever control program. The exact species involved in
this epidemic has been identified as An. arabiensis.[23]

Peptide hormones

edit

Kaufmann and Brown 2008 find the An. gambiae adipokinetic hormone (AKH) mobilizes carbohydrates
but not lipids. Meanwhile AKH/Corazonin Peptide (ACP) does not mobilize (or inhibit mobilization) of
either. Mugumbate et al., 2013 provides in solution and membrane bound structures from a nuclear
magnetic resonance investigation.[24]

References

edit

Giles, G. M. (1902). A handbook of the gnats or mosquitoes giving the anatomy and life history of the
Culicidae together with descriptions of all species noticed up to the present date. London, United
Kingdom: John Bale, Sons & Danielsson.

"Anopheles gambiae complex". Walter Reed Army Institute of Research. Archived from the original on
2007-09-29.

"Lymphatic filariasis". www.who.int. Retrieved 2020-04-10.

Besansky NJ, Powell JR, Caccone A, Hamm DM, Scott JA, Collins FH (July 1994). "Molecular phylogeny of
the Anopheles gambiae complex suggests genetic introgression between principal malaria vectors".
Proceedings of the National Academy of Sciences. 91 (15): 6885–8. Bibcode:1994PNAS...91.6885B.
doi:10.1073/pnas.91.15.6885. PMC 44302. PMID 8041714.
 Wilkins EE, Howell PI, Benedict MQ (2006). "IMP PCR primers detect single nucleotide polymorphisms
for Anopheles gambiae species identification, Mopti and Savanna rDNA types, and resistance to dieldrin
in Anopheles arabiensis". Malaria Journal. 5 (1): 125. doi:10.1186/1475-2875-5-125. PMC 1769388.
PMID 17177993.

Yakob, Laith (2011). "Epidemiological consequences of a newly discovered cryptic subgroup of

Anopheles gambiae". Biology Letters. 7 (6): 947–949. doi:10.1098/rsbl.2011.0453. PMC 3210673. PMID
21693489.

G.B. White (1974). "Anopheles gambiae complex and disease transmission in Africa". Transactions of
the Royal Society of Tropical Medicine and Hygiene. 68 (4): 278–298. doi:10.1016/0035-9203(74)90035-
2. PMID 4420769.

C. Fanello; F. Santolamazza; A. Della Torre (2002). "Simultaneous identification of species and molecular
forms of the Anopheles gambiae complex by PCR-RFLP". Medical and Veterinary Entomology. 16 (4):
461–4. doi:10.1046/j.1365-2915.2002.00393.x. PMID 12510902. S2CID 28983355.

"Anopheles gambiae: First genome of a vector for a parasitic disease". Genoscope. Archived from the
original on 2011-08-07.

Lawniczak, M. K.; et al. (Oct 22, 2010). "Widespread divergence between incipient Anopheles gambiae
species revealed by whole genome sequences". Science. 330 (6003): 512–4.
Bibcode:2010Sci...330..512L. doi:10.1126/science.1195755. PMC 3674514. PMID 20966253.

Pennetier, Cédric; Warren, Ben; Dabiré, K. Roch; Russell, Ian J.; Gibson, Gabriella (2010). ""Singing on
the Wing" as a Mechanism for Species Recognition in the Malarial Mosquito Anopheles gambiae".
Current Biology. 20 (2): 131–136. doi:10.1016/j.cub.2009.11.040. PMID 20045329. S2CID 15185976.

Gilbert, Clément; Peccoud, Jean; Cordaux, Richard (2021-01-07). "Transposable Elements and the
Evolution of Insects" (PDF). Annual Review of Entomology. 66 (1): 355–372. doi:10.1146/annurev-ento-
070720-074650. ISSN 0066-4170. PMID 32931312. S2CID 221747772.

Hay, Bruce A.; Oberhofer, Georg; Guo, Ming (2021-01-07). "Engineering the Composition and Fate of
Wild Populations with Gene Drive". Annual Review of Entomology. 66 (1): 407–434.
doi:10.1146/annurev-ento-020117-043154. ISSN 0066-4170. PMID 33035437. S2CID 222257628.

Gilbert, Lawrence I., ed. (2012). Insect molecular biology and biochemistry. Amsterdam Boston:
Academic Press. pp. x+563. ISBN 978-0-12-384747-8. OCLC 742299021.

Cheeseman, Iain M.; Desai, Arshad (2008). "Molecular architecture of the kinetochore–microtubule
interface". Nature Reviews Molecular Cell Biology. 9 (1). Nature Portfolio: 33–46. doi:10.1038/nrm2310.
ISSN 1471-0072. PMID 18097444. S2CID 34121605.
 Severson, David W.; Behura, Susanta K. (2012-01-07). "Mosquito Genomics: Progress and Challenges".
Annual Review of Entomology. 57 (1). Annual Reviews: 143–166. doi:10.1146/annurev-ento-120710-
100651. ISSN 0066-4170. PMID 21942845.

"Anopheles gambiae". Invasive Species Compendium. CABI. 2019-11-24. Retrieved 2022-01-26.

Alout, Haoues; Labbé, Pierrick; Chandre, Fabrice; Cohuet, Anna (2017). "Malaria Vector Control Still
Matters despite Insecticide Resistance". Trends in Parasitology. 33 (8): 610–618.
doi:10.1016/j.pt.2017.04.006. ISSN 1471-4922. PMID 28499699. S2CID 32524464.

Minetti, Corrado; Ingham, Victoria A; Ranson, Hilary (2020). "Effects of insecticide resistance and
exposure on Plasmodium development in Anopheles mosquitoes" (PDF). Current Opinion in Insect
Science. 39: 42–49. doi:10.1016/j.cois.2019.12.001. ISSN 2214-5745. PMID 32109860. S2CID 211563675.

Caragata, E.P.; Dong, S.; Dong, Y.; Simões, M.L.; Tikhe, C.V.; Dimopoulos, G. (2020-09-08). "Prospects
and Pitfalls: Next-Generation Tools to Control Mosquito-Transmitted Disease". Annual Review of
Microbiology. 74 (1). Annual Reviews: 455–475. doi:10.1146/annurev-micro-011320-025557. ISSN 0066-
4227. PMID 32905752. S2CID 221625690.

DeJong RJ, Miller LM, Molina-Cruz A, Gupta L, Kumar S, Barillas-Mury C (February 2007). "Reactive
oxygen species detoxification by catalase is a major determinant of fecundity in the mosquito Anopheles
gambiae". Proceedings of the National Academy of Sciences. 104 (7): 2121–6.
Bibcode:2007PNAS..104.2121D. doi:10.1073/pnas.0608407104. PMC 1892935. PMID 17284604.

Killeen GF (October 2003). "Following in Soper's footsteps: northeast Brazil 63 years after eradication of
Anopheles gambiae". The Lancet Infectious Diseases. 3 (10): 663–6. doi:10.1016/S1473-3099(03)00776-
X. PMID 14522266.

Parmakelis A, Russello MA, Caccone A, et al. (January 2008). "Historical analysis of a near disaster:
Anopheles gambiae in Brazil". The American Journal of Tropical Medicine and Hygiene. 78 (1): 176–8.
doi:10.4269/ajtmh.2008.78.176. PMID 18187802.

Strand, M.R.; Brown, M.R.; Vogel, K.J. (2016). "Mosquito Peptide Hormones". Advances in Insect
Physiology. Vol. 51. Elsevier. pp. 145–188. doi:10.1016/bs.aiip.2016.05.003. ISBN 9780128024577. ISSN
0065-2806. PMC 6338476. PMID 30662099. S2CID 58546659.

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Anopheles introlatus

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Edit

Anopheles introlatus (formerly Anopheles balabacensis introlatus) is the main vector for Plasmodium
cynomolgi (a simian malaria) in Malaya.[1]

Anopheles introlatus

Scientific classificationEdit this classification

Domain:

Eukaryota

Kingdom:

Animalia

Phylum:

Arthropoda

Class:

Insecta

Order:

Diptera

Family:

Culicidae

Genus:

Anopheles

Species:

A. introlatus

Binomial name

Anopheles introlatus

Colless, 1957

References

edit
 Eyles DE, Warren M, Guinn E, et al. (1963). "Identification of Anopheles balabacensis introlatus as a
vector of monkey malaria in Malaya". Bull World Health Organ. 28 (1): 134–5. PMC 2554658. PMID
14166986.

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Anopheles latens

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The Anopheles latens mosquito (part of the An. leucosphyrus group)[1] is an important vector for the
transmission of malaria in humans and monkeys in Southeast Asia.[2] It is an important vector for the
transmission of human malaria in Sarawak; but because it is attracted to both humans and to macaques
it is also responsible for the transmission of simian malarias to humans (Plasmodium knowlesi[3] and
possibly P. inui[2] as well).

Anopheles latens

Scientific classificationEdit this classification

Domain:

Eukaryota

Kingdom:

Animalia

Phylum:

Arthropoda

Class:

Insecta

Order:

Diptera

Family:
Culicidae

Genus:

Anopheles

Species:

A. latens

Binomial name

Anopheles latens

Sallum, Peyton, Wilkerson, 2005

A. latens tends to bite from 6 p.m. throughout the night, peaking at midnight.[3] It is found in forests
and at forest fringes, but tends not to enter human dwellings.[3]

References

edit

Sallum MA, Peyton EL, Wilkerson RC (2005). "Six new species of the Anopheles leucosphyrus group,
reinterpretation of An. elegans and vector implications". Med Vet Entomol. 19 (2): 158–99.
doi:10.1111/j.0269-283X.2005.00551.x. PMID 15958025. Archived from the original on September 27,
2017.

Wharton, R.H.; Eyles, DE; Warren, M; Moorhouse, DE (1962). "Anopheles leucosphyrus identified as a
vector of monkey malaria in Malaya". Science. 137 (3532): 758. doi:10.1126/science.137.3532.758.
PMID 14006429.

Vythilingam I, Tan CH, Asmad M, Chan ST, Lee KS, Singh B (2006). "Natural transmission of Plasmodium
knowlesi to humans by Anopheles latens in Sarawak, Malaysia" (PDF). Trans R Soc Trop Med Hyg. 100
(11): 1087–88. doi:10.1016/j.trstmh.2006.02.006. PMID 16725166.

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Anopheles maculipennis

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Anopheles maculipennis is a species of mosquito that can be found mostly in Europe and New Zealand.
It is one of the main vectors of malaria.[1][2][3]

Anopheles maculipennis

Scientific classificationEdit this classification

Domain:

Eukaryota

Kingdom:

Animalia

Phylum:

Arthropoda

Class:

Insecta

Order:

Diptera

Family:

Culicidae

Genus:

Anopheles

Species:

A. maculipennis

Binomial name

Anopheles maculipennis

Meigen, 1818

References

edit
 "Anopheles maculipennis Meigen, 1818 | Walter Reed Biosystematics Unit (WRBU)". wrbu.si.edu.
Retrieved 2022-03-11.

"Anopheles Maculipennis - an overview | ScienceDirect Topics". www.sciencedirect.com. Retrieved

2022-03-11.

"New map shows the presence of Anopheles maculipennis s.l. mosquitoes in Europe". European Centre
for Disease Prevention and Control. 2018-09-12. Retrieved 2022-03-11.

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Anopheles moucheti

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Anopheles moucheti is a species of mosquito. It is an anthropophile. It can be mainly located in Congo

Basin forest, Central Africa. It was the main vector of malaria.[1][2]

Anopheles moucheti

Scientific classificationEdit this classification

Domain:

Eukaryota

Kingdom:

Animalia

Phylum:

Arthropoda

Class:

Insecta

Order:

Diptera
Family:

Culicidae

Genus:

Anopheles

Species:

A. moucheti

Binomial name

Anopheles moucheti

Evan, 1923

References

edit

Antonio-Nkondjio, Christophe; Simard, Frédéric (2013), Manguin, Sylvie (ed.), "Highlights on Anopheles
nili and Anopheles moucheti, Malaria Vectors in Africa", Anopheles Mosquitoes: New Insights into
Malaria Vectors, Wellcome Trust–Funded Monographs and Book Chapters, Rijeka (HR): InTech, ISBN
978-953-51-1188-7, PMID 28045480, retrieved 2022-03-11

"Anopheles moucheti Evans, 1923 | Walter Reed Biosystematics Unit (WRBU)". wrbu.si.edu. Retrieved
2022-03-11.

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Anopheles nili

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Anopheles nili is a species of mosquito[1] in the Culicidae family.[2] It comprises the following elements:
An. carnevalei, An. nili, An. ovengensis and An. somalicus.[3] The scientific name of this species was first
published in 1904 by Theobald.[4] It is the main mosquito species found in the south Cameroon forest
zone which bites humans.[5] It is known as a problematic carrier of malaria,[6] although newly
discovered, closely related species in the same genus have also been found to interact with A. nili as a
disease vector.[7] In that, they both have similar feeding habits on local targets in the Cameroon region.
[8]

Anopheles nili

Scientific classificationEdit this classification

Domain:

Eukaryota

Kingdom:

Animalia

Phylum:

Arthropoda

Class:

Insecta

Order:

Diptera

Family:

Culicidae

Genus:

Anopheles

Species:

A. nili

Binomial name

Anopheles nili

Theobald, 1904

Life Cycle

edit

For more information, see mosquito life cycle.

Similar to all mosquitoes, Anopheles nili go through a life stages of egg, larva, pupa, then emerge as
adults.[9]

A. nili is a generalist species, meaning that they may adapt to different environments to survive.[10] This
has worked to the detriment of human health, as the mosquito species has become very well adapted to
spawning in dam reservoirs.[11] As A. nili shifts its habitats to areas of high human concentration, they
also begin to mate and spawn their young in nearby reservoirs, such as dams. This of course allows
spread of malaria to the nearby populace.[9]

As Malaria Carriers

edit

Mosquitoes are well known carriers of malaria, A. nili being without exception. As a native species that
has adapted to live with the environmental changes of Cameroon it marks itself as a common species
found to be carrying malarial parasites in the newly built dams of Cameroon.[12]

Dam development in Cameroon marks itself as a grave detriment to native populations health, through
the spread of malaria. Dam developments in particular, because mass environmental changes alter the
balance of disease hosts, vectors, and parasite development.[9] Studies have been made in recent
history before and after dams are built in Cameroon to better understand the method in how malaria is
transmitted. It was found in studies dating back as long as1979, where native mosquitoes, like A. nili,
favoured dams as breeding grounds; and thus ripe for malaria breakouts.[13] It was also found in those
researches that the malarial parasites are found in the midgut of mosquitoes, that being an example of
the parasites persistence.[14]

References

edit

Antonio-Nkondjio, Christophe; Simard, Frédéric (2013), Manguin, Sylvie (ed.), "Highlights on Anopheles
nili and Anopheles moucheti, Malaria Vectors in Africa", Anopheles Mosquitoes: New Insights into
Malaria Vectors, Wellcome Trust–Funded Monographs and Book Chapters, Rijeka (HR): InTech, ISBN
978-953-51-1188-7, PMID 28045480, retrieved 2020-08-03

Ossè, Razaki A.; Tokponnon, Filémon; Padonou, Germain Gil; Sidick, Aboubakar; Aïkpon, Rock; Fassinou,
Arsène; Koukpo, Come Z.; Sèwadé, Wilfrid; Akinro, Bruno; Sovi, Arthur; Aïssi, Melchior (2019-04-29).
"Involvement of Anopheles nili in Plasmodium falciparum transmission in North Benin". Malaria Journal.
18 (1): 152. doi:10.1186/s12936-019-2792-0. ISSN 1475-2875. PMC 6489317. PMID 31036025.

