Scrape

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Scrape
Madagascar hissing cockroach (Gromphadorrhina portentosa)

Madagascar hissing cockroach
( Gromphadorrhina portentosa )

Systematics
Sub-stem : Trachea (Tracheata)
Superclass : Six-footed (Hexapoda)
Class : Insects (Insecta)
Subclass : Flying insects (Pterygota)
Superordinate : Dictyoptera (parent)
Order : Scrape
Scientific name
Blattodea
Brunner von Wattenwyl , 1882
American cockroach ( Periplaneta americana )

The cockroaches (Blattodea) are an order of hemimetabolic insects that populate the tropics and subtropics with around 4600 species . There are also some species that have been introduced into human dwellings, which are then considered pests or at least annoying. Cockroaches have meanwhile also found their way into terraristics . Some of them are kept and bred here as food for other species, but some are also for their own sake.

features

size

The body length of the cockroaches varies between 5 and more than 100 mm. For a long time the largest recent species was Megaloblatta blaberoides with a total length of 100 mm, a body length of about 66 mm and a wingspan of a maximum of 170 mm. The Miroblatta baai from Borneo , which was only discovered in 2004, reaches a comparable size. In terms of body mass, the approximately 80 mm long, wingless Australian Macropanesthia rhinoceros leads the way with around 35 g. The hissing cockroaches of the genus Gromphadorhina , popular as pets, also reach an impressive size. In addition to the well-known Madagascar hissing cockroach ( Gromphadorrhina portentosa ), the slightly larger Gromphadorhina grandidieri with up to 85 mm should be mentioned. The body size often varies considerably within a species. In contrast to many other orders, no fossils have been found in the cockroaches that would exceed the current species in size.

Body shape and body sections

The body of the animals is almost always flattened. Some genera, especially many of the subfamily Perisphaeriinae , are higher arched and have the ability to ball themselves. There are also special deviations in body shape in species that drill into soil or wood, in cave dwellers or in ants or termite nests.
The mouthparts are biting and chewing and the antennae are usually long and thread-like, they are the most important sense organs in many species. In addition, many representatives have extremely sensitive vibration sensors (subgenual organs) in their legs. The head is very mobile and usually sits somewhat hanging under the neck shield (Scutum or pronotum ), the mouthparts are directed downwards (hypognath). The complex eyes can be present in different sizes depending on the species. They are usually flat and do not protrude from the contour of the head. In species with larger eyes, these often surround the antennae roots in a kidney shape. The large pronotum that covers the entire head is striking. It is usually shaped transversely oval, but can also be rounded triangular and widened towards the rear. Especially with the males it often has humps, rarely horns, with which the males fight for females with mutual rams. The three pairs of legs usually have flattened hips and thighs. The bases of the hips are very close together on the ventral side. The thighs can often be placed in recesses in the hips. The tarsi are five-part. The first four parts of the tarsi usually have pillow-shaped projections called plantula or pulvilli, another extension of this type, the arolium , lies between the two claws. This allows the animals to find hold on smooth surfaces effectively, some species can easily walk on vertical glass panes. Inside the body, a muscular chewing stomach (Proventriculus) is characteristic, which has teeth on its inside. The broadly seated abdomen ( abdomen ) has ten segments ( sclerite ), whereby individual segments can be greatly reduced. The first abdominal sternite is often reduced on the underside and the eighth and ninth tergite on the upper side of the abdomen . In males nine abdominal sternites are usually visible from the outside, in females seven. The reason for this is that the eighth and ninth segments are drawn into the abdomen. The abdominal appendages ( cerci ), which are usually multi-limbed, are clearly visible . They are less common or absent. The cerci are designed as mechanical sensory organs that warn above all of the approach of predators . The end of the abdomen is designed differently in males and females. The females have a brood chamber in which the egg packet ( ootheca ) is formed and often carried. The non-functional remainder of the original laying apparatus ( ovipositor ) is hidden in the incubation chamber . In the males, the ninth sternite is drawn far back as a subgenital plate, with various, often hook-shaped appendages on the upper side. The attachments are always extremely asymmetrically built differently between the left and right side. They are used to hold the female in place during mating. The shape of these genital appendages is the most important feature for species identification.

