Curculionoidea
| Curculionoidea | ||||||||||||
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Apoderus longicollis |
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| Systematics | ||||||||||||
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| Scientific name | ||||||||||||
| Curculionoidea | ||||||||||||
| Latreille , 1802 |
The Curculionoidea are a global superfamily of beetles within the suborder Cucujiformia . In addition to the well-known weevils and bark beetles , it includes a number of families, most of which were previously considered part of the weevils. The superfamily comprises 62,000 species (as of 2007), including 51,000 weevils alone. It is assumed, however, that there are a particularly large number of scientifically undescribed species in this family, mostly in the tropical rainforests, so that the actual number of species is probably many times that. The number and delimitation of the families of the Curculionoidea has not yet been finally clarified within science and is controversial.
features
Beetle
The Curculionoidea comprises an enormous variety of beetles with almost all conceivable modifications and changes in body structure, from tiny, millimeter-sized species to species about 10 centimeters long. What they all have in common is the nutrition of the larvae as herbivores (phytophagus) or, alternatively, from living or dead wood. A characteristic and most striking common feature is the trunk, anatomically called rostrum, an extension of the head capsule forwards or downwards, at the tip of which the mouthparts are located. The trunk can be long and extremely thin (up to several body lengths) or short and stocky. A few groups, in particular the Scolytinae (bark beetles) and the Platypodinae (heartwood beetles), have regressed the proboscis secondarily. On the other hand, trunk-like extensions of the head capsule also occur in other families, so that the trunk alone is not a sure distinguishing feature. The main biological purpose of the long and thin trunk shapes is mostly seen in the laying of eggs. With its help, the female can eat a long and thin cavity in the plant tissue, in which it can then lay its eggs in a protected place (with a long and flexible, fold-out tube). Accordingly, the trunk is often developed differently between the sexes ( sexual dimorphism ). Sitting at the tip of the proboscis mouthparts . These are of the normal, biting-chewing type, but have a number of characteristic transformations. A free labrum is only formed in the original families Nemonychidae and Attelabidae , mostly it is fused with the clypeus . The mandibles are almost always of the normal, biting type, they can be short and stocky or long and slender. In many Curculionoidea they have a horn-shaped or cone-shaped extension on the outside, which is only present in newly hatched beetles and is shed later (at a predetermined breaking point), so that only a rough scar remains. The function of the extension is that the newly hatched imago can free itself from its pupa chamber. Some groups, such as the subfamily Rhynchitinae, have externally toothed (exodonte) mandibles. In the genus Curculio and relatives, the mandibles do not work against each other, but are moved up and down. There are two types of labium and maxilla formation . Either the maxillae are covered by the enlarged prementum of the labium, or they sit on the side of the prementum and are freely visible from below. While the labial palps are mostly normal, the maxillary palps in all but the most primitive families are transformed into a short and rigid, one-part process. The antennas are usually located on the side of the trunk, they can also be pivoted on the top or bottom. They consist of eleven members (with some species also less secondary). The last three members are usually transformed into a clear, widened antennae lobe, which is tightly closed in the Curculionidae , but rather loosely structured in the other families. It is seldom indistinct, then its limbs stand out from the other shiny segments by a matt surface. In the original families, called "Orthoceri" (also "Recticornes") after this characteristic, the base member ( Scapus ) of the antenna is short and the antenna is attached to it. In the modified "Gonatoceri" (or "Infracticornes") the scapus is elongated, the antennae flagella adjoins it at a clear angle (just like the antennae of the ants ). These angled ("kneeled") feelers allow the animal to examine the surface in front of the mouthparts. In addition, the scapus can be kicked back, often even inserted into a groove in the trunk, and is thus not in the way of gnawing cavities. There is also a pair of complex eyes on the head , which can be varied in many ways. The head is almost always held forward so that the mouthparts point forward and not downward (prognath). To do this, it is closed on the belly by an additional sclerite called a "gula". In the Curculionoidea, with the exception of the basal families, the gula is shifted towards the inside of the head, so that only one seam remains visible on the outside, or fused with the head capsule without any noticeable seams.