"Anopheles (Cellia) nili species complex". MAP. Retrieved 2020-08-03.

Harbach, Ralph (2013) Mosquito Taxonomic Inventory - Valid Species (Last updated May 5, 2013)
 Carnevale, P.; Goff, G. Le; Toto, J.-C.; Robert, V. (1992). "Anopheles nili as the main vector of human
malaria in villages of southern Cameroon" (PDF). Medical and Veterinary Entomology. 6 (2): 135–138.
doi:10.1111/j.1365-2915.1992.tb00590.x. ISSN 1365-2915. PMID 1421483. S2CID 13558360.

Antonio-Nkondjio, Christopher (October 26, 2019). "Review of malaria situation in Cameroon: technical
viewpoint on challenges and prospects for disease elimination". Parasites & Vectors. 12 (1): 501.
doi:10.1186/s13071-019-3753-8. PMC 6815446. PMID 31655608 – via Web of Science.

Sharakhova, Maria V.; Peery, Ashley; Antonio-Nkondjio, Christophe; Xia, Ai; Ndo, Cyrille; Awono-
Ambene, Parfait; Simard, Frederic; Sharakhov, Igor V. (2013-06-06). "Cytogenetic analysis of Anopheles
ovengensis revealed high structural divergence of chromosomes in the Anopheles nili group". Infection,
Genetics and Evolution. 16: 341–348. doi:10.1016/j.meegid.2013.03.010. PMC 3669242. PMID
23523820.

Wanji, Samuel; Tanke, Theodore; Atanga, Sali N.; Ajonina, Caroline; Nicholas, Tendongfor; Fontenille,
Didier (2003-07-07). "Anopheles species of the mount Cameroon region: biting habits, feeding
behaviour and entomological inoculation rates". Tropical Medicine and International Health. 8 (7): 643–
649. doi:10.1046/j.1365-3156.2003.01070.x. ISSN 1360-2276. PMID 12828548. S2CID 7328386.

Prevention, CDC-Centers for Disease Control and (2020-07-16). "CDC - Malaria - About Malaria -
Biology". www.cdc.gov. Retrieved 2020-10-05.

Antonio-Nkondjio, Christophe; Ndo, Cyrille; Costantini, Carlo; Awono-Ambene, Parfait; Fontenille,

Didier; Simard, Frédéric (2009-11-05). "Distribution and larval habitat characterization of Anopheles
moucheti, Anopheles nili, and other malaria vectors in river networks of southern Cameroon". Acta
Tropica. 112 (3): 270–276. doi:10.1016/j.actatropica.2009.08.009. PMID 19682965.

Mbakop, Lili R.; Awono-Ambene, Parfait H.; Mandeng, Stanislas E.; Ekoko, Wolfgang E.; Fesuh, Betrand
N.; Antonio-Nkondjio, Christophe; Toto, Jean-Claude; Nwane, Philippe; Fomena, Abraham; Etang,
Josiane (2019-05-09). "Malaria Transmission around the Memve'ele Hydroelectric Dam in South
Cameroon: A Combined Retrospective and Prospective Study, 2000–2016". International Journal of
Environmental Research and Public Health. 16 (9): 1618. doi:10.3390/ijerph16091618. ISSN 1660-4601.
PMC 6539953. PMID 31075820.

Ndo, Cyrille; Antonio-Nkondjio, Christophe; Cohuet, Anna; Ayala, Diego; Kengne, Pierre; Morlais,
Isabelle; Awono-Ambene, Parfait H; Couret, Daniel; Ngassam, Pierre; Fontenille, Didier; Simard, Frédéric
(2010-06-12). "Population genetic structure of the malaria vector Anopheles nili in sub-Saharan Africa".
Malaria Journal. 9 (1): 161. doi:10.1186/1475-2875-9-161. ISSN 1475-2875. PMC 2898787. PMID
20540796.

Mbakop, Lili R.; Awono-Ambene, Parfait H.; Mandeng, Stanislas E.; Ekoko, Wolfgang E.; Fesuh, Betrand
N.; Antonio-Nkondjio, Christophe; Toto, Jean-Claude; Nwane, Philippe; Fomena, Abraham; Etang,
Josiane (May 2019). "Malaria Transmission around the Memve'ele Hydroelectric Dam in South
Cameroon: A Combined Retrospective and Prospective Study, 2000–2016". International Journal of
Environmental Research and Public Health. 16 (9): 1618. doi:10.3390/ijerph16091618. ISSN 1661-7827.
PMC 6539953. PMID 31075820.

Robert, V.; le Goff, G.; Gouagna, L. C.; Sinden, M.; Kieboom, J.; Kroneman, R.; Verhave, J. P. (1998-01-
15). "Kinetics and efficiency of Plasmodium falciparum development in the midguts of Anopheles
gambiae, An. funestus and An. nili". Annals of Tropical Medicine & Parasitology. 92 (1): 115–118.
doi:10.1080/00034983.1998.11813268. ISSN 0003-4983. PMID 9614461. S2CID 37768559.

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Anopheles punctipennis

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Anopheles punctipennis is a species of mosquito in genus Anopheles. It is native to North America.[2]

Anopheles punctipennis

Scientific classificationEdit this classification

Domain:

Eukaryota

Kingdom:

Animalia

Phylum:

Arthropoda

Class:

Insecta

Order:

Diptera

Family:
Culicidae

Genus:

Anopheles

Species:

A. punctipennis

Binomial name

Anopheles punctipennis

(Say, 1823)

Synonyms[1]

Culex hyemalis

Culex punctipennis

The larvae of this species can be found in many kinds of natural and artificial water bodies, especially
cool, clear waters such as streams. Females feed on blood, including the blood of humans, and may bite
during the day or night. They generally stay outdoors and are rarely found inside dwellings.[2] This
species is a vector of P. vivax, one of the protozoans that cause malaria.[3]

References

edit

Anopheles punctipennis. Integrated Taxonomic Information System (ITIS)

Anopheles punctipennis. Walter Reed Biosystematics Unit, Walter Reed Army Institute of Research.

Jensen, T., et al. (1998). Lake Vera revisited: parity and survival rates of Anopheles punctipennis at the
site of a malaria outbreak in the Sierra Nevada foothills of California. The American Journal of Tropical
Medicine and Hygiene, 59(4), 591-594.

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Anopheles quadrimaculatus

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Anopheles quadrimaculatus is a species of mosquito mainly located in Eastern United States, America.
The species is a main vector of malaria.[1]

Anopheles quadrimaculatus

Scientific classificationEdit this classification

Domain:

Eukaryota

Kingdom:

Animalia

Phylum:

Arthropoda

Class:

Insecta

Order:

Diptera

Family:

Culicidae

Genus:

Anopheles

Species:

A. quadrimaculatus

Binomial name

Anopheles quadrimaculatus

Say, 1824
References

edit

"Anopheles quadrimaculatus Say, 1824 | Walter Reed Biosystematics Unit (WRBU)". www.wrbu.si.edu.
Retrieved 2023-07-14.

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Anopheles stephensi

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Anopheles stephensi is a primary mosquito vector of malaria in urban India and is included in the same
subgenus as Anopheles gambiae, the primary malaria vector in Africa.[1] A. gambiae consists of a
complex of morphologically identical species of mosquitoes, along with all other major malaria vectors;
however, A. stephensi has not yet been included in any of these complexes.[2] Nevertheless, two races
of A. stephensi exist based on differences in egg dimensions and the number of ridges on the eggs; A. s.
stephensi sensu stricto, the type form, is a competent malaria vector that takes place in urban areas,
and A. s. mysorensis, the variety form, exists in rural areas and exhibits considerable zoophilic
behaviour, making it a poor malaria vector.[3] However, A. s. mysorensis is a detrimental vector in Iran.
[4] An intermediate form also exists in rural communities and peri-urban areas, though its vector status
is unknown.[4] About 12% of malaria cases in India are due to A. stephensi.[5]

Anopheles stephensi

Scientific classificationEdit this classification

Domain:

Eukaryota

Kingdom:

Animalia

Phylum:

Arthropoda
Class:

Insecta

Order:

Diptera

Family:

Culicidae

Genus:

Anopheles

Species:

A. stephensi

Binomial name

Anopheles stephensi

Liston, 1901

In November 2015, an American research group demonstrated that an A. stephensi with genetic
modifications could be rendered incapable of transmitting malaria, and that 99.5% of the mutant
mosquitoes' offspring were also immune.[6]

In April of 2023, a malaria outbreak at Dire Dawa University in Ethiopia affected 1,300 students. The
outbreak was a mystery because it occurred in the dry season and in an urban area, both atypical
conditions for common cases of malaria in this area of the world. Blood tests confirming malaria's ring-
shaped parasite ultimately led researchers to conclude that it was the work of A. stephensi, which
thrives in urban areas and dry seasons, and has a resistance to insecticides.[7]

A team of scientists, headed by an entomologist from the University of Oxford, conducted an evaluation
of Africa's environments to determine if they provide suitable conditions for the A. stephensi mosquito.
Their findings indicate that the ongoing spread of this species could potentially expose an additional 126
million persons to the risk of malaria.[7]

Habitat

edit

In rural areas, the larvae of A. stephensi may exist in many aquatic habitats, such as ponds, streams,
swamps, marshes, and other sources of standing water.[8] They may also occupy smaller environments,
such as tree holes, leaf axils, and man-made containers.[9] The larvae of A. s. mysorensis exclusively
prefer to occupy stone pots and earthenware containers.[4] This species is also able to endure high
levels of salinity, and have been found to breed readily in water where the salinity is equal to or even
surpassing that of sea water.[10] Furthermore, A. stephensi breeds in a number of different water-
bodies in urban areas, but predominantly in artificial containers, walls, overhead tanks, and ground level
water tanks.[11]

Most larvae feed on microorganisms and particle matter suspended in water.[9] However, later in
development, adult males feed on the nectar of flowers, whereas females take blood meals, which help
produce viable eggs.[3]

Hosts

edit

Hosts include Bos taurus, Mus musculus and Pimephales promelas.[12]

Parasites

edit

A. stephensi is a vector of bovine leukemia virus[12] and Plasmodium berghei.[13] Mack and
Vandenberg performed a series of experiments 1978-1979 finding that P. berghei derives nutrients from
A. stephensi hemolymph during the sporogonic phase.[13] A. stephensi is an important vector for the
human malaria species Plasmodium falciparum.[14]

Biochemistry

edit

Mack and Vandenberg characterized A. stephensi's hemolymph composition in the late 1970s.[13]

Distribution

edit

A. stephensi is a subtropical species that predominates in the Indian subcontinent (except Nepal and Sri
Lanka)[3] and is also distributed across the Middle East and South Asia region, existing in countries such
as: Afghanistan, Bahrain, Bangladesh, China, Egypt, India, Iran, Iraq, Oman, Pakistan, Saudi Arabia, and
Thailand.[2] A. stephensi was discovered to be established on the continent of Africa, in Djibouti on the
Horn of Africa in 2012[15] or 2013,[16] in 2016 in Ethiopia,[17][15] Sri Lanka in 2017,[15] and in 2019 in
the Republic of the Sudan/North Sudan.[15]

Seasonal activity

edit
A. stephensi is considered to be endophilic and endophagic, regardless that it may feed outdoors during
the summer, when weather is warmer and humans and animals are more likely to sleep outside in the
open air.[10] Although indoor feeding habits have shown no variation between seasons, adult females
tend to feed more often at night during the summer rather than during the day in winter.[4] A.
stephensi shows a greater preference for humans over animals in urban areas, where they can be found
year-round.[4]

Insecticide resistance

edit

The introduced African population appears to have arrived with several insecticide resistances.[15]

References

edit

Valenzuela, Jesus G.; Francischetti, Ivo M.B.; Pham, Van My; Garfield, Mark K.; Ribeiro, José M.C. (2003).
"Exploring the salivary gland transcriptome and proteome of the Anopheles stephensi mosquito". Insect
Biochemistry and Molecular Biology. 33 (7). Elsevier: 717–732. doi:10.1016/s0965-1748(03)00067-5.
ISSN 0965-1748. PMID 12826099.

Dash, A. P.; Adak, T.; Raghavendra, K.; Singh, O. P. (2007). "The biology and control of malaria vectors in
India". Current Science. 92 (11). Current Science Association + Indian Academy of Sciences: 1571–1578.
ISSN 0011-3891. JSTOR 24097721.

Malhotra, Prithvi Raj; Jatav, Prakash Chandra; Chauhan, Ram Singh (2000). "Surface morphology of the
egg of anopheles stephensi stephensi sensu stricto (diptera, culicidae)". Italian Journal of Zoology. 67
(2). Taylor & Francis: 147–151. doi:10.1080/11250000009356307. ISSN 1125-0003.

Sinka, M.E., Bangs, M.J., Manguin, S., Chareonviriyaphap, T., Patil, A.P., Temperley, W.H., Gething, P.
W., Elyazar, I.R.F., Kabaria, C.W., Harbach, R.E., & Hay, S.I. (2011). The dominant Anopheles vectors of
human malaria in the Asia-Pacific region: occurrence data, distribution maps and bionomic précis.
Parasites & Vectors, 4, 1-46.

Tikar, S. N.; Mendki, M.J.; Sharma, A. K.; Sukumaran, D.; Veer, Vijay; Prakash, Shri; Parashar, B. D.
(2011). "Resistance Status of the Malaria Vector Mosquitoes, Anopheles stephensi and Anopheles
subpictus Towards Adulticides and Larvicides in Arid and Semi-Arid Areas of India". Journal of Insect
Science. 11 (85). Entomological Society of America (OUP): 1–10. doi:10.1673/031.011.8501. ISSN 1536-
2442. PMC 3281435. PMID 21870971.