Most cockroaches are inconspicuously brown or yellowish in color. In different species there are bright, red or orange, sometimes metallic iridescent blue or green colors; as a rule, these are poisonous species or those equipped with defensive glands. Different tropical species imitate beetles in body shape (e.g. genus Prosoplecta ).

Wings, flight and locomotion

Like most insects, cockroaches have two pairs of membranous wings, the front, somewhat coarser and leather-like cover wings being designed as tegmina and covering the more delicate rear wings. The rear wings have a considerably larger area because the rear part of them is expanded into a large anal fan. They are essential to flight. In the rest position, the wings are carried flat on top of one another on the abdomen, with the rear part of the rear wings being folded in like a fan. The forewings usually overlap a little, with the left always lying above the right. Some species (e.g. Diploptera punctata ) fold their hind wings once across when they are at rest. The wing veins of the cockroaches are characterized by very many longitudinal veins. A curved vein can be seen particularly clearly near the front edge, which is known as the subcosta . Often wings are only developed in males, while these are stunted in females. In addition, completely wingless species are not uncommon. Most cockroach species have little flight ability. Some species have no flight ability despite fully developed wings (e.g. Blattella germanica ), they lack the necessary, strong flight muscles. Some types, such as B. the American cockroach ( Periplaneta americana ), but are quite good fliers. Since cockroaches have very well developed legs, they are usually good and above all fast runners. Saltoblattella montistabularis was described from South Africa in 2009 , a wingless species with jumping ability that was only recently discovered.

Occurrence and nutrition

There are currently around 4,600 known species of cockroach. As a very warmth-loving group, they have their distribution focus in the tropics. In the subtropics already fewer species live in the European Mediterranean z. B. only about 130 species. Although they generally prefer humid habitats, there are also species that specialize in dry habitats, including deserts (e.g. family Corydiidae ). These are then buried during the day and only come out at night when the humidity is high. Only 15 species live
in Central Europe . In Germany only six species of the subfamily Ectobiinae ( forest cockroaches ) are free- living . Depending on the species, habitats are forests, forest edges or dry grasslands. The species living in Germany have a two-year development cycle, the discarded ootheca overwinter, the young nymphs hatch in the following spring. The other species have been introduced and live exclusively in heated houses or, under special conditions, outdoors (e.g. on rubbish dumps with heat generation). Often such species are only found in permanently very warm surroundings. They are particularly found in bakeries, large kitchens, greenhouses and terrariums. They can also occur in apartments and become very annoying here; this is much less common in Central Europe than in warmer countries. The best-known species in Central Europe are the common cockroach ( Blatta orientalis ), also known as the cockroach, and the German cockroach ( Blattella germanica ). They often find ideal living conditions in human dwellings and have therefore been deported all over the world. Even native species such as the common wood cockroach ( Ectobius lapponicus ) can be a nuisance in houses in exceptional cases.
Cockroaches feed on a wide variety of substances of vegetable and animal origin. The synanthropic species in particular are omnivores. Most of the species living in the wild are saprophagous and eat z. B. Fall foliage on the forest floor. A number of species (e.g. Cryptocercus ) specialize in rotting deadwood as food. Others scrape off algae and lichen from wood and other surfaces. Some species use soft parts of plants or seedlings in addition to other food. They can occasionally become harmful in greenhouses. However, there are no known species that would specifically eat green leaves or other parts of living plants. Many canopy tropical species use pollen and honeydew . Some species are facultative predators from other arthropods . Cannibalism is widespread in animals kept in terrariums .

Reproduction and development

Reproductive strategy, dating and mating

Two oothecae of the American cockroach ( Periplaneta americana )

The females of the cockroaches attract males over long distances with a fragrance ( pheromone ). This substance has been identified in the German cockroach ( Blatella germanica ) (Blattellachinon: gentisyl-quinone-isovalerate). The males also have "scent glands" on the first tergites of the abdomen, from which they emit a pairing-stimulating scent. In many species, the male lures the female onto his back by means of this fragrance. If both animals are aligned in parallel, the male grasps the female from below with his hook-shaped genital appendages. At the end of the pairing, both animals are then oriented in opposite directions with their genitals coupled to each other. In many species there is a complicated mating game in which drum signals from the male can also play a role. The sperm of a mating is enough for several egg packets.
Only a few cockroach species reproduce thelytok, i.e. parthenogenetically . That is, they only have females in some or all of their populations . Facultative parthenogenesis has been demonstrated in various synanthropic "pest" species, such as Pycnoscelus surinamensis . Here the bisexual form is often regarded as a separate species and called Pycnoscelus indicus . Facultative parthenogenesis has also been demonstrated in Periplaneta species. In 2008 it was also detected in the Phyllodromica subaptera, which is native to Europe .