The Curculionoidea are mostly hard sclerotized animals. The body outline is mostly oval, with high arched wing covers (elytra) and a flat underside. The wing covers are firmly attached to the rest of the body contour. In wingless animals such as the Brachyceridae , the upper side (abdomen and rear thoracic section) underneath is even soft-skinned and not sclerotized. In some groups, such as the Cryptorhynchinae , legs and trunk can be placed in recesses, which results in a hard-armored shape on the outside without the possibility of predators. In the morphologically modified families, the abdomen of the abdomen is reshaped in response to the armor. The anterior sternites , at least the first and second, are firmly and immovably fused together. The third and fourth sternite form a kind of hinge joint, which enables the animal to open the rear end (for dropping excrement or for laying eggs). In some groups the wing covers are a little shortened at the end. The last tergite , then called “ pygidium ”, is exposed here; in this case it is also hard sclerotized. The legs largely correspond to the basic plan of the insects. In some families (e.g. Brentidae ) the trochanter is noticeably elongated, in the Curculionidae it is short and triangular with angled attachment of the legs. The thighs ( femora ) are often club-shaped, sometimes toothed. The attachment of the tarsi to the rails is peculiarly changed in many groups, often with conspicuous hook-shaped appendages. The construction of the tarsi is typical. These are five-limbed, the third limb is greatly expanded at the bottom and has a striking trimming of hair that is used to hold on to smooth surfaces. The fourth link is greatly shortened and largely covered by the third, but only very rarely completely regressed or fused. Only the small family Raymondionymidae has four-membered tarsi.
Five sternites are freely visible on the abdomen of the Curculionoidea. The first visible sternite is that of the third segment, the first two are reduced and hidden in the cavity of the rear hips. Accordingly, seven tergites are recognizable, occasionally eight (only nine in the females of the Nemonychidae ). The eighth sternite is drawn in and hidden in the genital chamber. In the original families of the Curculionoidea, the ninth tergite within the genital chamber is also preserved in the males; it is referred to here as the "tectum". A modified group is characterized by a different structure of the genital segments of the male, in which the tectum has regressed. By and large, the families with tectum correspond to those with straight antennae, those without tectum to those with kneeling antennae. However, there is a group of families with kneeling antennae without tectum (families Nanophyidae , Brachyceridae , Erirhinidae , Dryophthoridae , Raymondionymidae and Cryptolaryngidae ), whose taxonomic position is often disputed.
Larvae
The larvae of the Curculionoidea are more or less elongated and rolled and usually slightly curved on the abdomen. Usually the entire body with the exception of the head capsule and the appendages, or at least the abdomen, is soft-skinned and hardly sclerotized. Almost always, with the exception of a few original families, they are legless. For example, in some Nemonychidae and Attelabidae, larvae with multi-limbed legs (with a claw at the end) occur. Many free-living larvae have warty appendages that are known as "ambulatory ampoules" for locomotion. The larvae of most Curculionoidea, however, live inside plant tissue or wood and have no appendages whatsoever for movement. They can crawl by propping themselves up inside the cavity and pushing forward. In addition to segmentation, the body is usually divided into secondary ringlets ("plicae").
The larvae have a clearly sclerotized head capsule, which in some groups can be withdrawn into the trunk. The mouthparts are directed downwards (hypognathic). Larval eyes (stemmata) are usually missing. The antennae are also largely reduced, mostly they consist of an unsclerotized, pillow-shaped structure that carries a large cone and several small sensillae. Below the mouthparts there is a heavily sclerotized process that consists of the reshaped hypopharynx. The larvae usually have prominent, biting mandibles. The two stores are fused at the maxilla. As a rule, the maxillae sit on both sides of the labium and lie in one plane with it. There are two- or three-part maxillary and labial palps.
There are usually eight pairs of spiracles on the larvae's abdomen , but not all of them are always functional. In many aquatic larval forms, they have thorn-shaped appendages. These serve to pierce the air conduction tissue ( aerenchyma ) of aquatic plants. Processes (styli or urogomphi) at the rear end of the larvae are never developed.
Way of life
Curculionoidea are common in all habitats in which plants grow and insects can occur at all. Almost every plant species is eaten by them to varying degrees. Many species specialize in one or a few related plant species ( monophag or oligophag ). Some families and subfamilies, especially the presumably closely related Cossoninae , Scolytinae and Platypodinae drill in wood. To unlock this difficult substrate, they cultivate wood-degrading fungi, which they usually carry with them in special structures on their bodies. Most of the other Curculionoidea investigated thereupon also have symbiotic microorganisms that support digestion. Most larval forms drill inside plant tissue, only a few forms live freely on the surface. The adult beetles usually feed on plant tissue as well. Numerous forms feed on pollen and visit flowers to do so. With a few exceptions, however, they play a much smaller role in pollination than other groups, especially hymenoptera (Hymenoptera). The Attelabidae family specializes in the fruiting bodies of mushrooms. Numerous species live on the ground as adults and larvae, they feed on z. Some of them eat roots, others eat algae or lichen coatings.