Gemuteerde mug moet malaria bestrijden Nederlandse Omroep Stichting 24 November 2015 (in Dutch)

Nolen, Stephanie; Negeri, Tiksa (29 September 2023). "An Invasive Mosquito Threatens Catastrophe in
Africa". The New York Times. Retrieved 18 February 2024.
 Rueda, Leopoldo M. (2007-12-18). "Global diversity of mosquitoes (Insecta: Diptera: Culicidae) in
freshwater". Hydrobiologia. 595 (1). Springer: 477–487. doi:10.1007/s10750-007-9037-x. ISSN 0018-
8158. S2CID 43415941.

Harbach, Ralph E. (2007-12-21). "The Culicidae (Diptera): a review of taxonomy, classification and
phylogeny". Zootaxa. 1668 (1). Magnolia Press: 591–638. doi:10.11646/zootaxa.1668.1.28. ISSN 1175-
5334. S2CID 17311770.

Manouchehri, AV; Javadian, E; Eshighy, N; Motabar, M (1976). "Ecology of Anopheles stephensi Liston in
southern Iran". Tropical and Geographical Medicine. 28 (3): 228–32. CiteSeerX 10.1.1.455.2712. ISSN
0041-3232. PMID 1006792.

Jeyabalan, D; Arul, N; Thangamathi, P (2003). "Studies on effects of Pelargonium citrosa leaf extracts on
malarial vector, Anopheles stephensi Liston". Bioresource Technology. 89 (2). Elsevier: 185–189.
doi:10.1016/s0960-8524(03)00036-1. ISSN 0960-8524. PMID 12699939.

"Anopheles stephensi". Invasive Species Compendium (ISC). CABI (Centre for Agriculture and Bioscience
International). 2019-11-22. Retrieved 2022-01-22.

Vanderberg, Jerome P. (2009). "Reflections". Vaccine. 27 (1). Edward Jenner Society + Japanese Society
for Vaccinology (Elsevier): 2–9. doi:10.1016/j.vaccine.2008.10.028. ISSN 0264-410X. PMC 2637529. PMID
18973784.

"Mosquitos, mosquito bite, mosquito transmitted diseases".

"Vector alert: Anopheles stephensi invasion and spread". WHO. 2019-08-26. Retrieved 2020-11-11.

Faulde, Michael K.; Rueda, Leopoldo M.; Khaireh, Bouh A. (2014). "First record of the Asian malaria
vector Anopheles stephensi and its possible role in the resurgence of malaria in Djibouti, Horn of Africa".
Acta Tropica. 139. Elsevier BV: 39–43. doi:10.1016/j.actatropica.2014.06.016. ISSN 0001-706X. PMID
25004439.

Carter, Tamar E.; Yared, Solomon; Gebresilassie, Araya; Bonnell, Victoria; Damodaran, Lambodhar;
Lopez, Karen; Ibrahim, Mohammed; Mohammed, Seid; Janies, Daniel (December 2018). "First detection
of Anopheles stephensi Liston, 1901 (Diptera: culicidae) in Ethiopia using molecular and morphological
approaches". Acta Tropica. 188: 180–186. doi:10.1016/j.actatropica.2018.09.001. ISSN 0001-706X. PMID
30189199.

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Anopheles subpictus

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Anopheles (Cellia) subpictus is a species complex of four species (A, B, C and D) of mosquito belonging to
the genus Anopheles.[1]

Anopheles subpictus

Scientific classification

Kingdom:

Animalia

Phylum:

Arthropoda

Class:

Insecta

Order:

Diptera

Superfamily:

Culicoidea

Family:

Culicidae

Subfamily:

Anophelinae

Genus:

Anopheles

Meigen 1818

Species:

Anopheles subpictus

(Grassi, 1899)
Synonyms

Anopheles error Theobald, 1903

Anopheles rossii Giles, 1899

Subgenus classification

edit

The genus Anopheles has been subdivided into seven subgenera – Anopheles, Baimaia, Cellia, Kerteszia,
Lophopodomyia, Nyssorhynchus and Stethomyia – based primarily on the number and positions of
specialized setae on the gonocoxites of the male genitalia. Within the subgenus but above the level of
species a number of additional taxonomic ranks have been created. While not officially recognised they
are in widespread use. The full classification of this species complex is:

Subgenus Cellia

Series Pyretophorus

Complex Anopheles subpictus

Bionomics

edit

The larvae are typically found in fresh water or rainwater pools often near urban peripheries or rural
houses. They are also found in barrow pits, buffalo wallows and artificial containers. Species B is the only
species restricted to coastal brackish-water habitats. Species A, C, and D generally found in fresh-water
sites.[2]

Geographic occurrence

edit

It has been reported in nineteen countries of the Oriental and Australasian Zones and is an important
vector of malaria in the Australasian Zone, Sulawesi, West Timor, South East Asia and Sri Lanka. It is
most common anopheline in most parts of the Indian subcontinent and has a widespread distribution
eastwards and southwards to Papua New Guinea, westwards to Iran and northwards to China.[3]

It has been reported from Afghanistan, Bangladesh, Cambodia, China, India, Indonesia, Iran, Malaysia,
Maldives, Mariana Islands, Myanmar, Nepal, Pakistan, Papua New Guinea, Philippines, Sri Lanka[4] and
Thailand.

Medical importance

edit
In addition to its role as a vector of malaria it is also a vector of some helminths (including Wuchereria
bancrofti) and arboviruses (including Japanese encephalitis virus, Sindbis virus and West Nile virus). It
has also been shown to transmit seadornaviruses but the importance of this observation is not yet clear.
[5]

References

edit

Surendran, Sinnathamby N.; Sarma, Devojit K.; Jude, Pavilupillai J.; Kemppainen, Petri; Kanthakumaran,
Nadarajah; Gajapathy, Kanapathy; Peiris, Lalanthika BS; Ramasamy, Ranjan; Walton, Catherine (2013).
"Molecular characterization and identification of members of the Anopheles subpictus complex in Sri
Lanka". Malaria Journal. 12: 304. doi:10.1186/1475-2875-12-304. PMC 3766661. PMID 24001126.

"Anopheles (Cellia) subpictus species complex". Malaria Atlas Project. Retrieved 30 January 2017.

"Anopheles (Cel.) subpictus". Walter Reed Biosystematics Unit. Retrieved 30 January 2017.

"An annotated checklist of mosquitoes of Sri Lanka" (PDF). Man and Biosphere Reserve of Sri Lanka.
Archived from the original (PDF) on 25 June 2016. Retrieved 30 January 2017.

Kumar, Ashwani; Hosmani, Rajeshwari; Jadhav, Shivaji; De Sousa, Trelita; Mohanty, Ajeet; Naik, Milind;
Shettigar, Adarsh; Kale, Satyajit; Valecha, Neena; Chery, Laura; Rathod, Pradipsinh K. (2016). "Anopheles
subpictus carry human malaria parasites in an urban area of Western India and may facilitate perennial
malaria transmission". Malaria Journal. 15: 124. doi:10.1186/s12936-016-1177-x. PMC 4769513. PMID
26919828.

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Anopheles sundaicus

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Anopheles sundaicus is a zoophilic mosquito of southeast Asia.

Anopheles sundaicus

Scientific classificationEdit this classification

Domain:
Eukaryota

Kingdom:

Animalia

Phylum:

Arthropoda

Class:

Insecta

Order:

Diptera

Family:

Culicidae

Genus:

Anopheles

Species:

A. sundaicus

Binomial name

Anopheles sundaicus

(Rodenwaldt, 1948)

Species complex

edit

Sukowati et al. 1999 finds there to be several cryptic species in an An. sundaicus complex. They use
protein electrophoresis to reveal populations defined by their characteristic allozymes.[1][2]

Hosts

edit

Hosts include Bubalus bubalis.[3] Kumari et al. 1993 finds An. sundaicus exclusively feeding on B. bubalis
on Car Nicobar Island, Moorhouse and Wharton 1965 finds the same in Malaysia and Gould et al. 1966
in Thailand.[3][4]
Control

edit

Insecticide

edit

Sargassum wightii and Bacillus thuringiensis var. israelensis can be combined to produce antifeedant,
insecticidal and growth inhibitor effects.[5]

Insecticide resistance

edit

Early studies of the inheritance of insecticide resistance were performed by Davidson 1957[6][7]: 97 [8]
and Soerono et al. 1965.[7]: 97 Davidson 1957 found An. sundaicus DDT resistance was provided by only
a single allele.[8]

References

edit

Krzywinski, Jaroslaw; Besansky, Nora J. (2003). "Molecular Systematics of Anopheles: From Subgenera
to Subpopulations". Annual Review of Entomology. 48 (1). Annual Reviews: 111–139.
doi:10.1146/annurev.ento.48.091801.112647. ISSN 0066-4170. PMID 12208816.

Manguin, Sylvie; Garros, C.; Dusfour, Isabelle; Harbach, Ralph E.; Coosemans, M. (2008). "Bionomics,
taxonomy, and distribution of the major malaria vector taxa of Anopheles subgenus Cellia in Southeast
Asia: An updated review". Infection, Genetics and Evolution. 8 (4). Elsevier: 489–503.
doi:10.1016/j.meegid.2007.11.004. ISSN 1567-1348. PMID 18178531.

Dusfour, Isabelle; Harbach, Ralph E.; Manguin, Sylvie (2004-10-01). "Bionomics and systematics of the
oriental Anopheles sundaicus complex in relation to malaria transmission and vector control" (PDF). The
American Journal of Tropical Medicine and Hygiene. 71 (4): 518–524. doi:10.4269/ajtmh.2004.71.518.
ISSN 0002-9637. PMID 15516652. S2CID 37387298.

Tananchai, Chatchai; Manguin, Sylvie; Bangs, Michael J.; Chareonviriyaphap, Theeraphap (2019).
"Malaria Vectors and Species Complexes in Thailand: Implications for Vector Control". Trends in
Parasitology. 35 (7). Cell Press: 544–558. doi:10.1016/j.pt.2019.04.013. ISSN 1471-4922. PMID
31182384. S2CID 184485189.

Song, Chenggang; Yang, Jian; Zhang, Mingzhe; Ding, Gang; Jia, Chengguo; Qin, Jianchun; Guo, Lanping
(2021-04-14). "Marine Natural Products: The Important Resource of Biological Insecticide". Chemistry &
Biodiversity. 18 (5). Wiley: e2001020. doi:10.1002/cbdv.202001020. ISSN 1612-1872. PMID 33855815.
S2CID 233243039.
 Davidson, G.; Mason, G. F. (1963). "Genetics of Mosquitoes". Annual Review of Entomology. 8 (1).
Annual Reviews: 177–196. doi:10.1146/annurev.en.08.010163.001141. ISSN 0066-4170. PMID
14025359.

Davidson, G. (1974). Genetic control of insect pests. London New York City: Academic Press. p. +158.
ISBN 978-0-323-16080-3. OCLC 761181727.

Brown, A. W. A. (1960). "Mechanisms of Resistance Against Insecticides". Annual Review of

Entomology. 5 (1). Annual Reviews: 301–326. doi:10.1146/annurev.en.05.010160.001505. ISSN 0066-
4170.

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Anopheles walkeri

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Anopheles walkeri is a species of mosquito found predominantly throughout the Mississippi River Valley,
[1] with its habitat ranging as far north as southern Quebec, Canada.[2] The eggs of A. walkeri are laid
directly on the water surface in freshwater swamp habitats.[3] Since its eggs are not resistant to
desiccation, this species is restricted to swampy regions with plenty of water. Anopheles walkeri, as with
many other anophelines, begins to become active later in the evening than most other mosquito species
in its range. This species becomes especially active late at night when in search of a blood meal.[4]
Feeding activity is affected greatly by environmental conditions within its microclimate. Wind, low
humidity and cool temperatures (around 10 °C or 50 °F and below), are all negatively correlated with
feeding aggression.[4]

Anopheles walkeri

Scientific classificationEdit this classification

Domain:

Eukaryota

Kingdom:

Animalia

Phylum:
Arthropoda

Class:

Insecta

Order:

Diptera

Family:

Culicidae

Genus:

Anopheles

Species:

A. walkeri

Binomial name

Anopheles walkeri

Theobald, 1901

Life cycle

edit

Anopheles walkeri mouthparts as compared with other Anopheles mouthparts and wings: 66, A.
punctipennis; 67, A. crucians; 68, A. quadrimaculatus; 69, A. walkeri.

Anopheles walkeri has a multivoltine life cycle.[3] It produces a hardy winter egg which differs
morphologically from the more vulnerable summer eggs by having enlarged floats on the dorsal side.[3]
By overwintering in egg form, this species is able to mature through one full larval generation before
hibernating adults of other species are able to become active. The multivoltine life cycle means this
species is active during both the swampy, open water conditions of early spring, as well as later in the
year after the swampland has become thickened with plant growth. It takes about 10 days to mature
through the larval stages and pupate, dependent on temperature and water conditions.[5] Adults will
typically mate within a few hours of emergence from their pupated form.[6] The females then begin to
seek out a blood meal to provide necessary protein to facilitate egg development. The female will then
rest while the eggs develop. Once mature, the eggs are oviposited and the female begins the process all
over again, feeding and laying for the 40 days or so that it lives for.[6]

Epidemiology
edit

Due to habitat preferences, coupled with particularly low rates of virus detection, Anopheles walkeri is
considered to be an unlikely vector of West Nile virus to humans.[7] In addition, it has not been shown
to demonstrate any capacity of transmitting avian malaria (Plasmodium circumflexum and P. polare).[8]
However, specimens of A. walkeri in the southern United States have been shown to harbor the human
infecting strains of malaria, Plasmodium vivax and P. falciparum, albeit infrequently (freq= <0.005).[9]

References

edit

Matheson, R.; Hurlbut, H. S. (1937). "Notes on Anopheles walkeri Theobald". American Journal of
Tropical Medicine and Hygiene. 17 (2): 237–243. doi:10.4269/ajtmh.1937.s1-17.237.

Ellis, R. A.; Chapman, H. C. (1980). "Mermithid parasites of Canadian anophelines". Mosquito News. 40
(1): 115–116.

Crans, W. J. (2004). "A classification system for mosquito life cycles: life cycle types for mosquitoes of
northeastern United States". Journal of Vector Ecology. 29 (1): 1–10. PMID 15266736.

Grimstad, P. R.; DeFoliart, G. R. (1975). "Mosquito nectar feeding in Wisconsin in relation to twilight and
microclimate". Journal of Medical Entomology. 11 (6): 691–698. doi:10.1093/jmedent/11.6.691. PMID
235648.