Egg packets (oothecae) ​​and development

The eggs are grouped in egg packets ( oothecae ). These are usually somewhat elongated and have a keel on one side, this forms a seam at which the hatched young animals hatch. Their shape and the number of eggs contained is species-specific and can be used to determine. The shell of the ootheca protects the eggs against enemies and water loss. However, it limits the number of eggs, which usually remains very low at around 20 to 50. The ootheca is thrown off by many species, sometimes also artfully camouflaged or buried, the latter especially by many representatives of the Blattinae and Ectobiinae. The oothecae of these species are usually hard and often protected against predators with calcium oxalate storage. The female of the common cockroach ( Blatta orientalis ) produces about 12 mm long, dark brown, smooth-walled egg capsules on which a seam is noticeable. It lays 16 eggs in each ootheca that it wears on its body for less than two days. However, many species carry their ootheca with them for a long time, sometimes until the nymphs hatch (e.g. the German cockroach ( Blattella germanica )). Some representatives turn the ootheca 90 degrees on its side. The females of the Blaberidae carry the ootheca inside the body after they have squeezed it out once, turned it and then pulled it in again. With them, finished young animals are finally born ( ovoviviparia ). A few species, such as Diploptera punctate , the eggs not only hold back, but feed them through a high-protein , milk-like substance in the uterus, which means they are viviparous ( viviparous ).
The development of the nymphs runs over a maximum of 13 stages, whereby the number within a species can vary depending on the living conditions. The females often grow a little larger and need more moults to reach sexual maturity. In exceptional cases, development can take several years. Species such as the synanthropic cockroaches can also develop significantly faster under favorable conditions, especially when there is sufficient heat, ideally 30 ° C.

Natural enemies

In addition to the predators common for insects of this size and way of life, there are also some that specialize in cockroaches or their oothecae. The representatives of the hymenoptera family hunger wasps (Evaniidae) are parasitoids that specialize in the oothecae of certain cockroach species . In Germany the species Bradygaster minuta is widespread and parasitizes in Ectobius oothecae. A notable enemy of the American cockroach ( Periplaneta americana ) is the jewel wasp . This injects a poison into the cockroach's brain, which suppresses its flight reflex . Then she builds the immobile animal in a brood cave and lays an egg on her body. The hatched wasp larva penetrates the cockroach and feeds on its internal organs until the cockroach dies. After pupation , the adult jewel wasp finally hatches from the cockroach's exoskeleton that has been eaten empty .

Systematics

Scientific naming

There is no complete agreement on the scientific name of the order. In addition to the name Blattodea, which goes back to a publication by Karl Brunner-von Wattenwyl in 1882, the name Blattaria, which goes back to Pierre André Latreille , 1810 , can also be found in current scientific publications . The latter is often written in older literature as in the original Blattariae. Occasionally, especially in the German-speaking area, the name Blattoptera appears, the taxonomic basis of which is puzzling. It probably goes back to Horst Bohn. The Dictyoptera , which are mostly understood as a superordinate order, are also regarded by some authors as an order, which then makes the cockroaches subordinate. The solution goes back to Hennig , to name the recent cockroaches Blattaria and to reserve the name Blattodea for the higher-level taxon, including the termites . However, Hennig was convinced of the monophyly of recent cockroaches. According to more recent findings, the Blattaria in this form would be an invalid ( paraphyletic ) taxon according to the rules of phylogenetic systematics .