Systematics
The monophyly of the superfamily Curculionoidea is generally considered to be well established and is not doubted in any more modern work. The superfamily Chrysomeloidea (with the large families of the leaf beetles (Chrysomelidae) and the longhorn beetles (Cerambycidae)), with which they share the special structure of the tarsi, is considered a sister group . The grouping is called "Phytophaga" according to the way of life. However, according to molecular family trees (based on the comparison of homologous DNA sequences), it is not certain whether the undeniably closely related superfamilies are really sister groups.
The traditional subdivision of the Curculionoidea into families has been rearranged in recent years due to new findings, especially regarding morphology. Numerous groups, especially from the more basal department "Orthoceri" (with straight antennae), which had traditionally been run as subfamilies, are now considered independent families. At the same time, the traditional families Scolytidae and Platypodidae, which differ mainly in the lack of proboscis, were downgraded to subfamilies within the Curculionidae. The subdivision of the large family Curculionidae is still problematic. Many of the traditional subfamilies are almost certainly polyphyletic, but will continue to be used because no one has yet been able to imagine a better system.
Two systems are currently in use side by side, one with 16 more narrowly defined families (plus two extinct families that are only known to be fossilized), one with only 8 more defined families. The difference between the two is in most cases less due to completely different phylogenetic systems, but more to a different understanding of the ranks, so that the differences of opinion are smaller than initially assumed. The more detailed system is based on the catalog by Alonso-Zarazaga and Lyal; it was used for most recent German-language publications. The other division into Oberprieler, Marvaldi and Anderson is at least as common. Your z. B. followed in the well-founded current list of beetle families. The authors of the catalog (Alonso-Zarazaga and Lyal), in which 18 families are distinguished, are also co-authors of this list, which only recognizes nine families. It is then to be expected that the system will experience changes in the next few years.
In the following list, all recent families that are listed in the catalog are taken into account. This appears necessary regardless of the authorization of the classification, since these names are used in numerous current works. Those families whose status is still controversial are placed indented below the family in which they may be classified as a subfamily by other authors. If one follows this alternative view, the name ending would change from -idae to -inae (e.g. Apioninae instead of Apionidae). The families that join below the Curculionidae are united by numerous authors in a broad family Curculionidae.
- Nemonychidae
- Anthribidae
- Attelabidae
- Caridae
- Belidae
- Brentidae
- Curculionidae
- Dryophthoridae
- Brachyceridae
Fossils and Evolution
The oldest fossils that can be assigned to the superfamily come from the Jurassic . They were found in the Karatau fossil deposit (today's spelling Qaratau ) in Kazakhstan, from which almost 50 species have already been discovered (some still undescribed). Even older finds from the Triassic are assigned to the extinct family Obrieniidae , which, although it has a proboscis and culled antennae, is no longer considered to belong to the Curculionoidea today. The Obrieniidae were probably Archostemata , the trunk formation probably occurred convergent . In the fossil Curculionoidea, the trunk sits on the underside of the head. Most fossils can be classified according to their characteristic expression in the recent family Nemonychidae, but represent extinct subfamilies. Some fossil species from the same deposit have been assigned to the Belidae family. It is possible that they are members of the core group (of the subfamily Eobelinae), from which many of the modern families descended. There are numerous fossils from the chalk , particularly those preserved in amber, which appeared for the first time in this epoch. The first weevil with kneeling antennae comes from Burmese amber from the early Cretaceous.
The fossil Curculionoidea from Jurassic and Chalk are presumably bound exclusively or predominantly to gymnosperms as food plants. The surviving representatives of the families that were widespread at the time, which today form species-poor relic groups, are predominantly tied to conifers. Numerous researchers support the hypothesis that the great diversity of species of today's Curculionoidea (and also the Chrysomeloidea) can be traced back to the origin of the angiosperms in the Cretaceous. The first species that passed over to them as herbivores had almost no competitors and then, together with their food plants, could have undergone rapid adaptive radiation . This hypothesis, although convincing in principle, is considered by some researchers to be insufficiently proven and ultimately unprovable.