Shaman, J.; Steiglitz, M.; Stark, C.; Le Blancq S.; Cane, M. (2002). "Using a dynamic hydrology model to
predict mosquito abundances in flood and swamp water". Emerging Infectious Diseases. 8 (1): 8–13.
doi:10.3201/eid0801.010049. PMC 2730265. PMID 11749741.

Shaman, J.; Spiegelman, M.; Crane, M.; Stieglitz, M. (2006). "A hydrologically driven model of swamp
water mosquito population dynamics". Ecological Modelling. 194 (4): 395–404.
doi:10.1016/j.ecolmodel.2005.10.037.

Andreadis, T. G.; Anderson, J. F.; Vossbrinck, C. R.; Main, A. J. (2004). "Epidemiology of West Nile virus in
Connecticut: A five-year analysis of mosquito data 1999-2003". Vector-Borne and Zoonotic Diseases. 4
(4): 360–378. doi:10.1089/vbz.2004.4.360. PMID 15682518.

Meyer, C. L.; Bennett, G. F. (1976). "Observations on the sporogony of Plasmodium circumflexum Kikuth
and Plasmodium polare Manwell in New Brunswick". Canadian Journal of Zoology. 54 (2): 133–141.
doi:10.1139/z76-014. PMID 3278.

Bang, F. B.; Quimby, G. E.; Simpson, T. W. (1940). "Anopheles walkeri (Theobald) a wild-caught
specimen harboring malarial Plasmodia". Public Health Reports. 55 (3): 119–120. doi:10.2307/4583155.
JSTOR 4583155.
YELLOW FEVER MOSQUITO

Domain: Eukaryota

Kingdom: Animalia

Phylum: Arthropoda

Class: Insecta

Order: Diptera

Family: Culicidae

Genus: Aedes

Subgenus: Stegomyia

Species: A. aegypti

Binomial name

Aedes aegypti

(Linnaeus in Hasselquist, 1762) [1]

Subspecies[2][3]

Aedes aegypti aegypti

Aedes aegypti formosus

Global Aedes aegypti predicted distribution in 2015,

(blue=absent, red=present)

Synonyms[1]

Culex aegypti Linnaeus in Hasselquist, 1762

Culex fasciatus Fabricius, 1805

Culex bancrofti Skuse, 1889

Biology
Male (left) and female (center and right) Ae. aegyptiE.A. Goeldi, 1905

Aedes aegypti is a 4–7 millimetres (5⁄32–35⁄128 in) long, dark mosquito which can be recognized by white
markings on its legs and a marking in the form of a lyre on the upper surface of its thorax. Females are
larger than males. Microscopically females possess small palps tipped with silver or white scales, and
their antennae have sparse short hairs, whereas those of males are feathery. Aedes aegypti can be
confused with Aedes albopictus without a magnifying glass: the latter have a white stripe on the top of
the mid thorax.[4]

Males live off fruit[5] and only the female bites for blood, which she needs to mature her eggs. To find a
host, she is attracted to chemical compounds emitted by mammals, including ammonia,[6] carbon
dioxide,[7] lactic acid, and octenol.[8] Scientists at The United States Department of Agriculture (USDA)
Agricultural Research Service studied the specific chemical structure of octenol to better understand
why this chemical attracts the mosquito to its host and found the mosquito has a preference for "right-
handed" (dextrorotatory) octenol molecules.[9] The preference for biting humans is dependent on
expression of the odorant receptor AaegOr4.[10] The white eggs are laid separately into water and not
together, unlike most other mosquitoes, and soon turn black. The larvae feed on bacteria, grow over a
period of weeks until they reach the pupa stage.[5]

The lifespan of an adult Ae. aegypti is two to four weeks depending on conditions,[11] but the eggs can
be viable for over a year in a dry state, which allows the mosquito to re-emerge after a cold winter or
dry spell.[12]

Hosts

Mammalian hosts include domesticated horses, and feral and wild horses and equids more generally.
[13] As of 2009 birds were found to be the best food supply for Ae. aegypti among all taxa.[14]

Distribution

Ae. aegypti mosquito distribution in the United States, 2016

Aedes aegypti originated in Africa and was spread to the New World through slave trade,[15] but is now
found in tropical, subtropical and temperate regions[16] throughout the world.[17]Ae. aegypti's
distribution has increased in the past two to three decades worldwide, and it is considered to be among
the most widespread mosquito species.[18]

In 2016, Zika virus-capable mosquito populations have been found adapting for persistence in warm
temperate climates. Such a population has been identified to exist in parts of Washington, DC, and
genetic evidence suggests they survived at least the last four winters in the region. One of the study
researchers noted, "...some mosquito species are finding ways to survive in normally restrictive
environments by taking advantage of underground refugia".[19] As the world's climate becomes
warmer, the range of Aedes aegypti and a hardier species originating in Asia, the tiger mosquito Aedes
albopictus, which can expand its range to relatively cooler climates, will inexorably spread north and
south. Sadie Ryan of the University of Florida was the lead author in a 2019 study that estimated the
vulnerability of naïve populations in geographic regions that currently do not harbor vectors i.e., for Zika
in the Old World. Ryan's co-author, Georgetown University's Colin Carlson remarked,"Plain and simple,
climate change is going to kill a lot of people."[20] As of 2020, the Northern Territory Government
Australia and the Darwin City Council have recommended tropical cities initiate rectification programs to
rid their cities of potential mosquito breeding stormwater sumps.[21] A 2019 study found that
accelerating urbanization and human movement would also contribute to the spread of Aedes
mosquitoes.[22]

In continental Europe, Aedes aegypti is not established but it has been found in localities close to Europe
such as the Asian part of Turkey.[23] However, a single adult female specimen was found in Marseille
(Southern France) in 2018. On the basis of a genetic study and an analysis of the movements of
commercial ships, the origin of the specimen could be traced as coming from Cameroon, in Central
Africa.[23]

Genomics

In 2007, the genome of Aedes aegypti was published, after it had been sequenced and analyzed by a
consortium including scientists at The Institute for Genomic Research (now part of the J. Craig Venter
Institute), the European Bioinformatics Institute, the Broad Institute, and the University of Notre Dame.
The effort in sequencing its DNA was intended to provide new avenues for research into insecticides and
possible genetic modification to prevent the spread of virus. This was the second mosquito species to
have its genome sequenced in full (the first was Anopheles gambiae). The published data included the
1.38 billion base pairs containing the insect's estimated 15,419 protein-encoding genes. The sequence
indicates the species diverged from Drosophila melanogaster (the common fruit fly) about 250 million
years ago, and Anopheles gambiae and this species diverged about 150 million years ago.[24][25]
Matthews et al., 2018 finds A. aegypti to carry a large and diverse number of transposable elements.
Their analysis suggests this is common to all mosquitoes.[26]

Vector of disease

Aedes aegypti is a vector for transmitting numerous pathogens. According to the Walter Reed
Biosystematics Units as of 2022,[27] it is associated with the following 54 viruses and 2 species of
Plasmodium:

Aino virus (AINOV),

African horse sickness virus (AHSV),

Bozo virus (BOZOV),

Bussuquara virus (BSQV),

Bunyamwera virus (BUNV),

Catu virus (CATUV),

Chikungunya virus (CHIKV),

Chandipura vesiculovirus (CHPV),

Cypovirus (unnamed),

Cache Valley virus (CVV),

Dengue virus (DENV),

Eastern Equine Encephalitis virus (EEEV),

Epizootic hemorrhagic disease virus (EHDV),

Guaroa virus (GROV), Hart Park virus (HPV),

Ilheus virus (ILHV), Irituia virus (IRIV),

Israel Turkey Meningoencephalitis virus (ITV),

Japanaut virus (JAPV),

Joinjakaka (JOIV),

Japanese encephalitis virus (JBEV),

Ketapang virus (KETV),

Kunjin virus (KUNV),

La Crosse virus (LACV),

Mayaro virus (MAYV),

Marburg virus (MBGV),

Marco virus (MCOV),

Melao virus (MELV),

Marituba virus (MTBV),

Mount Elgon bat virus (MEBV),

Mucambo virus (MUCV),

Murray Valley Encephalitis virus (MVEV),

Navarro virus (NAVV),

Nepuyo virus (NEPV),

Nola virus (NOLV),

Ntaya virus (NTAV),

Oriboca virus (ORIV),

Orungo virus (ORUV),

Restan virus (RESV),

Rift Valley fever virus (RVFV),

Semliki Forest virus (SFV),

Sindbis virus (SINV),

Tahyna virus(TAHV),

Tsuruse virus (TSUV),

Tyuleniy virus (TYUV),

Venezuelan equine encephalitis virus (VEEV),

Vesicular stomatitis virus (Indiana serotype),

Warrego virus (WARV),

West Nile virus (WNV),

Wesselsbron virus (WSLV),

Yaounde virus (YAOV),

Yellow fever virus (YFV),

Zegla virus (ZEGV),

Zika virus,

And also Plasmodium gallinaceum and Plasmodium lophurae.

This mosquito also mechanically transmits some veterinary diseases. In 1952 Fenner et al., found it
transmitting the myxoma virus between rabbits[28] and in 2001 Chihota et al., the lumpy skin disease
virus between cattle.[28][29]

The yellow fever mosquito can contribute to the spread of reticular cell sarcoma among Syrian
hamsters.[30]
Bite prevention methods

The Centers for Disease Control and Prevention traveler's page on preventing dengue fever suggests
using mosquito repellents that contain DEET (N, N-diethylmetatoluamide, 20% to 30%). It also suggests:

Although Aedes aegypti mosquitoes most commonly feed at dusk and dawn, indoors, in shady areas, or
when the weather is cloudy, "they can bite and spread infection all year long and at any time of
day."[31][32]

Once a week, scrub off eggs sticking to wet containers, seal or discard them. The mosquitoes prefer to
breed in areas of stagnant water, such as flower vases, uncovered barrels, buckets, and discarded tires,
but the most dangerous areas are wet shower floors and toilet tanks, as they allow the mosquitos to
breed in the residence. Research has shown that certain chemicals emanating from bacteria in water
containers stimulate the female mosquitoes to lay their eggs. They are particularly motivated to lay eggs
in water containers that have the correct amounts of specific fatty acids associated with bacteria
involved in the degradation of leaves and other organic matter in water. The chemicals associated with
the microbial stew are far more stimulating to discerning female mosquitoes than plain or filtered water
in which the bacteria once lived.[33]

Wear long-sleeved clothing and long pants when outdoors during the day and evening.

Use mosquito netting over the bed if the bedroom is not air conditioned or screened, and for additional
protection, treat the mosquito netting with the insecticide permethrin.

Insect repellents containing DEET (particularly concentrated products) or p-menthane-3,8-diol (from

lemon eucalyptus) were effective in repelling Ae. aegypti mosquitoes, while others were less effective or
ineffective in a scientific study.[34] The Centers for Disease Control and Prevention article on "Protection
against Mosquitoes, Ticks, & Other Arthropods" notes that "Studies suggest that concentrations of DEET
above approximately 50% do not offer a marked increase in protection time against mosquitoes; DEET
efficacy tends to plateau at a concentration of approximately 50%".[35] Other insect repellents
recommended by the CDC include Picaridin (KBR 3023/icaridin), IR3535, and 2-undecanone.[36]

Population control efforts

Insecticides

Pyrethroids are commonly used.[37] This widespread use of pyrethroids and DDT has caused
Knockdown resistance (kdr) mutations. Almost no research has been done on the fitness implications.
studies by Kumar et al., 2009 on deltamethrin in India, Plernsub et al., 2013 on permethrin in Thailand,
by Jaramillo-O et al., 2014 on λ-cyhalothrin in Colombia, by Alvarez-Gonzalez et al., 2017 on
deltamethrin in Venezuela, are all substantially confounded. As of 2019, understanding of selective
pressure under withdrawal of insecticide is hence limited.[37]

Genetic modification
Ae. aegypti has been genetically modified to suppress its own species in an approach similar to the
sterile insect technique, thereby reducing the risk of disease. The mosquitoes, known as OX513A, were
developed by Oxitec, a spinout of Oxford University. Field trials in the Cayman Islands,[38] in Juazeiro,
[39][40] Brazil,[38] by Carvalho et al., 2015,[39][40] and in Panama[38] by Neira et al., 2014[39] have
shown that the OX513A mosquitoes reduced the target mosquito populations by more than 90%. This
mosquito suppression effect is achieved by a self-limiting gene that prevents the offspring from
surviving. Male modified mosquitoes, which do not bite or spread disease, are released to mate with the
pest females. Their offspring inherit the self-limiting gene and die before reaching adulthood—before
they can reproduce or spread disease. The OX513A mosquitoes and their offspring also carry a
fluorescent marker for simple monitoring. To produce more OX513A mosquitoes for control projects,
the self-limiting gene is switched off (using the Tet-Off system) in the mosquito production facility using
an antidote (the antibiotic tetracycline), allowing the mosquitoes to reproduce naturally. In the
environment, the antidote is unavailable to rescue mosquito reproduction, so the pest population is
suppressed.[41]

The mosquito control effect is nontoxic and species-specific, as the OX513A mosquitoes are Ae. aegypti
and only breed with Ae. aegypti. The result of the self-limiting approach is that the released insects and
their offspring die and do not persist in the environment.[42][43]

In Brazil, the modified mosquitoes were approved by the National Biosecurity Technical Commission for
releases throughout the country. Insects were released into the wild populations of Brazil, Malaysia, and
the Cayman Islands in 2012.[44][45] In July 2015, the city of Piracicaba, São Paulo, started releasing the
OX513A mosquitoes.[46][47] In 2015, the UK House of Lords called on the government to support more
work on genetically modified insects in the interest of global health.[48] In 2016, the United States Food
and Drug Administration granted preliminary approval for the use of modified mosquitoes to prevent
the spread of the Zika virus.[49]

Another proposed method consists in using radiation to sterilize male larvae so that when they mate,
they produce no progeny.[50] Male mosquitoes do not bite or spread disease.

