Belidae
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Belidae is a
family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...
of weevils, called belids or primitive weevils because they have straight antennae, unlike the "true weevils" or
Curculionidae The Curculionidae are a family of weevils, commonly called snout beetles or true weevils. They are one of the largest animal families, with 6,800 genera and 83,000 species described worldwide. They are the sister group to the family Brentidae. T ...
which have geniculate (elbowed) antennae. They are sometimes known as "
cycad Cycads are seed plants that typically have a stout and woody (ligneous) trunk (botany), trunk with a crown (botany), crown of large, hard, stiff, evergreen and (usually) pinnate leaves. The species are dioecious, that is, individual plants o ...
weevils", but this properly refers to a few species from the
genera Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomenclat ...
'' Parallocorynus'' and ''
Rhopalotria ''Rhopalotria'' is a genus of cycad weevils in the beetle family Belidae. There are about six described species in ''Rhopalotria''. Species These six species belong to the genus ''Rhopalotria'': * ''Rhopalotria dimidiata'' Chevrolat, 1878 * '' ...
''.


Distribution

The Belidae today have an essentially
Gondwana Gondwana () was a large landmass, often referred to as a supercontinent, that formed during the late Neoproterozoic (about 550 million years ago) and began to break up during the Jurassic period (about 180 million years ago). The final stages ...
n distribution, occurring only in the Australia–
New Guinea New Guinea (; Hiri Motu Hiri Motu, also known as Police Motu, Pidgin Motu, or just Hiri, is a language of Papua New Guinea, which is spoken in surrounding areas of Port Moresby (Capital of Papua New Guinea). It is a simplified version of ...
–New Zealand region up to Southeast Asia, South and Central America (barely reaching North America), some Pacific islands (notably the
Hawaiian Islands The Hawaiian Islands ( haw, Nā Mokupuni o Hawai‘i) are an archipelago of eight major islands, several atolls, and numerous smaller islets in the North Pacific Ocean, extending some from the island of Hawaii in the south to northernmost Kur ...
) and a few places in Africa. Many lineages of belids are notable for their highly
relict A relict is a surviving remnant of a natural phenomenon. Biology A relict (or relic) is an organism that at an earlier time was abundant in a large area but now occurs at only one or a few small areas. Geology and geomorphology In geology, a r ...
ual distribution; for example the
Aglycyderini Aglycyderini is a tribe of belids, primitive weevils of the family Belidae. Like in other belids, their antennae are straight, not elbowed as in the true weevils ( Curculionidae). They occur only on the Pacific Islands and in the Macaronesian ...
are found in two areas on opposite sides of the Earth, with no such beetles known from anywhere in between. Belids were more widespread during the
Late Jurassic The Late Jurassic is the third epoch of the Jurassic Period, and it spans the geologic time from 163.5 ± 1.0 to 145.0 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic strata.Owen 1987. In European lithostratigraphy, the name ...
to
Early Cretaceous The Early Cretaceous ( geochronological name) or the Lower Cretaceous (chronostratigraphic name), is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 145  Ma to 100.5 Ma. Geology Pro ...
, about , when they were found at least in Central Asia, Spain and Brazil.
Paleogene The Paleogene ( ; British English, also spelled Palaeogene or Palæogene; informally Lower Tertiary or Early Tertiary) is a geologic period, geologic period and system that spans 43 million years from the end of the Cretaceous Period million yea ...
species belonging to still-living
tribe The term tribe is used in many different contexts to refer to a category of human social group. The predominant worldwide usage of the term in English language, English is in the discipline of anthropology. This definition is contested, in p ...
s are known from Europe and North America.