External system

There is now broad consensus among researchers that cockroaches, termites, and fishing rodents form a common family group. This is mostly viewed as a superordinate order and called Dictyoptera. Hennig recognized this group as early as 1969 and named it Blattopteroidea. Morphologically, the group is z. B. characterized by a special structure of the tentorium . In addition, the eggs of this group are protected in an ootheca. For this reason, the cockroaches and the fishing fright were also summarized in the past in the taxon Oothecaria or together with the termites (and the ground lice ) as Oothecariformia.
Within the Dictyoptera, the ootheca is only reduced in the termites, but is also found here in one species: Mastotermes darwiniensis . Molecular pedigrees also regularly showed that the Dictyoptera belong together. The Dictyoptera themselves therefore belong to a kinship group of morphologically primitive insect orders, which are referred to as Polyneoptera . The exact relationship within the Polyneoptera is still uncertain and not stable between different studies.
While the existence of the Dictyoptera is hardly doubted, there has been a long controversy about the placement and position of the three classical orders within this group. The monophyly of termites and terrors is hardly disputed within research. However, the position of the cockroaches is problematic. Molecular pedigrees in particular have long indicated that cockroaches are paraphyletic with respect to termites. This means that the termites are probably nothing more than a special line of development within the cockroaches. They are therefore no longer regarded as order, but as a superfamily or family and as belonging to the cockroaches. Although there are opposing views, the majority of studies have identified the cockroach family Cryptocercidae as a sister group of termites. The sister group of this clade from Cryptocercus and termites is not known with certainty. Other researchers have set up other groupings.

This results in the following possible family tree:

 Dictyoptera  
  Blattodea  

 remaining cockroaches ("Blattaria")


  NN  

 Termites


   

 Cockroaches (genus Cryptocercus )




   

 Mantis (Mantodea)



Internal system

Skull cockroach
( Blaberus craniifer )
subfamily Blaberinae
German cockroach
( Blattella germanica )
subfamily Blattellinae
Common wood cockroach
( Ectobius lapponicus )
subfamily Ectobiinae
Amber cockroach
( Ectobius vittiventris )
subfamily Ectobiinae
Common cockroach ( Blatta orientalis )
subfamily Blattinae
Australian cockroach ( Macropanesthia rhinoceros )

In the classic system, the cockroaches are divided into a number of families within at least three superfamilies. In addition to the subfamilies listed here, most of these families also contain a considerable number of genera that are currently not classified in any of these subfamilies. The following overview shows the division down to the subfamily level (as of January 2012). In addition, some of the more well-known species are classified or shown in the picture:

Fossil evidence

Exuvia of a Blattodea in Baltic amber (image width approx. 6.5 mm)

The oldest known fossils of this insect order come from the Carboniferous . The finds from the Upper Carboniferous and the Permian testify to a great variety of forms, which points to a dominance of this order in the late Paleozoic . Although some taxons are controversial, several hundred species can be assumed to have been found from this period. Some of these species are suitable as guide fossils. The Paleozoic forms had already reached a stage of development that did not differ significantly from that of the more recent representatives of this order. In addition to the finds in sediments of almost all geological periods since the Carboniferous, fossil evidence from various Cretaceous and Tertiary amber deposits is known. In the Eocene / Oligocene Baltic amber forty species were identified about eight families. Since large adult animals were usually strong enough to break free from the resin, juvenile specimens are represented above average in amber.

Origin of name

The name “cockroaches” was transferred from the pest species to the rest of the order. The name cockroaches for the Blattodea can be traced back to the end of the 18th century. Before, and sometimes even later, all possible material pests were grouped under cockroaches. Primarily the caterpillar of the clothes moth, which belongs to the butterflies, was meant, only later the name was also transferred to the Blattodea. The reason for the naming was probably in all cases the scraping diet and the resulting damage.

Web links

Wiktionary: cockroach  - explanations of meanings, word origins, synonyms, translations
Commons : Cockroaches  - Collection of images, videos and audio files

