Coccinellidae (/ˌkɒksɪˈnɛlɪdaɪ/)[3] is a widespread family of small beetles ranging from 0.8 to 18 mm (0.0315 to 0.708 inches).[4]
They are commonly yellow, orange, or red with small black spots on their wing covers, with black legs, heads and antennae. However such colour patterns vary greatly. For example, a minority of species, such as Vibidia duodecimguttata,
a twelve-spotted species, have whitish spots on a brown background.
Coccinellids are found worldwide, with over 6,000 species described.[5][6]
Coccinellids are known as ladybugs in North America, and ladybirds in other areas. Entomologists widely prefer the names ladybird beetles or lady beetles as these insects are not classified as true bugs.[7]
The majority of coccinellid species are generally considered useful insects, because many species prey on herbivorous homopterans such as aphids or scale insects,
which are agricultural pests. Many coccinellids lay their eggs directly
in aphid and scale insect colonies in order to ensure their larvae have
an immediate food source.[8] However, some species do have unwelcome effects; among these, the most prominent are of the subfamilyEpilachninae,
which are herbivorous themselves. Usually, epilachnines are only mild
agricultural pests, eating the leaves of grain, potatoes, beans, and
various other crops, but their numbers can increase explosively in years
when their natural enemies, such as parasitoid
wasps that attack their eggs, are few. In such situations, they can do
major crop damage. They occur in practically all the major
crop-producing regions of temperate and tropical countries.
The name coccinellids is derived from the Latin word coccineus meaning "scarlet".[9] The name "ladybird" originated in Britain where the insects became known as "Our Lady's bird" or the Lady beetle.[10][11]Mary
(Our Lady) was often depicted wearing a red cloak in early paintings,
and the spots of the seven-spot ladybird (the most common in Europe)
were said to symbolise her seven joys and seven sorrows.[10]
In the United States, the name was adapted to "ladybug". Common names
in other European languages have the same association, for example, the
German name Marienkäfer translates to Marybeetle.[12]
Physical appearance
Most
coccinellids have oval, dome-shaped bodies with six short legs.
Depending on the species, they can have spots, stripes, or no markings
at all. Seven-spotted coccinellids are red or orange with three spots on
each side and one in the middle; they have a black head with white
patches on each side.
As well as the usual yellow and deep red colourings, many coccinellid
species are mostly, or entirely, black, dark grey, gray, or brown, and
may be difficult for an entomologist/nonentomologists to recognise as
coccinellids at all. Conversely, non-entomologists might easily mistake
many other small beetles for coccinellids. For example, the tortoise beetles,
like the ladybird beetles, look similar because they are shaped so that
they can cling to a flat surface so closely that ants and many other
enemies cannot grip them.
Non-entomologists are prone to misidentify a wide variety of beetle
species in other families as "ladybirds", i.e. coccinellids. Beetles are
particularly prone to such misidentification if they are spotted in
red, orange or yellow and black. Examples include the much larger scarabaeidgrapevine beetles and spotted species of the Chrysomelidae, Melyridae
and others. Conversely, laymen may fail to identify unmarked species of
Coccinellidae as "ladybirds". Other beetles that have a defensive
hemispherical shape, like that of the Coccinellidae (for example the Cassidinae), also are often taken for ladybirds.
A common myth, totally unfounded, is that the number of spots on the insect's back indicates its age.[13][14]
In fact, the underlying pattern and colouration are determined by the
species and genetics of the beetle, and develop as the insect matures.
In some species its appearance is fixed by the time it emerges from its pupa,
though in most it may take some days for the colour of the adult beetle
to mature and stabilise. Generally, the mature colour tends to be
fuller and darker than the colour of the callow.