Description

Members of the subfamily Belinae are typically elongated and cylindrical. Some belines resemble other weevils, like the genus ''
Lixus Lixus may refer to: * ''lixus'', the Latin word for "boiled" * Lixus (ancient city) in Morocco * ''Lixus (beetle)'', a genus of true weevils * Lixus, one of the sons of Aegyptus and Caliadne Caliadne (; Ancient Greek: Καλιάδνης ) or Cali ...
'' of the true weevil family (
Curculionidae The Curculionidae are a family of weevils, commonly called snout beetles or true weevils. They are one of the largest animal families, with 6,800 genera and 83,000 species described worldwide. They are the sister group to the family Brentidae. T ...
), or brentids. Others are
mimetic Mimesis (; grc, μίμησις, ''mīmēsis'') is a term used in literary criticism and philosophy that carries a wide range of meanings, including '' imitatio'', imitation, nonsensuous similarity, receptivity, representation, mimicry, the ...
and imitate less closely related
Polyphaga Polyphaga is the largest and most diverse suborder of beetles. It comprises 144 families in 16 superfamilies, and displays an enormous variety of specialization and adaptation, with over 350,000 described species, or approximately 90% of the b ...
beetles such as
Lycidae The Lycidae are a family in the beetle order Coleoptera, members of which are commonly called net-winged beetles. These beetles are cosmopolitan, being found in Nearctic, Palearctic, Neotropical, Afrotropical, Oriental, and Australian ecoregions ...
or Pyrochroidae. The adults are distinguished from other ancient weevil lineages by some characteristic traits: The fore
tibia The tibia (; ), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outside of the tibia); it connects ...
of belines is unusual in that it has a comb of bristles (
seta In biology, setae (singular seta ; from the Latin word for "bristle") are any of a number of different bristle- or hair-like structures on living organisms. Animal setae Protostomes Annelid setae are stiff bristles present on the body. Th ...
e) in an
apical Apical means "pertaining to an apex". It may refer to: *Apical ancestor, refers to the last common ancestor of an entire group, such as a species (biology) or a clan (anthropology) *Apical (anatomy), an anatomical term of location for features loc ...
groove opposite the tarsal articulation; this is used for grooming. Microscopically, the
spermatheca The spermatheca (pronounced plural: spermathecae ), also called receptaculum seminis (plural: receptacula seminis), is an organ of the female reproductive tract in insects, e.g. ants, bees, some molluscs, oligochaeta worms and certain other ...
l gland is several times longer than the spermathecal capsule. Most also have only a few (four or fewer) setae on each
mandible In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower tooth, teeth in place. The mandible sits beneath the maxilla. It is the only movabl ...
(more in
Aglycyderini Aglycyderini is a tribe of belids, primitive weevils of the family Belidae. Like in other belids, their antennae are straight, not elbowed as in the true weevils ( Curculionidae). They occur only on the Pacific Islands and in the Macaronesian ...
), and the
pronotum The prothorax is the foremost of the three segments in the thorax of an insect, and bears the first pair of legs. Its principal sclerites (exoskeletal plates) are the pronotum (dorsal), the prosternum (ventral), and the propleuron (lateral) on ea ...
is constricted at the tip (not in '' Oxycorynus'', '' Parallocorynus'' and ''
Rhopalotria ''Rhopalotria'' is a genus of cycad weevils in the beetle family Belidae. There are about six described species in ''Rhopalotria''. Species These six species belong to the genus ''Rhopalotria'': * ''Rhopalotria dimidiata'' Chevrolat, 1878 * '' ...
''). The endocarina is V-shaped. In most, the antennae have a retractable membrane at the base (not in '' Parallocorynus'' and ''
Rhopalotria ''Rhopalotria'' is a genus of cycad weevils in the beetle family Belidae. There are about six described species in ''Rhopalotria''. Species These six species belong to the genus ''Rhopalotria'': * ''Rhopalotria dimidiata'' Chevrolat, 1878 * '' ...
''), and the
alimentary canal The gastrointestinal tract (GI tract, digestive tract, alimentary canal) is the tract or passageway of the digestive system that leads from the mouth to the anus. The GI tract contains all the major organs of the digestive system, in humans ...
has caeca distributed all over at random (in two clean bundles in
Aglycyderini Aglycyderini is a tribe of belids, primitive weevils of the family Belidae. Like in other belids, their antennae are straight, not elbowed as in the true weevils ( Curculionidae). They occur only on the Pacific Islands and in the Macaronesian ...
and
Metrioxenini Metrioxenini (metrioxenines) are a tribe of belids, primitive weevils of the family Belidae, containing about 30 species. They are only found in two widely separated areas, Southeast Asia extending to Indonesia, and South Africa. In the Paleo ...
).