Using CRISPR/Cas9 based genome editing to engineer the genome of Aedes aegypti genes like ECFP
(enhanced cyan fluorescent protein), Nix (male-determining factor gene), Aaeg-wtrw (Ae. aegypti water
witch locus), Kmo (kynurenine 3-monoxygenase), loqs (loquacious), r2d2 (r2d2 protein), ku70 (ku
heterodimer protein gene) and lig4 (ligase4) were targeted to modify the genome of Aedes aegypti. The
new mutant will become incapable of pathogen transmission or result in population control.[51]

Infection with Wolbachia

In 2016 research into the use of a bacterium called Wolbachia as a method of biocontrol was published
showing that invasion of Ae. aegypti by the endosymbiotic bacteria allows mosquitos to be resistant to
certain arboviruses such as dengue fever and Zika virus strains currently circulating.[52][53][54] In 2017
Alphabet, Inc. started the Debug Project to infect males of this species with Wolbachia bacteria,
interrupting the reproductive cycle of these animals.[55]
Fungus infection

Fungal species Erynia conica (from the family Entomophthoraceae) infects (and kills) two types of
mosquitos: Aedes aegypti and Culex restuans. Studies on the fungus have been carried out on its
potiential use as a biological control of the mosquitos.[56]

Taxonomy

The species was first named (as Culex aegypti) in 1757 by Fredric Hasselquist in his treatise Iter
Palaestinum.[57] Hasselquist was provided with the names and descriptions by his mentor, Carl
Linnaeus. This work was later translated into German and published in 1762 as Reise nach Palästina.[58]
Since the latter is an uncritical reproduction of the former, they are both considered to antedate the
starting point for zoological nomenclature in 1758. Nonetheless, the name Aedes aegypti was frequently
used, starting with H. G. Dyar in 1920.[[]]

Ae. aegypti feeding on a human

To stabilise the nomenclature, a petition to the International Commission on Zoological Nomenclature

was made by P. F. Mattingly, Alan Stone, and Kenneth L. Knight in 1962.[59] It also transpired that,
although the name Aedes aegypti was universally used for the yellow fever mosquito, Linnaeus had
actually described a species now known as Aedes (Ochlerotatus) caspius.[59] In 1964, the commission
ruled in favour of the proposal, validating Linnaeus' name, and transferring it to the species for which it
was in general use.[60]

The yellow fever mosquito belongs to the tribe Aedini of the dipteran family Culicidae and to the genus
Aedes and subgenus Stegomyia. According to one recent analysis, the subgenus Stegomyia of the genus
Aedes should be raised to the level of genus.[61] The proposed name change has been ignored by most
scientists;[62] at least one scientific journal, the Journal of Medical Entomology, has officially
encouraged authors dealing with aedile mosquitoes to continue to use the traditional names, unless
they have particular reasons for not doing so.[63] The generic name comes from the Ancient Greek
ἀηδής, aēdēs, meaning "unpleasant"[64] or "odious".

Subspecies

Two subspecies are commonly recognized:

Aedes aegypti subsp. aegypti[2][3]

Aedes aegypti subsp. formosus[2][3]

This classification is complicated by the results of Gloria-Soria et al., 2016. Although confirming the
existence of these two major subspecies, Gloria-Sora et al. finds greater worldwide diversity than
previously recognized and a large number of distinct populations separated by various geographic
factors.[2][3]

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Currently Circulating Zika Virus Isolates in Brazilian Aedes aegypti Mosquitoes". Cell Host & Microbe. 19
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Hancock, Penelope A.; White, Vanessa L.; Callahan, Ashley G.; Godfray, Charles H. J.; Hoffmann, Ary A.;
Ritchie, Scott A.; Clough, Yann (June 2016). "Density ‐dependent population dynamics in Aedes aegypti
slow the spread of wMel Wolbachia". Journal of Applied Ecology. 53 (3): 785–793.
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Utarini, Adi; Indriani, Citra; Ahmad, Riris A.; Tantowijoyo, Warsito; Arguni, Eggi; Ansari, M. Ridwan;
Supriyati, Endah; Wardana, D. Satria; Meitika, Yeti; Ernesia, Inggrid; Nurhayati, Indah; Prabowo,
Equatori; Andari, Bekti; Green, Benjamin R.; Hodgson, Lauren; Cutcher, Zoe; Rancès, Edwige; Ryan, Peter
A.; O'Neill, Scott L.; Dufault, Suzanne M.; Tanamas, Stephanie K.; Jewell, Nicholas P.; Anders, Katherine
L.; Simmons, Cameron P. (10 June 2021). "Efficacy of Wolbachia-Infected Mosquito Deployments for the
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Erynia conica (Entomophthorales: Entomophthoraceae)". J Am Mosq Control Assoc. 8 (4): 367–71. PMID
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proposed validation and interpretation under the plenary powers of the species so named. Z.N.(S.)
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doi:10.3201/eid2210.ET2210. PMC 5038420.

ASIAN TIGER MOSQUITO

Aedes albopictus (Stegomyia albopicta), from the mosquito (Culicidae) family, also known as the (Asian)
tiger mosquito or forest mosquito, is a mosquito native to the tropical and subtropical areas of
Southeast Asia. In the past few centuries, however, this species has spread to many countries through
the transport of goods and international travel.[1] It is characterized by the white bands on its legs and
body.

Aedes albopictus

Scientific classification

Domain: Eukaryota

Kingdom: Animalia

Phylum: Arthropoda

Class: Insecta

Order: Diptera

Family: Culicidae

Genus: Aedes

Subgenus: Stegomyia

Species: A. albopictus

Binomial name

Aedes albopictus

(Skuse, 1894)

Synonyms

Culex albopictus Skuse, 1894

This mosquito has become a significant pest in many communities because it closely associates with
humans (rather than living in wetlands), and typically flies and feeds in the daytime in addition to at
dusk and dawn. The insect is called a tiger mosquito for its striped appearance, which resembles that of
the tiger. Ae. albopictus is an epidemiologically important vector for the transmission of many viral
pathogens, including the yellow fever virus, dengue fever, and Chikungunya fever,[2] as well as several
filarial nematodes such as Dirofilaria immitis.[3] Aedes albopictus is capable of hosting the Zika virus[4]
[5] and is considered a potential vector for Zika transmission among humans.

Description

Tiger mosquito leg at the microscope 200x

Name and systematics

In 1894, a British-Australian entomologist, Frederick A. Askew Skuse, was the first to scientifically
describe the Asian tiger mosquito, which he named Culex albopictus (lat. culex "gnat", "midge" and
albopictus "white-painted").[6][7] Later, the species was assigned to the genus Aedes (gr. άηδής,
"unpleasant")[8] and referred to as Aedes albopictus.[9] Like the yellow fever mosquito, it belongs to
the subgenus Stegomyia (Gr. στέγος, "covered, roofed", referring to the scales that completely cover the
dorsal surface in this subgenus, and μυία, "fly") within the genus Aedes.[10] In 2004, scientists explored
higher-level relationships and proposed a new classification within the genus Aedes and Stegomyia was
elevated to the genus level, making Aedes albopictus now Stegomyia albopicta. This is, however, a
controversial matter, and the use of Stegomyia albopicta versus Aedes albopictus is continually debated.
[11][12][13]

Characteristics

The adult Asian tiger mosquito is less than 10 mm (0.39 in) long from end to end with a striking white
and black pattern.[6][14][15] The variation of the body size in adult mosquitoes depends on the density
of the larval population and food supply within the breeding water. Since these circumstances are
seldom optimal, the average body size of adult mosquitoes is considerably smaller than 10 mm. For
example, the average length of the abdomen was calculated to be 2.63 mm (0.104 in), the wings 2.7 mm
(0.11 in), and the proboscis 1.88 mm (0.074 in).[16]

The males are roughly 20% smaller than the females, but they are morphologically very similar.
However, as in all mosquito species, the antennae of the males in comparison to the females are
noticeably bushier and contain auditory receptors to detect the characteristic whine, almost inaudible to
humans, of the female. The maxillary palps of the males are also longer than their proboscis, whereas
the females' maxillary palps are much shorter. (This is typical for the males of the Culicinae.) In addition,
the tarsus of the hind legs of the males is more silvery. Tarsomere IV is roughly 75% silver in the males
whereas the females' is only about 60% silver.[[]]

The other characteristics do not differentiate between sexes. A single silvery-white line of tight scales
begins between the eyes and continues down the dorsal side of the thorax. This characteristic marking is
the easiest and surest way to identify the Asian tiger mosquito.[[]]

The proboscis is dark colored, the upper surface of the end segment of the palps is covered in silvery
scales, and the labium does not feature a light line on its underside. The compound eyes are distinctly
separated from one another. The scute, the dorsal portion of an insect's thoracic segment, is black
alongside the characteristic white midline. On the side of the thorax, the scutellum, and the abdomen
are numerous spots covered in white-silvery scales.

Such white-silvery scales can also be found on the tarsus, particularly on the hind legs that are
commonly suspended in the air. The bases of tarsomeres I through IV have a ring of white scales,
creating the appearance of white and black rings. On the forelegs and middle legs, only the first three
tarsomeres have the ring of white scales, whereas tarsomere V on the hind legs is completely white. The
femur of each leg is also black with white scales on the end of the "knee". The femora of the middle legs
do not feature a silver line on the base of the upper side, whereas, the femora on the hind legs have
short white lines on base of the upper side. The tibiae are black on the base and have no white scales.
[[]]

The terga on segments II through VI of the abdomen are dark and have an almost triangular silvery-
white marking on the base that is not aligned with the silvery bands of scales on the ventral side of the
abdomen. The triangular marking and the silvery band are only aligned on abdominal segment VII. The
transparent wings have white spots on the base of the costae. With older mosquito specimens, the
scales could be partially worn off, making these characteristics not stand out as much.[14][16]

As with other members of the mosquito family, the female is equipped with an elongated proboscis that
she uses to collect blood to feed her eggs. The Asian tiger mosquito has a rapid bite and an agility that
allows it to escape most attempts by people to swat it. By contrast, the male member of the species
primarily feeds on nectar and does not bite.

The female lays her eggs near water, not directly into it as other mosquitoes do, but typically near a
stagnant pool. However, any open container containing water will suffice for larvae development, even
with less than one US fl oz (30 ml) of water. It can also breed in running water, so stagnant pools of
water are not its only breeding sites. It is more likely to lay eggs in water sources near flowers than in
water sources without flowers. It has a short flight range (less than 200 m (220 yd)), so breeding sites
are likely to be close to where this mosquito is found.[17][18]

Other mosquito species may be visually confused with the tiger mosquito. Comparison with approved
pictures is the best way to determine the species with certainty.[19] Behavioral cues like almost-silent
flight and difficulty in catching combined with knowledge of the range of local endemic mosquitoes may
also aid this process.

Similar species

Some mosquitoes in North America, such as Ochlerotatus canadensis, have a similar leg pattern. In
North and South America, Ae. albopictus can be distinguished from Aedes taeniorhynchus since only Ae.
albopictus has back markings.

In Europe, the mosquito Culiseta annulata, which is very common, but does not occur in high densities,
can be mistaken for an Asian tiger mosquito because of its black-and-white-ringed legs. However, this
species is missing the distinctive white line that runs from the middle of its head and down the thorax. It
is also considerably larger than Ae. albopictus, is not black and white, but rather beige and grey striped,
and has wings with noticeable veins and four dark, indistinct spots. The Tree Hole mosquito or Aedes
geniculatus – a native to Europe and North Africa – has also been mistaken for Ae. albopictus. This is
because the Tree Hole mosquito has very white scales on a very similar body.[20]

In the eastern Mediterranean area, Ae. albopictus species can be mistaken for Aedes cretinus, which
also belongs to the subgenus Stegomyia and uses similar breeding waters. Aedes cretinus also has a
white stripe on the scute, but it ends shortly before the abdomen, and also has two additional stripes to
the left and right of the middle stripe. So far Aedes cretinus is only located in Cyprus, Greece, North
Macedonia, Georgia and Turkey.[21]

In Asia, the Asian tiger mosquito can be mistaken for other members of the subgenus Stegomyia,
particularly the yellow fever mosquito Aedes aegypti (the most prevalent species in the tropics and
subtropics), because both species display a similar black and white pattern. It can be hard to distinguish
Ae. albopictus from the closely related Aedes scutellaris (India, Indonesia, Papua New Guinea, and the
Philippines), Aedes pseudoalbopictus (India, Indonesia, Malaysia, Myanmar, Nepal, Taiwan, Thailand,
and Vietnam) and Aedes seatoi (Thailand).[14][22]

Diet and host location

Bloated female at the end of a meal

Like other mosquito species, only the females require a blood meal to develop their eggs. Apart from
that, they feed on nectar and other sweet plant juices just as the males do. In regards to host location,
carbon dioxide and organic substances produced from the host, humidity, and optical recognition play
important roles.[[]]

The search for a host takes place in two phases. First, the mosquito exhibits a nonspecific searching
behavior until it perceives host stimulants, whereupon it secondly takes a targeted approach.[23] For
catching tiger mosquitoes with special traps, carbon dioxide and a combination of chemicals that
naturally occur in human skin (fatty acids, ammonia, and lactic acid) are the most attractive.[24]

The Asian tiger mosquito particularly bites in forests during the day, so has been known as the forest day
mosquito. Depending upon region and biotype, activity peaks differ, but for the most part, they rest
during the morning and night hours. They search for their hosts inside and outside human dwellings, but
are particularly active outside. The size of the blood meal depends upon the size of the mosquito, but it
is usually around 2 μl. Their bites are not necessarily painful, but they are more noticeable than those
from other kinds of mosquitoes. Tiger mosquitoes generally tend to bite a human host more than once if
they are able to.[23][25]

Ae. albopictus also bites other mammals besides humans, as well as birds.[23][25] The females are
always on the search for a host and are persistent but cautious when it comes to their blood meal and
host location. Their blood meal is often broken off before enough blood has been ingested for the
development of their eggs, so Asian tiger mosquitoes bite multiple hosts during their development cycle
of the egg, making them particularly efficient at transmitting diseases. The mannerism of biting diverse
host species enables the Asian tiger mosquito to be a potential bridge vector for certain pathogens that
can jump species boundaries, for example the West Nile virus.[[]]

Natural enemies

Primarily, other mosquito larvae, flatworms, swimming beetles, fungi, ciliates, paramecia, protozoans
which act as parasites, predatory copepods, and spiders are natural enemies of the larval stage of Asian
tiger mosquitoes.[[]]
Toxorhynchites speciosus larvae (an adult is shown here) feed on the larvae of Aedes albopictus.