Coccinellids are best known as predators of Sternorrhyncha such as aphids and scale insects, but the range of prey species that various Coccinellidae may attack is much wider. A genus of small black ladybirds, Stethorus, presents one example of predation on non-Sternorrhyncha; they specialise in mites as prey, notably Tetranychus spider mites. Stethorus species accordingly are important in certain examples of biological control.[15]
Various larger species of Coccinellidae attack caterpillars and other beetle larvae.[15] Several genera feed on various insects or their eggs; for example, Coleomegilla species are significant predators of the eggs and larvae of moths such as species of Spodoptera and the Plutellidae.[16]Larvae and eggs of ladybirds,
either their own or of other species, can also be important food
resources when alternative prey are scarce. As a family, the
Coccinellidae used to be regarded as purely carnivorous,[17]
but they are now known to be far more omnivorous than previously
thought, both as a family and in individual species; examination of gut
contents of apparently specialist predators commonly yield residues of
pollen and other plant materials. Besides the prey they favour, most
predatory coccinellids include other items in their diets, including
honeydew, pollen, plant sap, nectar, and various fungi. The significance
of such nonprey items in their diets is still under investigation and
discussion.[18]
Apart from the generalist aphid and scale predators and incidental
substances of botanical origin, many Coccinellidae do favour or even
specialise in certain prey types. This makes some of them particularly
valuable as agents in biological control programmes. Determination of
specialisation need not be a trivial matter, though; for example the
larva of the Vedalia ladybird Rodolia cardinalis is a specialist predator on a few species of Monophlebidae, in particular Icerya purchasi, which is the most notorious of the cottony cushion scale species. However, the adult R. cardinalis can subsist for some months on a wider range of insects plus some nectar.[19]
Certain species of coccinellids are thought to lay extra infertile
eggs with the fertile eggs, apparently to provide a backup food source
for the larvae when they hatch. The ratio of infertile to fertile eggs
increases with scarcity of food at the time of egg laying. Such a
strategy amounts to the production of trophic eggs.[20]
Some species in the subfamily Epilachninae are herbivores, and can be very destructive agricultural pests (e.g., the Mexican bean beetle). Again, in the subfamily Coccinellinae, members of the tribe Halyziini and the genus Tythaspis are mycophagous.
While predatory species are often used as biological control agents, introduced species of coccinellids are not necessarily benign. Species such as Harmonia axyridis or Coccinella septempunctata in North America outcompete and displace native coccinellids and become pests themselves.[21]
The main predators of coccinellids are usually birds, but they are
also the prey of frogs, wasps, spiders, and dragonflies. The bright
colours of many coccinellids discourage some potential predators from
making a meal of them. This phenomenon, called aposematism, works because predators learn by experience to associate certain prey phenotypes with a bad taste. A further defence, known as "reflex bleeding",
exists in which an alkaloid toxin is exuded through the joints of the
exoskeleton, triggered by mechanical stimulation (such as by predator
attack) in both larval and adult beetles, deterring feeding.
Coccinellids in temperate regions enter diapause during the winter, so they often are among the first insects to appear in the spring. Some species (e.g., Hippodamia convergens) gather into groups and move to higher elevations, such as a mountain, to enter diapause.
Most coccinellids overwinter
as adults, aggregating on the south sides of large objects such as
trees or houses during the winter months, dispersing in response to
increasing day length in the spring.[22]
Predatory coccinellids are usually found on plants which harbour
their prey. They lay their eggs near their prey, to increase the
likelihood the larvae will find the prey easily. In Harmonia axyridis,
eggs hatch in three to four days from clutches numbering from a few to
several dozen. Depending on resource availability, the larvae pass
through four instars over 10–14 days, after which pupation occurs. After a teneral
period of several days, the adults become reproductively active and are
able to reproduce again, although they may become reproductively
quiescent if eclosing late in the season. Total life span is one to two years on average.