Ecology

Adults usually eat
pollen Pollen is a powdery substance produced by seed plants. It consists of pollen grains (highly reduced microgametophytes), which produce male gametes (sperm cells). Pollen grains have a hard coat made of sporopollenin that protects the gametophyt ...
; the
larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. The ...
e feed on the wood of diseased or dying plants or on deadwood,
flower bud In botany, a bud is an undeveloped or embryonic shoot and normally occurs in the axil of a leaf or at the tip of a stem. Once formed, a bud may remain for some time in a dormant condition, or it may form a shoot immediately. Buds may be speci ...
s or fruits. Though they may appear to be pests because of their association with plants dying off, belids tend to avoid healthy plants. They may occur as a secondary complication in plants already affected by some pest or
disease A disease is a particular abnormal condition that negatively affects the structure or function of all or part of an organism, and that is not immediately due to any external injury. Diseases are often known to be medical conditions that a ...
however. But altogether, belids (as opposed to many other weevils) are normally harmless and can be beneficial, the adults as
pollinator A pollinator is an animal that moves pollen from the male anther of a flower to the female stigma of a flower. This helps to bring about fertilization of the ovules in the flower by the male gametes from the pollen grains. Insects are the maj ...
s and the larvae as
indicator species A bioindicator is any species (an indicator species) or group of species whose function, population, or status can reveal the qualitative status of the environment. The most common indicator species are animals. For example, copepods and other sma ...
for bad condition of their host plants. In
Florida Florida is a state located in the Southeastern region of the United States. Florida is bordered to the west by the Gulf of Mexico, to the northwest by Alabama, to the north by Georgia, to the east by the Bahamas and Atlantic Ocean, and to ...
for example, ''
Rhopalotria slossoni ''Rhopalotria'' is a genus of cycad weevils in the beetle family Belidae. There are about six described species in ''Rhopalotria''. Species These six species belong to the genus ''Rhopalotria'': * ''Rhopalotria dimidiata'' Chevrolat, 1878 * ''R ...
'' is important in maintaining populations of the rare
Coontie ''Zamia integrifolia'', also known as coontie palm is a small, tough, woody cycad native to the southeastern United States (in Florida and Georgia), the Bahamas, Cuba, and the Cayman Islands. Description ''Zamia integrifolia'' produces reddish ...
(''Zamia pumila''); indirectly, the Atala butterfly (''
Eumaeus atala ''Eumaeus atala'', also known as the Atala butterfly or coontie hairstreak, is a small colorful butterfly in the family Lycaenidae. It is found in southeastern Florida (including the Florida Keys) in the United States, and the Bahamas, Cuba, and ...
'') also depends on this beetle. The original host plants of belids were probably
Araucariaceae Araucariaceae – also known as araucarians – is an extremely ancient family of coniferous trees. The family achieved its maximum diversity during the Jurassic and Cretaceous periods and the early Cenozoic, when it was distributed almost worldw ...
,
conifer Conifers are a group of conifer cone, cone-bearing Spermatophyte, seed plants, a subset of gymnosperms. Scientifically, they make up the phylum, division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single ...
s which were extremely common during the time when the family evolved. Some modern
genera Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomenclat ...
also feed on
angiosperm Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...
s, usually rather old groups like
Balanophoraceae The Balanophoraceae are a subtropical to tropical family of obligate parasitic flowering plants, notable for their unusual development and formerly obscure affinities. In the broadest circumscription, the family consists of 16 genera. Alternativ ...
,
Myrtaceae Myrtaceae, the myrtle family, is a family of dicotyledonous plants placed within the order Myrtales. Myrtle, pōhutukawa, bay rum tree, clove, guava, acca (feijoa), allspice, and eucalyptus are some notable members of this group. All speci ...
or
Vitaceae The Vitaceae are a family of flowering plants, with 14 genera and around 910 known species, including common plants such as grapevines (''Vitis'' spp.) and Virginia creeper (''Parthenocissus quinquefolia''). The family name is derived from the ge ...
, on
cycad Cycads are seed plants that typically have a stout and woody (ligneous) trunk (botany), trunk with a crown (botany), crown of large, hard, stiff, evergreen and (usually) pinnate leaves. The species are dioecious, that is, individual plants o ...
s, or on
palm Palm most commonly refers to: * Palm of the hand, the central region of the front of the hand * Palm plants, of family Arecaceae **List of Arecaceae genera * Several other plants known as "palm" Palm or Palms may also refer to: Music * Palm (ba ...
s. The Pacific genus '' Proterhinus'' has undergone a vigorous
adaptive radiation In evolutionary biology, adaptive radiation is a process in which organisms diversify rapidly from an ancestral species into a multitude of new forms, particularly when a change in the environment makes new resources available, alters biotic int ...
on the
Hawaiian Islands The Hawaiian Islands ( haw, Nā Mokupuni o Hawai‘i) are an archipelago of eight major islands, several atolls, and numerous smaller islets in the North Pacific Ocean, extending some from the island of Hawaii in the south to northernmost Kur ...
and evolved to utilize a wide range of the limited diversity of plants found there.