Toxorhynchites larvae, a mosquito genus that does not suck blood, feeds upon other mosquito larvae
and are often found with tiger mosquito larvae. Flatworms and small swimming beetles are considered
natural predators.[25]

Fungi from the genus Coelomomyces (order Blastocladiales) develop inside the visceral cavity of
mosquito larvae. The species Coelomomyces stegomyiae was first found on the Asian tiger mosquito.
[25]

Paramecia, or ciliates, can also affect Ae. albopictus larvae, and the first detected species was
Lambornella stegomyiae (Hymenostomatida: Tetrahymenidae).[25] The virulence, mortality rate, and
subsequent possibilities of Lambornella being implemented as a biological remedy to control Ae.
albopictus, however, has conflicting views.[26][27]

Sporozoans of the genus Ascogregarina (Lecudinidae) infect the larval stage of mosquitoes. The species
Ascogregarina taiwanensis was found in Asian tiger mosquitoes.[25] When the adult mosquitoes emerge
from their pupal case, they leave the infectious intermediary stage of parasites in the water and close off
the infection cycle. Infected adults are generally smaller than non-infected adults and have an
insignificantly higher mortality rate; therefore, food supply and larval density apparently play a role. In
competitive situations, an infection with sporozoans can also reduce the biological fitness of other
uninfected mosquitoes. However, the use of the parasites as an effective biological remedy to control
mosquito populations is implausible because the host must reach the adult stage for the transmission of
the parasites.[28]

Though they do not commonly occur in the natural habitats of Asian tiger mosquitoes, predatory
copepods from the family Cyclopidae seem to willingly feed on them given the opportunity.[25]
Relatives of different genera could therefore present a possibility in the control of tiger mosquitoes.[29]

Predators of adult Ae. albopictus in Malaysia include various spider species. Up to 90% of the gathered
spiders from rubber plantations and a cemetery fed upon Asian tiger mosquitoes. Whether the spiders
would have an effect on the mosquito population is still unclear. Tiger mosquitoes were abundantly
present despite the existence of the spiders.[30]

Distribution

Estimated distribution of Ae. albopictus in the United States, CDC 2016

Climatic adaptations

Ae. albopictus eggs

The Asian tiger mosquito originally came from Southeast Asia. In 1966, parts of Asia and the island
worlds of India and the Pacific Ocean were denoted as the area of circulation for the Asian tiger
mosquito.[31] Ae. albopictus as a native to tropical and subtropical regions with warm and humid
climate, is active all year long; however, it has been adapting successfully to cooler, temperate regions,
where they hibernate over winter. Eggs from strains in the temperate zones are more tolerant to the
cold than ones from warmer regions.[32][33] The species can even tolerate snow and temperatures
under freezing. Adult tiger mosquitoes can survive throughout winter in suitable microhabitats.[34]

Invasive species

Since the mid-1960s, the tiger mosquito has spread to Europe, the Americas, the Caribbean, Africa, and
the Middle East. As of 2008 Ae. albopictus was one of the 100 world's worst invasive species according
to the Global Invasive Species Database.[35]

As of 2006, Ae. albopictus was not native to Australia and New Zealand.[36][37] The species was
introduced there multiple times, but has yet to establish itself. This is due to the well-organized
entomological surveillance programs in the harbors and airports of these countries. Nevertheless, as of
2006 it has become domestic on the islands in the Torres Strait between Queensland, Australia, and
New Guinea.[38]

In Europe, Asian tiger mosquitos first emerged in Albania in 1979, introduced through a shipment of
goods from China. In 1990–1991, they were most likely brought to Italy in used tires from Georgia (USA),
and since then have spread throughout the entire mainland of Italy, as well as parts of Sicily and
Sardinia. Since 1999, they have established themselves on the mainland of France, primarily southern
France. In 2002, they were also discovered in a vacation town on the island of Corsica, but did not
completely establish themselves there until 2005. In Belgium, they were detected in 2000 and 2013,[39]
in 2001 in Montenegro, 2003 in Canton Ticino in southern Switzerland, and Greece, 2004 in Spain and
Croatia, 2005 in the Netherlands and Slovenia, 2006 in Bosnia and Herzegovina[1] and 2022 in Cyprus.
[40] In the fall of 2007, the first tiger mosquito eggs were discovered in Rastatt (Baden-Wuerttemberg,
Germany).[41] Shortly before, they were found in the northern Alps of Switzerland in Canton Aargau.
[42] since 2010, it has also been sighted increasingly in Malta during summer.[[]] In September 2016,
Public Health England found eggs, though no mosquitos, in a lorry park at Folkestone service station on
the M20, near Westenhanger, which is 6 miles West of the Eurotunnel.[43] The Swiss Autobahns are
especially of concern.[44] Governments and universities in Switzerland cooperate every year to monitor
the invasion using traps at Autobahn rest stations, and also at airports and commercial hubs.[44]

In the United States, this species invaded the Southern United States in the 1980s and rapidly spread
northward into novel climate compared to its native range.[45] It was initially found in 1983 in
Memphis, Tennessee.[46] then at the Port of Houston in a 1985 shipment of used tires,[47] and spread
across the South up the East Coast to become prevalent in the Northeast.[48] It was not discovered in
Southern California until 2001, then eradicated for over a decade; however, by 2011, it was again being
found in Los Angeles County traps, then over the next two years expanded its range to Kern County and
San Diego County.[49][50][51] As of 2013, North American land favoring the environmental conditions
of the Asian tiger mosquito was expected to more than triple in size in the coming 20 years, especially in
urban areas.[52] As of 2017 Aedes albopictus mosquitoes have been identified in 1,368 counties in 40
U.S states.[53][54] A 2019 study in Nature Microbiology that modeled expansion of Aedes albopictus
due to climate change, urbanization, and human movement found that the species would likely continue
to spread throughout the coming decades.[55]

In Latin America, the Asian tiger mosquito was first discovered 1986 in Brazil[56] and in 1988 in
Argentina and Mexico,[57] as well. Other parts of Latin America where the Asian tiger mosquito was
discovered are the Dominican Republic in 1993, Bolivia, Cuba, Honduras, and Guatemala in 1995, El
Salvador in 1996, Paraguay in 1999, Panama in 2002, and Uruguay and Nicaragua in 2003.[58]

In Africa, the species was first detected in 1990 in South Africa.[59] In Nigeria, it has been domestic since
at least 1991.[60] It spread to Cameroon in 1999/2000,[61] to the Bioko Island of Equatorial Guinea in
2001,[62] and to Gabon in 2006.[63]

In the Middle East, the species was detected in Lebanon in 2003 and in Syria in 2005; the first record in
Israel was published in 2003.[64]

Competition with established species

Ae. albopictus

Ae. albopictus can outcompete and even eradicate other species with similar breeding habitats from the
very start of its dispersal to other regions and biotopes.[65] In Kolkata, for example, it was observed in
the 1960s that egg depositing containers were being settled by the Asian tiger mosquito in city districts
where the malaria mosquito (genus Anopheles) and yellow fever mosquito (Aedes aegypti) had both
been eliminated by the application of DDT.[66] This may be because primarily the inner walls of the
houses were treated with DDT to kill the mosquitoes resting there and fight the malaria mosquito. The
yellow fever mosquito also lingers particularly in the inside of buildings and would have been also
affected. The Asian tiger mosquito rests in the vicinity of human dwellings would therefore have an
advantage over the other two species. In other cases where the yellow fever mosquito was repressed by
the Asian tiger mosquito, for instance in Florida, this explanation does not fit.[67][68] Other hypotheses
include competition in the larval breeding waters, differences in metabolism and reproductive biology,
or a major susceptibility to sporozoans (Apicomplexa).[69]

Another species which was suppressed by the migrating Ae. albopictus was Ae. guamensis in Guam.[70]

The Asian tiger mosquito is similar, in terms of its close socialization with humans, to the common house
mosquito (Culex pipiens). Among other differences in their biology, Culex pipiens prefers larger breeding
waters and is more tolerant to cold. In this respect, no significant competition or suppression between
the two species likely occurs.[69]

A possible competition among mosquito species that all lay their eggs in knotholes and other similar
places (Ae. cretinus, Ae. geniculatus, and Anopheles plumbeus) has yet to be observed.[[]]

In Europe, the Asian tiger mosquito apparently covers an extensive new niche. This means that no
native, long-established species conflict with the dispersal of Ae. albopictus.[[]]
Role as disease vector

For humans

Ae. albopictus is known to transmit pathogens and viruses, such as the yellow fever virus, dengue fever,
Chikungunya fever,[2] and Usutu virus.[71] There is some evidence supporting the role of Ae. albopictus
in the transmission of Zika virus, which is primarily transmitted by the related Ae. aegypti.[5]

The Asian tiger mosquito was responsible for the Chikungunya epidemic on the French Island La Réunion
in 2005–2006. By September 2006, an estimated 266,000 people were infected with the virus, and 248
fatalities occurred on the island.[72] The Asian tiger mosquito was also the transmitter of the virus in the
first outbreak of Chikungunya fever on the European continent. This outbreak occurred in the Italian
province of Ravenna in the summer of 2007, and infected over 200 people.[73][74] Evidently, mutated
strains of the Chikungunya virus are being directly transmitted through Ae. albopictus particularly well
and in such a way that another dispersal of the disease in regions with the Asian tiger mosquito is
feared.[75]

On the basis of experimental evidence and probability estimates, the likelihood of mechanical or
biological transmission of HIV by insects is virtually nonexistent.[76]

For animals

The tiger mosquito is relevant to veterinary medicine. For example, tiger mosquitoes are transmitters of
Dirofilaria immitis, a parasitic roundworm that causes heart failure in dogs and cats.[77]

For arthropods

Wolbachia infection are the most common infection in arthropods today, and over 40% of arthropods
have contracted it.[78] Wolbachia can be transmitted from parent to offspring or between breeding
individuals. Wolbachia is easily transmitted within the Ae. albopictus mosquito due to the effects it has
on fecundity in females.[79] Once female Asian tiger mosquitos have contracted the infection, they
produce more eggs, give birth more frequently, and live longer than uninfected females. In this way,
Wolbachia provides a fitness advantage to the infected females and prevents uninfected females from
reproducing. This allows control of the spread of diseases that many species carry by suppressing
reproduction of the individuals with the harmful disease, but without the Wolbachia infection.
Wolbachia can also be used to transfer certain genes into the population to further control the spread of
diseases.[80]

Cytoplasmic incompatibility

In the natural environment, Wolbachia and the Asian tiger mosquito are in a symbiotic relationship, so
both species benefit from each other and can evolve together. The relationship between Wolbachia and
its host might not have always been mutualistic, as Drosophila populations once experienced decreased
fecundity in infected females, suggesting that Wolbachia evolved over time so that infected individuals
would actually reproduce much more.[81] The mechanism by which Wolbachia is inherited through
maternal heredity is called cytoplasmic incompatibility.[79] This changes the gamete cells of males and
females, making some individuals unable to mate with each other. Although little is known about why
cytoplasmic incompatibility exists, Wolbachia infection creates a fitness advantage for infected females,
as they can mate with either infected or uninfected males. Despite this, infected males cannot
reproduce with uninfected females. Therefore, over time, a population exposed to Wolbachia
transitions from a few infected individuals to all individuals becoming infected, as the males that cannot
reproduce successfully do not contribute to future generations. This is called population replacement,
where the population's overall genotype is replaced by a new genotype. This shows how populations of
Asian tiger mosquitoes can vary in number of Wolbachia-infected individuals, based on how often the
infection is transmitted.[82] Due to Wolbachia's ability to transmit from one host to the next, it can
change the average genotype of a population, potentially reducing the population's gene flow with
other nearby populations.[[]]

Unidirectional cytoplasmic incompatibility

This type of cytoplasmic incompatibility where an infected male cannot reproduce successfully with an
uninfected female is called unidirectional cytoplasmic incompatibility. It occurs because Wolbachia
modifies the paternal chromosomes during sperm development, leading to complications for these
offspring during embryonic development.[83]

Bidirectional cytoplasmic incompatibility

Also, bidirectional cytoplasmic incompatibility occurs when an infected male carrying one strain of
Wolbachia reproduces with an infected female carrying a different strain of Wolbachia. This also results
in failed reproduction. Bidirectional cytoplasmic incompatibility also has evolutionary implications for
populations of Ae. albopictus and other vectors of the infection.[84] This is because bidirectional
cytoplasmic incompatibility in Wolbachia creates unviable offspring, reducing gene flow between two
populations, which can eventually lead to speciation.[[]]

Control and suppression

Litter in roadside ditches makes an ideal breeding ground for the Asian tiger mosquito.

Ae. albopictus is very difficult to suppress or to control due to its remarkable ability to adapt to various
environments, its close contact with humans, and its reproductive biology.[[]]

An Ovitrap, a tool for the detection of Asian tiger mosquitoes: Their presence is confirmed through the
eggs they lay on the wooden paddle. The brown granules in the water are a Bti preparation that kills
hatching mosquito larvae.

The containment of infestations is generally effected by public health services through area-wide
integrated control plans, which aim to reduce the nuisance perceived by populations and the risks of
viraemic transmission. Such plans consist of different activities that include entomological surveillance,
larvicide treatments in public and private areas, information campaigns, and treatments against adult
mosquitoes in the zones affected by suspected cases of transmissible viruses.[85]
Efficient monitoring or surveillance is essential to prevent the spread and establishment of this species.
In addition to the monitoring of ports, warehouses with imported plants, and stockpiles of tires, rest
areas on highways and train stations should be monitored with appropriate methods.[86]

Stagnant water removal

The control of Asian tiger mosquitoes begins with destroying the places where they lay their eggs, which
are never far from where people are being bitten, since they are weak fliers, with only about a 180-
metre (590 ft) lifetime flying radius. Puddles that last more than three days, sagging or plugged roof
gutters, old tires holding water, litter, and any other possible containers or pools of standing water
should be drained or removed. Bird baths, inlets to sewers and drainage systems holding stagnant
water, flower pots, standing flower vases, knotholes, and other crevices that can collect water should be
filled with sand or fine gravel to prevent mosquitoes from laying their eggs in them.[[]]

Any standing water in pools, catchment basins, etc., that cannot be drained, or dumped, can be
periodically treated with properly labeled insecticides or Bacillus thuringiensis israelensis (Bti), often
formed into doughnut-shaped "mosquito dunks". Bti produces toxins which are effective in killing larvae
of mosquitoes and certain other dipterans, while having almost no effect on other organisms. Bti
preparations are readily available at farm, garden, and pool suppliers.[[]]

Flowing water will not be a breeding spot,[contradictory] and water that contains minnows is not usually
a problem, because the fish eat the mosquito larvae.[[]]

Dragonflies

Dragonflies are also an excellent method of control. Dragonfly larvae eat mosquito larvae in the water,
and adults snatch adult mosquitoes as they fly.[[]]

Ovitraps

In any case, an efficient surveillance is essential to monitor the presence of tiger mosquitoes and the
effect of control programs. Ovitraps are normally used for the monitoring of Ae. albopictus. They are
black water containers with floating Styrofoam blocks or small wooden paddles that are in contact with
the surface of the water. Female tiger mosquitoes lay their eggs on these surfaces. Through the
identification of these eggs or of the larvae that hatch from these eggs in the laboratory, the presence
and abundance of mosquito species can be estimated. Versions of these traps with an adhesive film
(sticky traps) that catch the egg-depositing mosquitoes make the analysis much easier and quicker, but
are more complicated in terms of handling.[87][88] The results of ovitraps are often variable and
depend on the availability of alternative egg-depositing waters. Due to this, it is best to use them in
large numbers and in conjunction with other monitoring methods.[[]]