Infestations and impacts
Coccinellids covering a branch
In the United States, coccinellids usually begin to appear indoors in
the autumn when they leave their summer feeding sites in fields,
forests, and yards and search out places to spend the winter. Typically,
when temperatures warm to the mid-60s F (around 18 °C) in the late
afternoon, following a period of cooler weather, they will swarm onto or
into buildings illuminated by the sun. Swarms of coccinellids fly to
buildings in September through November depending on location and
weather conditions. Homes or other buildings near fields or woods are
particularly prone to infestation.[23]
After an abnormally long period of hot, dry weather in the summer of 1976 in the UK,
a marked increase in the aphid population was followed by a "plague" of
ladybirds, with many reports of people being bitten as the supply of
aphids dwindled.[24][25]
The presence of coccinellids in grape harvests can cause ladybird taint in wines produced from the grapes.[26]
As an alien species
Harmonia axyridis
(the harlequin ladybird) is an example of how an animal might be partly
welcome and partly harmful. It was introduced into North America from
Asia in 1916 to control aphids, but is now the most common species,
outcompeting many of the native species.[27] It has since spread to much of western Europe, reaching the UK in 2004.[27][28]
It has become something of a domestic and agricultural pest in some
regions, and gives cause for ecological concern. It similarly has turned
up in parts of Africa, where it has proved variously unwelcome, perhaps
most prominently in vine-related crops.
The atlas Ladybirds (Coccinellidae) of Britain and Ireland published in 2011[29]
showed a decline of more than 20% in native species due to
environmental changes and competition from foreign invaders. The
distribution maps, compiled over a 20-year period with help from
thousands of volunteers, showed a decline in the numbers of the common
10-spot and 14-spot ladybirds and a number of other species, including
the 11-spot, 22-spot, cream-spot, water and hieroglyphic ladybirds, Coccidula rufa, Rhyzobius litura and Nephus redtenbacheri. Conversely, increases were seen in the numbers of harlequin, orange, pine, and 24-spot ladybirds, as well as Rhyzobius chrysomeloides.
The kidney spot ladybird was recorded in Scotland for the first time in
recent years, and the 13-spot was found to have recolonised Cornwall, Devon, and the New Forest.
The most commonly recorded species was the 7-spot, closely followed by
the Asian harlequin — an invader that arrived from continental Europe in
2003 after being introduced to control pests. An 'explosion' in the
number of orange ladybirds, which feed on mildew, is thought to have
been due to the warmer, damper conditions that now prevail in parts of
England.[30]
Midfielder Hakan Çalhanoğlu says Turkey have been
"waiting impatiently" for their opening game against Croatia – and
believes "showing our talent" will give them a good shot at glory.
Hakan Çalhanoğlu on ... a long wait We've
been waiting impatiently for this day to come. This will be a big day,
for all our country. We are ready for the match and we've prepared well.
We've done what was necessary; our coach Fatih Terim has prepared us
well and we feel good. Now let's hope we get a good result. Hakan Çalhanoğlu on ... the preparation period We've
gone 14 or 15 matches with only one defeat. As a team, we've developed
very well. I am hoping that, against good opposition, we can maintain
the same team spirit on the field. We shouldn't forget that Croatia are
tough opponents. But if we play in our own style, if we show our talent,
we have a good shot.
Hakan Çalhanoğlu on ... Turkey's match winners There
are lots of players with that capability. Anyone can do anything on the
pitch, including me. I hope we're lucky enough for that opportunity to
arise. We'd like to keep going beyond the group and we are hoping to get
three points. Hakan Çalhanoğlu on ... his free-kick technique Of
course there have been players I’ve looked up to in the past. I love
David Beckham and Juninho Pernambucano. But now I have found my own
style. Hakan Çalhanoğlu on ... the tournament prospects All
the UEFA EURO 2016 groups are difficult. Spain might look like the
favourites in ours but for me anything can happen in this group. All the
members of our team take great pleasure in playing for Turkey. We have
young players and experienced ones. We have a good atmosphere within the
squad.
Overview. Turkic
languages originated in the steppes of Central Asia from where they
were carried west by nomadic peoples to reach Turkey and the Balkans,
and north and east advancing into European Russia and Siberia.
Comprising some thirty members and spoken by more than 160 million
people, they are of the agglutinating type and typologically quite
similar.