Systematics

The
subfamilies In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus. Standard nomenclature rules end subfamily botanical names with "-oideae", and zoologi ...
have each, at various times, been considered as separate families, but they are grouped together in most recent classifications. There are three main living lineages, variously considered three or, as here, two subfamilies, with the
tribe The term tribe is used in many different contexts to refer to a category of human social group. The predominant worldwide usage of the term in English language, English is in the discipline of anthropology. This definition is contested, in p ...
Aglycyderini Aglycyderini is a tribe of belids, primitive weevils of the family Belidae. Like in other belids, their antennae are straight, not elbowed as in the true weevils ( Curculionidae). They occur only on the Pacific Islands and in the Macaronesian ...
sometimes considered a distinct subfamily. Other classifications treat the
Oxycoryninae Oxycoryninae are subfamily of primitive weevils of the family Belidae, but sometimes treated as a distinct family Oxycorynidae. Like in other belids, their antennae are straight, not elbowed as in the true weevils ( Curculionidae), and their l ...
as distinct family Oxycorynidae. A prehistoric subfamily only known from
Mesozoic The Mesozoic Era ( ), also called the Age of Reptiles, the Age of Conifers, and colloquially as the Age of the Dinosaurs is the second-to-last era of Earth's geological history, lasting from about , comprising the Triassic, Jurassic and Cretaceo ...
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
s are the Eobelinae.