Individual evidence
- ^ A b Rolf G. Oberprieler, Adriana E. Marvaldi, Robert S. Anderson (2007): Weevils, weevils, weevils everywhere. Zootaxa 1668: 491-520.
- ↑ cf.
- ↑ a b c R. T. Thompson (1992): Observations on the morphology and classification of weevils (Coleoptera, Curculionoidea) with a key to major groups. Journal of Natural History, 26 (4): 835 - 891. doi : 10.1080 / 00222939200770511
- ↑ MA Alonso-Zarazaga (2007): On terminology in Curculionoidea (Coleoptera). Boletín Sociedad Entomológica Aragonesa 40: 210.
- ↑ Brenda Μ. May (1993): Larvae of Curculionoidea (Insecta: Coleoptera): a systematic overview. In: Lincoln, NZ (editor): Fauna of New Zealand, No. 28. Manaaki Whenua Press.
- ↑ Brenda M. May (1994): An introduction in the immature stages of Australian Curculionoidea. In: Elwood C. Zimmerman: Australian Weevils (Coleoptera: Curculionoidea): Brentidae, Eurhynchidae, Apionidae and a Chapter on Immature Stages by Brenda May (Australian Weevils Series) (Volume II). CSIRO Publishing.
- ↑ Adriana E. Marvaldi (2003): Key to larvae of the South American subfamilies of weevils (Coleoptera, Curculionoidea). Revista Chilena de Historia Natural 76: 603-612.
- ↑ cf. Cladogram in Toby Hunt, Johannes Bergsten, Zuzana Levkanicova, Anna Papadopoulou, Oliver St. John, Ruth Wild, Peter M. Hammond, Dirk Ahrens, Michael Balke, Michael S. Caterino, Jesús Gómez-Zurita, Ignacio Ribera, Timothy G. Barraclough , Milada Bocakova, Ladislav Bocak, Alfried P. Vogler (2007): A Comprehensive Phylogeny of Beetles Reveals the Evolutionary Origins of a Superradiation. Science 318: 1913-1916. doi : 10.1126 / science.1146954
- ↑ MA Alonso-Zarazaga & CHC Lyal (1999): A World Catalog of families and genera of Curculionoidea (Insecta: Coleoptera) excluding (Scolytidae and Platypodidae). (Entomopraxis). 315 pp. ISBN 978-8-460-59994-4
- ↑ Patrice Bouchard, Yves Bousquet, Anthony E. Davies, Miguel A. Alonso-Zarazaga, John F. Lawrence, Chris HC Lyal, Alfred F. Newton, Chris AM Reid, Michael Schmitt, S. Adam Ślipiński, Andrew BT Smith (2011 ): Family-group names in Coleoptera (Insecta). ZooKeys 88: 1-972. doi : 10.3897 / zookeys.88.807
- ↑ Vadim G. Gratshev & Vladimir V. Zherikhin (2003): The fossil record of weevils and related beetle families (Coleoptera, Curculionoidea). Acta zoologica cracoviensia, 46 (suppl.- Fossil Insects): 129-138.
- ↑ George Poinar Jr. (2006): Mesophyletis calhouni (Mesophyletinae), a new genus, species, and subfamily of early cretaceous weevils (Coleoptera: Curculionoidea: Eccoptarthridae) in burmese amber. Proceedings of the Entomological Society of Washington 108: 878-884. [1]
- ^ BD Farrell (1998): “Inordinate fondness” explained: why are there so many beetles? Science 281: 553-557.
- ↑ AE Marvaldi, AS Sequeira, CW O'Brien, BD Farrell (2002): Molecular and morphological phylogenetics of weevils (Coleoptera: Curculionoidea): do niche shifts accompany diversification? Systematic Biology 51: 761-785.
- ↑ DD McKenna, AS Sequeira, AE Marvaldi, BD Farrell (2009): Temporal lags and overlap in the diversification of weevils and flowering plants. Proceedings of the National Academy of Sciences USA 106: 7083-7088.
- ↑ Nico M. Franz & Michael S. Engel (2010): Can higher-level phylogenies of weevils explain their evolutionary success? A critical review. Systematic Entomology 35: 597-606.