Distribution. Turkic
languages are distributed over a vast territory ranging from eastern
Europe to east Siberia and China. Their core area is in Central Asia
(Kazakhstan, Kyrgyzstan, Turkmenistan, Uzbekistan, and Xinjiang in
China) from where they spread west to northern Iran and the South
Caucasus, Turkey and part of the Balkans, and to the north into European
and Asiatic Russia straddling the Volga, Ob and Yenisei rivers reaching
northeast Siberia and the Arctic Ocean.
Map of Turkic languages distribution (click to enlarge it)
External Classification. The external classification of Turkic languages is disputed. Many scholars consider them one of the three subfamilies of the Altaic family (the other two are Tungusic and Mongolic).
However, the parallelisms between Turkic, Mongolic and Tungusic are too
few, according to others, to support the unity of Altaic and,
therefore, they may be independent families.
Internal Classification. In contrast, there is agreement regarding the internal classification of Turkic. It is divided into six branches:
1.The SouthwesternorOghuz branch contains a western group comprising Turkish, Gagauz, and Azerbaijanian and an eastern group comprising Turkmenand Khorasani Turkic. The Salar
language of China seems to be Oghuz in origin but during its eastward
migration it acquired features from northwestern and southeastern
languages.
2.The NorthwesternorKipchakbranch has a western group comprising Kumyk,Karachay-Balkar, Crimean Tatar, and Karaim, a northern group comprising Tatar and Bashkir, and a southern group comprising Kazakh, Karakalpak, Kipchak Uzbek (considered a dialect of Kazakh), Nogai, and Kirghiz (Kyrgyz).
3.The SoutheasternorUyghur-Karluk branch has a western group represented by Uzbek and an eastern group represented by Uyghur.
4.The Northeasternor Siberian branch has a southern heterogeneous group comprising Tuvan (Tuva), Karagas (Tofa), Khakas, Shor, Chulym, Altai, Yellow Uyghur (West Yugur) and a northern group comprising Yakut(Sakha) and Dolgan.
5.Chuvash situated in the northwestern area (in the Volga region).
6.Khalaj situated in the southwestern area (northwest Iran).
Speakers. The
total number of Turkic speakers is close to 164 million. The following
is a complete list of living Turkic languages with their number of
speakers and distribution:
Russia (border with China, Kazakhstan and Mongolia)
Russia (close to border with Mongolia, north of Altai)
Iran (Markazi Province, north of Arak)
China (northwest Gansu province)
Russia (north of Altai and west of Khakas)
Russia (Sakha Republic in north Siberia)
Lithuania, Ukraine (Halych)
Russia (southern Siberia, Chulym River basin)
Russia (southern Siberia)
Note: It is not possible to
estimate accurately the number of Khorasani Turkic speakers. There may
be between 400,000 and one million or more.
Oldest Documents.They
appear in the territory of the second Türk empire, written in Old
Turkic with the native 'runic' script. The earliest are engraved in an
ensemble of four stone stelae erected, between 720 and 735, in the
Orkhon valley (northern Mongolia) in honor of several rulers (the king
Kaghan Bilge, his brother the prince Kol Tegin and their commander
Tonyuquq). They tell about the origins of the Turks, their subjugation
by the Chinese and subsequent liberation by Bilge.
SHARED FEATURES
✦Phonology
-Syllable structure.
Most syllables have a (C)V(C) structure i.e. they contain a vowel that
may be preceded by an initial consonant and/or followed by a final
consonant. Initial consonant clusters are avoided as well as vowel
hiatus (two adjacent vowels in different syllables).
-Vowels.
Many Turkic languages (Turkish among them) have a completely
symmetrical vowel system regarding height (4 high and 4 low vowels),
frontness (4 front and 4 back), and roundness (4 unrounded and 4
rounded):
The
symbols are those current in writing, when they differ from those of
the International Phonetic Alphabet the latter are indicated between
brackets.
Some
languages (Turkmen, Khalaj, Yakut) exhibit a phonemic contrast between
short and long vowels; those in the Volga region (Chuvash, Bashkir,
Tatar) have reduced vowels.