To date, few effective traps for adult Asian tiger mosquitoes are available. Those traps that catch other
species of mosquitoes do not catch tiger mosquitoes efficiently. A form of an ovitrap called a lethal
ovitrap mimics the breeding site for Ae. albopictus just like the monitoring tool, but it has the added
benefit of containing chemicals that are toxic to the mosquitoes when they enter, but do not harm
humans. These traps have had success in some countries to control Aedes mosquito populations.[89] A
new trap type has now been shown to catch significant numbers of Ae. albopictus.[90][91] This device,
with the help of a ventilator, produces an upward air current of ammonia, fatty acids, and lactic acids
that takes a similar form and smell of a human body. With the addition of carbon dioxide, the efficacy of
the trap is increased. This means a suitable tool is available for trapping adult tiger mosquitoes, and for
example, examining the existence of viruses in the trapped mosquitoes. Previously, the mosquitoes had
to be collected from volunteers to be studied, which is ethically questionable, especially during
epidemics. Recent research also indicates this trap type may also have a use as a control tool; in a study
in Cesena, Italy, the number of biting tiger mosquitoes was reduced in places where traps were installed.
[92]

An amino acid substitution mutation – F1534C – is overwhelmingly the most common voltage-gated
sodium channel in A. albopictus in Singapore.[93] This channel being the target of pyrethroids,[93] this is
suspected to be a knockdown resistance (kdr) mutation,[93] and that that is the reason for its
prevalence.[93]

Public health benefits

Although the Wolbachia infection is prevalent in arthropod species, especially the Asian tiger mosquito,
it is a useful mechanism for inhibiting the spread of dengue.[94] Ae. aegypti individuals, a close relative
of Ae. albopictus, with an artificial Wolbachia infection, cannot transmit dengue, an infectious virus, but
they can pass on the Wolbachia infection to other populations. This could lead to many more discoveries
in disease control for Ae. albopictus and other mosquito species.[94] In addition, due to the cytoplasmic
incompatibility caused by Wolbachia, the artificial infection of males can serve as a biological control as
they are unable to reproduce successfully with uninfected females (unidirectional CI).[83] When
artificially infected males are unable to reproduce, the population size can be controlled, thereby
reducing the transmission of the harmful disease of interest. Artificial infection of males is achieved by
the removal of cytoplasm from infected oocytes, which is then transferred into embryos prior to the
blastoderm stage.[[]]

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Facchinelli, L; Valerio, L; Pombi, M; Reiter, P; Costantini, C; Della Torre, A (2007). "Development of a

novel sticky trap for container-breeding mosquitoes and evaluation of its sampling properties to monitor
urban populations of Aedes albopictus". Medical and Veterinary Entomology. 21 (2): 183–95.
doi:10.1111/j.1365-2915.2007.00680.x. PMID 17550438. S2CID 4237472.

Gama, Renata A.; Silva, Eric M.; Silva, Ivoneide M.; Resende, Marcelo C.; Eiras, Álvaro E. (2007).
"Evaluation of the sticky MosquiTRAP for detecting Aedes (Stegomyia) aegypti (L.) (Diptera: Culicidae)
during the dry season in Belo Horizonte, Minas Gerais, Brazil". Neotropical Entomology. 36 (2): 294–302.
doi:10.1590/S1519-566X2007000200018. PMID 17607465.

Zeichner, B. C.; Debboun, M. (2011). "The lethal ovitrap: A response to the resurgence of dengue and
chikungunya". U.S. Army Medical Department Journal: 4–11. PMID 21805450.

Wilhelmine H. Meeraus; Jennifer S. Armistead; Jorge R. Arias (2008). "Field comparison of novel and
gold standard traps for collecting Aedes albopictus in Northern Virginia". Journal of the American
Mosquito Control Association. 24 (2): 244–248. doi:10.2987/5676.1. PMID 18666532. S2CID 10287369.

Foley, K. (2007). "The BG-Sentinel Trap" (PDF). Presentation at the Annual Meeting of the Virginia
Mosquito Control Association. Archived from the original (PDF) on 13 October 2007.

Engelbrecht et al. (2009) Continuous trapping of adult Asian tiger mosquitoes (Aedes albopictus) with
BG-Sentinel traps reduced the human landing rate and density indices in an urban environment in
Cesena, Italy. Oral presentation at the 5th European Mosquito Control Association Workshop, Turin,
Italy, 12 March 2009. Session 10.5.

Kasai, S; Ng, LC; Lam-Phua, SG; Tang, CS; Itokawa, K; Komagata, O; Kobayashi, M; Tomita, T (2011). "First
detection of a putative knockdown resistance gene in major mosquito vector, Aedes albopictus".
Japanese Journal of Infectious Diseases. 64 (3): 217–21. doi:10.7883/yoken.64.217. ISSN 1344-6304.
PMID 21617306. S2CID 43513750.

Hoffmann, AA; Montgomery, BL; Popovici, J (25 August 2011). "Successful establishment of Wolbachia
in Aedes populations to suppress dengue transmission". Nature. 476 (7361): 454–457.
Bibcode:2011Natur.476..454H. doi:10.1038/nature10356. PMID 21866160. S2CID 4316652.
SOUTHERN HOUSE MOSQUITO

Culex quinquefasciatus (originally named Culex fatigans), commonly known as the southern house
mosquito, is a medium-sized mosquito found in tropical and subtropical regions of the world. It is a
vector of Wuchereria bancrofti, avian malaria, and arboviruses including St. Louis encephalitis virus,
Western equine encephalitis virus, Zika virus[1] and West Nile virus.[2][3] It is taxonomically regarded as
a member of the Culex pipiens species complex.[4][5] Its genome was sequenced in 2010, and was
shown to have 18,883 protein-coding genes.[6]

Culex quinquefasciatus

Scientific classification

Domain: Eukaryota

Kingdom: Animalia

Phylum: Arthropoda

Class: Insecta

Order: Diptera

Family: Culicidae

Genus: Culex

Species: C. quinquefasciatus

Binomial name

Culex quinquefasciatus

Say, 1823

Culex acer Walker, 1848

Culex aestuans Wiedemann, 1828

Culex aikenii Dyar & Knab, 1908

Culex albolineatus Giles, 1901

Culex anxifer Bigot, 1859

Culex aseyehae Dyar & Knab, 1915

Culex autumnalis Weyenbergh, 1882

Culex barbarus Dyar & Knab, 1906

Culex cartroni Ventrillon, 1905

Culex christophersii Theobald, 1907

Culex cingulatus Doleschall, 1856

Culex cubensis Bigot, 1857

Culex didieri Neveu-Lemaire, 1906

Culex fatigans Wiedemann, 1828

Culex fouchowensis Theobald, 1901

Culex hensemaeon Dyar, 1920

Culex luteoannulatus Theobald, 1901

Culex macleayi Skuse, 1889

Culex minor Theobald, 1908

Culex nigrirostris Enderlein, 1920

Culex pallidocephala Theobald, 1904

Culex penafieli Sanchez, 1885

Culex pungens Wiedemann, 1828

Culex pygmaeus Neveu-Lemaire, 1906

Culex quasilinealis Theobald, 1907

Culex quasipipiens Theobald, 1901

Culex raymondii Tamayo, 1907

Culex reesi Theobald, 1901

Culex revocator Dyar & Knab, 1909

Culex sericeus Theobald, 1901

Culex serotinus Philippi, 1865

Culex skusii Giles, 1900

Culex trillineatus Theobald, 1901

Culex zeltneri Neveu-Lemaire, 1906

Culicelsa fuscus Taylor, 1914

Etymology

American entomologist Thomas Say described Culex quinquefasciatus, which he collected along the
Mississippi River, in 1823. Originally written as "C. 5-fasciatus", the name refers to 5 (quinque) black,
broad, transverse bands ("fasciatus" or "fasciae") on the mosquito's dorsal abdomen. The name remains
despite later revelations of more than 5 fasciae, thanks to improved microscopy. Although
quinquefasciatus is the official scientific name, there are at least 5 synonymous names for this species.
[7]

Taxonomy

Cx. quinquefasciatus is a member of the Culex pipiens species complex.[4][5] Smith et al. 2004 develop
an assay specifically for this complex and use it to confirm that it does encompass this species.[4]

Description

The adult C. quinquefasciatus is a medium-sized mosquito and is brown in colour. The body is about 3.96
to 4.25 mm long. While the main body is brown, the proboscis, thorax, wings, and tarsi are darker than
the rest of the body. The head is light brown, with the lightest portion in the center. The antennae and
the proboscis are about the same length, but in some cases, the antennae are slightly shorter than the
proboscis. The flagellum has 13 segments that may have few or no scales. The scales of the thorax are
narrow and curved. The abdomen has pale, narrow, rounded bands on the basal side of each tergite.
Males can be differentiated from females in having large palps and feathery antennae.[8]

The larva has a short and stout head. The mouth brushes have long yellow filaments used for filtering
organic materials. The abdomen consists of eight segments, the siphon, and the saddle. Each segment
has a unique setae pattern. The siphon is on the dorsal side of the abdomen, and is four times longer
than its breadth. The siphon has multiple setae tufts. The saddle is barrel-shaped and located on the
ventral side of the abdomen, with four long anal papillae protruding from the posterior end.[2]

Lifecycle

Mature C. quinquefasciatus females fly at night to nutrient-rich standing water to lay eggs. They breed
profusely in dirty water collections, including stagnant drains, cesspools, septic tanks with leaks, burrow
pits, and almost all organic polluted water collections. A single female can lay up to five rafts of eggs in a
lifetime, with each raft containing 100 to 300 eggs.[9] The exact number varies depending on climatic
conditions.[2] The larvae feed on organic material in the water and require between five and eight days
to complete their development at 30 °C (86 °F). The larvae pass through four larval instars, and towards
the end of the fourth instar, they stop eating and undergo moulting to give rise to pupae. After 36 hours
at 27 °C (81 °F), adults emerge. The exact timing of development can vary depending on temperature. In
optimum temperature and humidity, the lifecycle will be completed in seven days, passing through the
egg, larval, pupal, and adult stages.[[]]

Both male and female adults take sugar meals from plants. After mating, the female seeks a blood meal
from a mammal or bird, as ingested blood is necessary for egg development. C. quinquefasciatus shows
a preference for the blood of birds, but will also commonly bite humans.[10]

Hosts

Known hosts include birds (Aves), cattle (Bos taurus), dogs (Canis familiaris), Equus including donkeys (E.
asinus), cats (Felis), mice (Mus musculus), house sparrows (Passer domesticus), rats (Rattus) and boars
(Sus scrofa).[11]

Distribution

"Quinx" are among the world's most abundant peridomestic mosquitoes, earning the nickname
"southern house mosquito". The species' place of origin is uncertain. It may have been native to the
lowlands of West Africa, or to Southeast Asia.[11] Cx. quinquefasciatus is now found throughout
subtropical and tropical areas worldwide, including the Americas, Australia and New Zealand,[12] except
for exceedingly dry or cold regions. Thomas Say described the species as "exceedingly numerous and
troublesome". It rests in trees and high places.[10]

As a vector

The southern house mosquito is a principal vector of numerous pathogens, transmitting the phlebovirus
Rift Valley fever virus, and the two flaviviruses St. Louis encephalitis virus and West Nile virus, plus
filarial worms and avian malarial parasites.[7]

It transmits zoonotic diseases that affect humans and wild and domestic animals, such as lymphatic
filariasis, avian malaria, St. Louis encephalitis, Western equine encephalitis, and West Nile fever, and
may be a vector of the Zika virus.[13] It causes infection through biting during blood meal. In the
southern U.S., it is the primary vector of St. Louis encephalitis virus. In India and Southeast Asia, it is the
primary vector of Wuchereria bancrofti, a nematode that causes lymphatic filariasis. It acts as an
intermediate host for the helminth parasite by harbouring the larval stages.[14] In Hawaii, it is the
principal vector of avian malaria (Plasmodium relictum), to which historic extinctions and significant
contemporary population declines in Hawaii's native honeycreeper species are attributed. It is the
definitive host for the malarial parasite as it harbours the sexual cycle.[15] In 2013 West Nile Virus
positive specimens were collected in Southern California.[16][failed verification]

References
 Nolen, Stephanie (July 21, 2016). ‘Very bad news for Brazil’: Zika virus found in second mosquito
species. The Globe and Mail.

Hill, Stephanie; Connelly, Roxanne (2009). "Features Creatures: Southern house mosquito". University
of Florida. Retrieved 19 March 2014.

Sandhu TS, Williams GA, Haynes BW, Dhillon MS. Evaluation of arboviral activity at Northwest Mosquito
and Vector Control District, Riverside County, California during 2008. Proc and Papers of the Mosq and
Vector Control Assoc of Calif, vol 77, 2009. p. 108-15.

Harbach, Ralph E. (2012). "Culex pipiens: Species Versus Species Complex – Taxonomic History and
Perspective". Journal of the American Mosquito Control Association. The American Mosquito Control
Association. 28 (4s): 10–23. doi:10.2987/8756-971x-28.4.10. ISSN 8756-971X. PMID 23401941. S2CID
31007129.

Turell, MJ (2012). "Members of the Culex pipiens complex as vectors of viruses". Journal of the
American Mosquito Control Association. 28 (4 Suppl): 123–6. doi:10.2987/8756-971X-28.4.123. PMID
23401952.

Arensburger, P.; Megy, K.; Waterhouse, R. M.; Abrudan, J.; Amedeo, P.; Antelo, B.; Bartholomay, L.;
Bidwell, S.; Caler, E.; Camara, F.; Campbell, C. L.; Campbell, K. S.; Casola, C.; Castro, M. T.;
Chandramouliswaran, I.; Chapman, S. B.; Christley, S.; Costas, J.; Eisenstadt, E.; Feschotte, C.; Fraser-
Liggett, C.; Guigo, R.; Haas, B.; Hammond, M.; Hansson, B. S.; Hemingway, J.; Hill, S. R.; Howarth, C.;
Ignell, R.; Kennedy, R. C.; Kodira, C. D.; Lobo, N. F.; Mao, C.; Mayhew, G.; Michel, K.; Mori, A.; Liu, N.;
Naveira, H.; Nene, V.; Nguyen, N.; Pearson, M. D.; Pritham, E. J.; Puiu, D.; Qi, Y.; Ranson, H.; Ribeiro, J. M.
C.; Roberston, H. M.; Severson, D. W.; Shumway, M.; Stanke, M.; Strausberg, R. L.; Sun, C.; Sutton, G.; Tu,
Z.; Tubio, J. M. C.; Unger, M. F.; Vanlandingham, D. L.; Vilella, A. J.; White, O.; White, J. R.; Wondji, C. S.;
Wortman, J.; Zdobnov, E. M.; Birren, B.; Christensen, B. M.; Collins, F. H.; Cornel, A.; Dimopoulos, G.;
Hannick, L. I.; Higgs, S.; Lanzaro, G. C.; Lawson, D.; Lee, N. H.; Muskavitch, M. A. T.; Raikhel, A. S.;
Atkinson, P. W. (2010). "Sequencing of Culex quinquefasciatus Establishes a Platform for Mosquito
Comparative Genomics". Science. 330 (6000): 86–88. doi:10.1126/science.1191864. PMC 3740384.
PMID 20929810.