Extinct taxa

* †'' Sinoeuglypheus'' Yu et al. 2019
Daohugou The Haifanggou Formation (), also known as the Jiulongshan Formation (), is a fossil-bearing rock deposit located near Daohugou () village of Ningcheng County, in Inner Mongolia, northeastern China. The formation consists of coarse conglomerates ...
, China,
Callovian In the geologic timescale, the Callovian is an age and stage in the Middle Jurassic, lasting between 166.1 ± 4.0 Ma (million years ago) and 163.5 ± 4.0 Ma. It is the last stage of the Middle Jurassic, following the Bathonian and preceding the ...
* Subfamily Belinae Schoenherr 1826 ** †tribe Davidibelini Legalov 2015 *** †'' Davidibelus'' Zherikhin and Gratshev 2004
Crato Formation The Crato Formation is a geologic formation of Early Cretaceous ( Aptian) age in northeastern Brazil's Araripe Basin. It is an important Lagerstätte (undisturbed fossil accumulation) for palaeontologists. The strata were laid down mostly duri ...
, Brazil,
Aptian The Aptian is an age in the geologic timescale or a stage in the stratigraphic column. It is a subdivision of the Early or Lower Cretaceous Epoch or Series and encompasses the time from 121.4 ± 1.0 Ma to 113.0 ± 1.0 Ma (million years ago), a ...
** †subfamily Montsecbelinae Legalov 2015 *** †tribe Montsecbelini Legalov 2015 **** †'' Montsecbelus'' Zherikhin and Gratshev 1997 La Pedrera de Rúbies Formation, Spain,
Barremian The Barremian is an age in the geologic timescale (or a chronostratigraphic stage) between 129.4 ± 1.5 Ma (million years ago) and 121.4 ± 1.0 Ma). It is a subdivision of the Early Cretaceous Epoch (or Lower Cretaceous Series). It is preceded ...
** Subfamily Oxycoryninae Schoenherr 1840 *** †'' Khetana'' Zherikhin 1993 Emanra Formation, Russia,
Turonian The Turonian is, in the ICS' geologic timescale, the second age in the Late Cretaceous Epoch, or a stage in the Upper Cretaceous Series. It spans the time between 93.9 ± 0.8 Ma and 89.8 ± 1 Ma (million years ago). The Turonian is preceded by t ...
***Clade Aglycyderitae Wollaston 1864 **** †tribe Distenorrhinoidini Legalov 2009 ***** †'' Distenorrhinoides'' Gratshev and Zherikhin 2000 La Pedrera de Rúbies Formation, Spain, Barremian *** Clade Allocorynitae Sharp 1890 **** Tribe Allocorynini Sharp 1890 ***** †'' Pleurambus'' Poinar and Legalov 2014
Dominican amber Dominican amber is amber from the Dominican Republic derived from resin of the extinct tree '' Hymenaea protera''. Dominican amber differentiates itself from Baltic amber by being nearly always transparent, and it has a higher number of fossil inc ...
,
Miocene The Miocene ( ) is the first geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and means "less recen ...
****†tribe Palaeorhopalotriini Legalov 2013 *****†'' Palaeorhopalotria'' Legalov 2013 Monteils Formation, France,
Priabonian The Priabonian is, in the ICS's geologic timescale, the latest age or the upper stage of the Eocene Epoch or Series. It spans the time between . The Priabonian is preceded by the Bartonian and is followed by the Rupelian, the lowest stage of t ...
***Clade Oxycorynitae Schoenherr 1840 ****Tribe
Metrioxenini Metrioxenini (metrioxenines) are a tribe of belids, primitive weevils of the family Belidae, containing about 30 species. They are only found in two widely separated areas, Southeast Asia extending to Indonesia, and South Africa. In the Paleo ...
Voss 1953 *****Subtribe Metrioxenina Voss 1953 ******†'' Archimetrioxena'' Voss 1953
Baltic amber The Baltic region is home to the largest known deposit of amber, called Baltic amber or succinite. It was produced sometime during the Eocene epoch, but exactly when is controversial. It has been estimated that these forests created more than 1 ...
,
Eocene The Eocene ( ) Epoch is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene' ...
*****Subtribe Zherichinixenina Legalov 2009 ******†'' Paltorhynchus'' Scudder 1893
Green River Formation The Green River Formation is an Eocene geologic formation that records the sedimentation in a group of intermountain lakes in three basins along the present-day Green River in Colorado, Wyoming, and Utah. The sediments are deposited in very fine ...
,
Florissant Formation The Florissant Formation is a sedimentary geologic formation outcropping around Florissant, Teller County, Colorado. The formation is noted for the abundant and exceptionally preserved insect and plant fossils that are found in the mudstones and ...
, United States, Eocene ******†'' Succinometrioxena'' Legalov 2012 Baltic amber, Eocene


References


External links

*
Belidae on web.archive.org

Images of Belidae species in New Zealand
{{DEFAULTSORT:Belidae Beetle families