-Consonants. Most
Turkic languages have a contrast between voiceless and voiced stops and
fricatives, though a few others, like Chuvash, have only voiceless
ones. They are usually articulated at four or five places: labial,
dental-alveolar, palatal, velar and uvular-glottal. Gagauz and Karaim
have palatalized consonants acquired by Slavic influence.
-Sound harmony. It
is widespread in Turkic. The most general type is intrasyllabic
affecting the vowel and consonant(s) of a given syllable. The whole
syllable is classified as front or back; in front syllables only front
vowels and front consonants are allowed; in back syllables the opposite
is true.
There
is also an intersyllabic type of harmony in which words tend to consist
of syllables produced with either a back or a front tongue position. In
some languages, like Yakut and Kirghiz, harmony may be extended to
vowel roundness.
-Accent. Turkic
languages have a pitch accent (increase of the tone height), which
tends to fall on the last syllable. Some suffixes are accentable while
others are not. There is also a stress accent which tends to fall on the
first syllable.
✦Morphology
It
is agglutinative and suffixing. Stems are expanded by adding
unchangeable and clear-cut monosyllabic suffixes, of which many serve to
express grammatical notions. Morphology is regular and predictable and
there is little or no fusion between morphemes. There are virtually no
prefixes. The order of suffixes is rigidly established, derivational
suffixes preceding inflectional ones. Each added suffix tends to modify
the whole preceding stem.
Nominal
-Nouns are marked for number, possession and case (in that order). There are no grammatical genders.
-The plural number is usually marked with the suffix lar/ler.
-Possession
may be indicated by possessive suffixes or by independent possessive
pronouns (genitive of personal and demonstrative pronouns). The latter
are usually employed for emphasis. Demonstrative pronouns distinguish
three deictic degrees (proximal, intermediate, and distal).
-There
are six general cases: nominative (unmarked), accusative, genitive,
dative, locative and ablative. Not all Turkic languages have this whole
six-case system. For example, Yakut lacks genitive and locative but has a
partitive case; in Chuvash the dative and accusative have merged; in
Kumyk, Karachai-Balkar and in some Uzbek dialects the accusative and the
genitive have merged. Other, subsidiary, cases are: equative ('like'),
terminative ('until'), comitative (company), instrumental ('by means
of'), directive ('towards'), etc.
-Adjectives are morphologically similar to nouns. Intensive adjectives may be formed by reduplication of the first syllable.
-Superlatives
are formed by adding particles meaning ‘most' or 'best'; comparatives
are expressed by particles meaning ‘more' or by putting the standard of
comparison in the ablative case.
-There is an indefinite article (identical to number one) but no definite articles. Demonstratives can, sometimes, act as such.
Verbal
-To
the verb root derivational suffixes can be added to build up the verb
stem conferring causative, reflexive, negative, passive meanings, etc.
To the stem, tense/aspect suffixes and personal endings usually follow.
The first and second persons are marked by personal suffixes, the third
person is usually unmarked.
-Non-finite forms include infinitives, participles, and verbal adverbs (converbs).
✦Syntax
-Word
order is Subject-Object-Verb in Turkic languages. They are head final,
modifiers preceding their head. In the nominal phrase the order is:
demonstrative pronoun-cardinal number-adjective-head noun. Attributes do
not agree in number or case with their heads.
-Postpositions,
corresponding to English prepositions, are placed after the words they
modify. Conjunctions are used sparingly and most of the existing ones
have been borrowed from non-Turkic languages. In genitive constructions
the possessor, which is the first element, carries a genitive suffix
while the possessed carries a possessive suffix.
Scripts
Turkic languages have
been written with a great variety of scripts in the past (Runic, Uyghur,
Brāhmī, Manichean, etc). The Arabic script was used for all Turkic
languages after the adoption of Islam and until the beginning of the
20th century when it was replaced in many countries by the Latin or
Cyrillic scripts. It is still used in China and Iran.
In the former
territory of the Soviet Union, Cyrillic-based scripts were imposed for
all Turkic languages until its dissolution when some of them started a
process of change into the Latin alphabet. The latter was introduced in
Turkey in 1923.