Guagliard, Sarah Anne J.; Levine, Rebecca S. (August 2021). "Etymologia: Culex quinquefasciatus".
Emerg Infect Dis. 27 (8): 2041. doi:10.3201/eid2708.et2708. PMC 8314802. Retrieved September 2,
2021. This article incorporates text from this source, which is in the public domain.

"Brown House Mosquito (Culex quinquefasciatus)". OzAnimals.com. Retrieved 19 March 2014.

"Culex Mosquito Life Cycle | CDC". Centers for Disease Control and Prevention. 2022-07-12. Retrieved
2023-07-15.

Prada, Paulo (2016). "Research indicates another common mosquito may be able to carry Zika".
Reuters.
"Culex quinquefasciatus (southern house mosquito)". Invasive Species Compendium. CAB International.
Retrieved 6 January 2022.

"Southern house mosquito – Culex quinquefasciatus Say". Featured Creatures. University of Florida.
Retrieved 6 January 2022.

Kathy Keatley Garvey. 2016. "Are Culex Mosquitoes Potential Vectors of the Zika Virus?", Bug Squad,
Happenings in the Insect World, March 2, 2016.

Albuquerque, Cleide MR; Cavalcanti, Vânia MS; Melo, Maria Alice V; Verçosa, Paulo; Regis, Lêda N;
Hurd, Hilary (1999). "Bloodmeal microfilariae density and the uptake and establishment of Wuchereria
bancrofti infections in Culex quinquefasciatus and Aedes aegypti". Memórias do Instituto Oswaldo Cruz.
94 (5): 591–596. doi:10.1590/S0074-02761999000500005. PMID 10464399.

Farajollahi, Ary; Fonseca, Dina M.; Kramer, Laura D.; Marm Kilpatrick, A. (October 2011). ""Bird biting"
mosquitoes and human disease: A review of the role of Culex pipiens complex mosquitoes in
epidemiology". Infection, Genetics and Evolution. 11 (7): 1577–1585. doi:10.1016/j.meegid.2011.08.013.
PMC 3190018. PMID 21875691.

Sandhu, Tejbir S.; Williams, Gregory W.; Haynes, Bryan W.; Dhillon, Major S. (2013). "Population
dynamics of blood-fed female mosquitoes and comparative efficacy of resting boxes in collecting them
from the northwestern part of riverside county, California". Journal of Global Infectious Diseases. 5 (1):
15–18. doi:10.4103/0974-777X.107168. ISSN 0974-777X. PMC 3628227. PMID 23599612.

Wyeomyia smithii, the pitcher plant mosquito, is an inquiline mosquito that completes its pre-adult life
cycle in the phytotelma of—that is, the water contained by—the purple pitcher plant, Sarracenia
purpurea. In this microcommunity of bacteria, rotifers, protozoa, and midges, W. smithii is the top-level
predator; its presence determines the bacterial species diversity within the pitcher.[1]

Wyeomyia smithii

Female

Wyeomyia smithii larva magnified 40×

Scientific classification

Domain: Eukaryota

Kingdom: Animalia

Phylum: Arthropoda

Class: Insecta
Order: Diptera

Family: Culicidae

Subfamily: Culicinae

Tribe: Sabethini

Genus: Wyeomyia

Species: W. smithii

Binomial name

Wyeomyia smithii

Coquillett, 1901

W. smithii is not a pest mosquito in general. The northern US population does not consume blood at all,
while the southern US populations only consume blood after laying an initial egg batch;[2][3] even then
they appear disinterested in feeding. In fact, it is the only known mosquito to have both obligatory biting
and non-biting populations in the same species.[4]

Life cycle

Description of the life cycle

The life cycle of Wyeomyia smithii begins in either late spring or early fall when the adult female lays her
eggs in the phytolema of a purple pitcher plant. The eggs then gestate in the pitcher plant anywhere
from 1–8 days before they hatch as larvae. The larvae remain in the phytolema feeding on bacteria,
micro-animals, and decaying insects. The mosquito larvae will live in the pitcher plant until it goes
through its fifth instar about 20 to 22 days after hatching. They then emerge as adult mosquitoes ready
to mate. Wyeomyia smithii females will produce a clutch of fertilized eggs generally within two days of
sexual maturity. Some populations in the southern United States have also been known to drink blood
after producing their first clutch to help produce a second clutch; there are no reports of northern
populations displaying this behavior, however.[5]

Photoperiodism

Wyeomyia smithii is a model organism for the study of photoperiodism, the biotic process of controlling
seasonal life history events by measuring day length as a reliable predictor of the seasons. W. smithii
enters a state of developmental arrest, larval diapause, that is initiated and maintained by short day
lengths and averted or terminated by long day lengths.[6] There is speculation that hypothetical global
warming and the shortening of winters, W. smithii has been observed to now require shorter days
before going dormant, this led to the theory that this is an example of microevolutionary selection;
mosquitoes that waited longer to go dormant and which had a greater fitness have been favored.[7]
Photoperiodism in Wyeomyia smithii is believed in nature to examine the startling finding that
proposed, but not proven, recent climate change can force genetic change in plants and animals.
Wyeomyia smithii lives through the winter as diapausing larvae in the leaves of the pitcher plant. The
pitcher-plant mosquito enters a hibernal diapause as larvae. Which means, short days initiate and
maintain diapause, and long days promote continuous development in diapausing larvae. The day length
promotes 50% development and 50% diapause (the critical photoperiod) as the same for the initiation
and termination of not developing in unchilled larvae. For the initiation of not developing, insects pass
through a "sensitive period" during which day length is interpreted as long or short, which results in a
diapause/no-diapause response. Wyeomyia smithii is photoperiodic while in diapause. The sensitive
period is indefinitely long, and the effect of manipulating different light. Dark cycles can be assessed
over weeks or months, instead of a few days. Wyeomyia smithii oviposits into and completes their entire
preadult development only within the water-filled leaves of the carnivorous pitcher plant Sarracenia
purpurea. Throughout this range, W. smithii occupies a uniform microhabitat whose community
composition remains highly consistent. The photoperiodic response has been exposed to various
seasonal changes, both in situ and during postglacial dispersal.[8]

Co-evolution of mosquito and plant

Plant adaptations

The purple pitcher plant (Sarracenia purpurea), which houses W. smithii, is known to be the most
habitable pitcher plant for many different inquilines. At least 165 various species of insects, protozoa,
algae, and other organisms can live within the purple pitcher plant. The plant has adapted to have fluid
with a higher pH than other, more acidic, carnivorous pitcher plants. There is little fluid secreted by this
plant in comparison to the rainwater it collects in its pitcher, which dilutes the solution and creates a
more habitable environment for outside organisms. The plant's adaptation allowing it to host diverse life
is an advantage to the mosquito because its larvae prey upon those organisms. The mosquitos also prey
on the food that the pitcher plant catches, giving them a steady food source.[9]

Mosquito evolution/adaptations

The pitcher plant mosquito, Wyeomyia smithii, has been featured in scientific studies because of its co-
evolution with the purple pitcher plant. The relationship between the two species highlights a genetic
response to climate change, which details how a population can evolve at a quicker pace in order to
make accommodations due to the changes in climate. The water-filled pitcher plant proved to be a
suitable home for the mosquito in its habitat range. It is because both species can endure a temperate
climate. This idea makes the pitcher plant the next option for this mosquito species to stay in its current
domain.[10]

The relationship is beneficial for both species and allows them to continue to evolve together. The
purple pitcher plant uses the mosquito for nutrients once the mosquitoes die off. And, these mosquitoes
are no different than any other subspecies, in terms of attraction, they are drawn to water. The pitcher
plant is filled with water and this is a type of environment that is used as a location for female
mosquitoes to lay their eggs. Most pitcher plant mosquitoes tend to frequent a pitcher plant younger in
age. Not only does the plant provide a steady source of food, it is a protective place for larvae to
develop.[11]

References

Celeste N. Peterson; Stephanie Day; Benjamin E. Wolfe; Aaron M. Ellison; Roberto Kolter & Anne Pringle
(2008). "A keystone predator controls bacterial diversity in the pitcher-plant (Sarracenia purpurea)
microecosystem" (PDF). Environmental Microbiology. 10 (9): 2257–2266. doi:10.1111/j.1462-
2920.2008.01648.x. PMID 18479443.

Bradshaw, William E. (1980). "Blood-feeding and capacity for increase in the pitcher-plant mosquito,
Wyeomyia smithii". Environmental Entomology. 9 (1): 86–89. doi:10.1093/ee/9.1.86.

D. Allen (2015). Carbon dioxide sensitivity in two disjunct populations of the pitcher-plant mosquito,
Wyeomyia smithii. University Honors Program Theses. Vol. 126. Georgia Southern University.

Armbruster, Peter A. (30 January 2018). "Molecular pathways to nonbiting mosquitoes". Proceedings of
the National Academy of Sciences. 115 (5): 836–838. doi:10.1073/pnas.1721209115. PMC 5798389.
PMID 29330331.

Donahue, Luke (2012). "Wyeomyia smithii". Animal Diversity Web. Retrieved 12 December 2017.

William E. Bradshaw & L. Philip Lounibos (1977). "Evolution of dormancy and its photoperiodic control
in pitcher-plant mosquitoes". Evolution. 31 (3): 546–567. doi:10.2307/2407521. JSTOR 2407521. PMID
28563474.

"Evolution 101: Examples of Microevolution". University of California Berkeley. Retrieved 2014-10-15.

Bradshaw, W.E.; Holzapfel, C. M. (2017). "Chapter 2: Natural Variation and Genetics of Photoperiodism
in Wyeomyia smithii.". Advances in Genetics. Vol. 99. Academic Press. pp. 39–71.

Adlassnig, W.; Peroutka, M. & Lendl, T. (15 December 2010). "Traps of carnivorous pitcher plants as a
habitat: composition of the fluid, biodiversity and mutualistic activities". Annals of Botany. 107 (2): 181–
194. doi:10.1093/aob/mcq238. PMC 3025736. PMID 21159782.

"Pitcher plant-dwelling mosquito shows effects of Earth's rapidly changing climate". Genetic Structure
of First Animal to Show Evolutionary Response to Climate Change Determined. National Science
Foundation. Retrieved 12 December 2019.

Nastase, Anthony; De La Rosa, Carlos & Newell, Sandra (1995). "Abundance of pitcher-plant
mosquitoes, Wyeomyia smithii (Coq.) (Diptera: Culicidae) and midges, Metriocnemus knabi Coq.
(Diptera: Chironomidae), in relation to pitcher characteristics of Sarracenia purpurea L.". The American
Midland Naturalist. 133 (1): 44–51. doi:10.2307/2426346. JSTOR 2426346.
Aedes sollicitans, the eastern saltmarsh mosquito (also known as Ochlerotatus sollicitans), is a species of
mosquito native to the eastern seaboard of the United States and Canada as well as the entire Gulf coast
and is also present in the Bahamas and Greater Antilles. While primarily found in coastal areas within a
few miles of the coast, it is occasionally found inland in areas with saline pools, the species was reported
as far west as Arizona. The species is a prime vector for Eastern equine encephalitis, Venezuelan equine
encephalitis and dog heartworm.

Aedes sollicitans

Scientific classification

Domain: Eukaryota

Kingdom: Animalia

Phylum: Arthropoda

Class: Insecta

Order: Diptera

Family: Culicidae

Genus: Aedes

Species: A. sollicitans

Binomial name

Aedes sollicitans

(Walker, 1856)

Aedes sollicitans has a conspicuous band of white scales around the central area of the proboscis and
the anterior portion of the hind tarsomeres upon which there is also band a band of yellow scales in the
middle. The abdomen has white basal bands and is divided by a medial longitudinal stripe. The thorax is
white on the sides and the top is brown, yellow, golden and white.[1]

Similar Species

A. sollicitans resembles Aedes taeniorhynchus but the two species can be distinguished at the larval and
adult stages. Larval A. sollicitans have longer breathing tubes, have scale patches with pointed tips, and
larger spines that line the edges of each scale patch. Adult A. sollicitans are golden brown while adult
Aedes taeniorhychus are black and smaller in size.[2]

Habitat
A. sollicitans tends to stay within 5 miles of the coast on average all the range can be greater dependent
upon a number of factors such as wind speed and duration.

Food Resources

It tends to feed most actively at twilight but is an opportunistic feeder which will feed a host species that
enters its area in daytime. The female requires one blood meal for each egg batch with the primary host
species being mammals, and birds as a secondary host.

Oviposition

The female Aedes sollicitans lays her eggs on the dried out substrate of salt pannes, depressions within
salt marshes which dry out between periods of very high tide (spring tide). The eggs hatch upon the
panne filling at the next spring tide in 4–5 days with optimal conditions.

In the south the peak amount of adults occurs in the spring and fall, and in the northern portion of its
range peak adult population occurs in the summer. The last batches of eggs laid in the fall remain in
diapause until the spring.[3]

References

"University of Florida Aedes sollicitans". Archived from the original on 2010-06-12. Retrieved 2010-07-
18.

Komp, W. H. W. (1923). "Guide to Mosquito Identification for Field Workers Engaged in Malaria Control
in the United States". Public Health Reports. 38 (20): 1061–1080. doi:10.2307/4576745. ISSN 0094-6214.
JSTOR 4576745.

O'Meara, G. F. 1992. The eastern saltmarsh mosquito Aedes sollicitans. Wing Beats, Vol. 3(4):5.

1. YELLOW FEVER MOSQUITO

2. ASIAN TIGER MOSQUITO

3. ANOPHELES MOSQUITO

4. SOUTHERN HOUSE MOSQUITO

5. PITCHER PLANT MOSQUITO

6. EASTERN SALTMARSH MOSQUITO