swell

  1. ^ A b c George William Beccaloni: Blattodea Species File Online . Version 1.0 / 4.0. World Wide Web electronic publication. http://Blattodea.SpeciesFile.org (accessed January 15, 2012)
  2. ^ William J. Bell, Louis Marcus Roth, Christine A. Nalepa: Cockroaches: ecology, behavior, and natural history . JHU Press, 2007. ISBN 0-8018-8616-3
  3. ↑ Horst Bohn, Mike Picker, Klaus-Dieter Klass, Jonathan Colville: A Jumping Cockroach from South Africa, Saltoblattella montistabularis, gen. Nov., Spec. nov. (Blattodea: Blattellidae) . Arthropod Systematics & Phylogeny, 2009, 68 (1): 53-69.
  4. ^ H. Weidner: Lapland cockroaches as potential leisure pests. Anzeiger für Schädlingskunde, 45 (5), 1972: pp. 75-76. doi : 10.1007 / BF01886032
  5. ↑ Hannes Baur, Isabelle Landau Lüscher, Gabi Müller, Marcus Schmidt, Armin Coray: Taxonomy of the amber wood cockroach Ectobius vittiventris ( A. Costa , 1847) (Blattodea: Blattellidae) and its distribution in Switzerland . Revue Suisse de Zoologie, 111 (2), 2004: pp. 395–424.
  6. ↑ Thomas Knebelsberger: Geographical parthenogenesis in the cockroach Phyllodromica subaptera (Blattoptera, Blattellidae, Ectobiinae) and revision of the subaptera species complex . Diss., Univ. Munich, 2008.
  7. ↑ Anna Williford, Barbara Stay, Debashish Bhattacharya: Evolution of a novel function: nutritive milk in the viviparous cockroach, Diploptera punctata . Evolution and Development 6 (2), 2004: pp. 67-77.
  8. ↑ Carl Brunner von Wattenwyl: Prodromus of the European Orthopteren. Verlag Wilhelm Engelmann (Leipzig) 1882. Full text online
  9. ^ PA Latreille: Considérations générales sur l'ordre naturel des animaux composant les classes des Crustacés, des Arachnides, et des Insectes avec un tableau méthodique de leur genres disposés en familles. Pans, Schoell, 1810; 444 pp.
  10. ↑ a b Willi Hennig : The tribal history of the insects . Kramer Verlag, 1969.
  11. ↑ Klaus-Dieter Klass : The tentorium and anterior head sulci in Dictyoptera and Mantophasmatodea (Insecta) . Zoologischer Anzeiger Volume 246, Issue, 2007, 3: pp. 205-234. doi : 10.1016 / j.jcz.2007.06.001
  12. ↑ K. Günther, H.-J. Hannemann, F. Hieke, E. Königsmann & H. Schuman: Urania animal kingdom - insects. Urania-Verlag, Leipzig, Jena, Berlin 1994, ISBN 3-332-00498-0
  13. ↑ Oliver Zompro : Mantophasmatodea - Gladiators in the insect kingdom. , Arthropoda 16 (1) March 2008, Sungaya-Verlag Kiel. Pp. 24-25 ISSN  0943-7274
  14. ^ Ward C. Wheeler, Michael Whiting, Quentin D. Wheeler, James M. Carpenter: The Phylogeny of the Extant Hexapod Orders . Cladistics 17, 2001: pp. 113-169. doi : 10.1006 / clad.2000.0147
  15. ↑ Daegan Inward, George William Beccaloni, Paul Eggleton: Death of an order: a comprehensive molecular phylogenetic study confirms that termites are eusocial cockroaches . Biology Letters 3, 2007: pp. 331-335. doi : 10.1098 / rsbl.2007.0102
  16. ↑ Nathan Lo, Michael S Engel, Stephen Cameron, Christine A Nalepa, Gaku Tokuda, David Grimaldi, Osamu Kitade, Kumar Krishna, Klaus-Dieter Klass, Kiyoto Maekawa, Toru Miura and Graham J. Thompson: Save Isoptera: A comment on Inward et al . Biol. Letters 3, 2007: pp. 562-563. doi : 10.1098 / rsbl.2007.0264
  17. ↑ Steffen Roth, Bastian Fromm, Gerd Gäde, Reinhard Predel: A proteomic approach for studying insect phylogeny: CAPA peptides of ancient insect taxa (Dictyoptera, Blattoptera) as a test case . BMC Evolutionary Biology 9, 2008: p. 50 doi : 10.1186 / 1471-2148-9-50 open access
  18. ↑ LL Deitz, C. Nalepa, Klaus-Dieter Klass: Phylogeny of the Dictyoptera re-examined (Insecta) . Entomologische Abhandlungen (Dresden) 61 (1), 2003: pp. 69–91.
  19. ↑ Barbara L. Thorne & James M. Carpenter: Phylogeny of the Dictyoptera. Systematic Entomology 17, 1992: pp. 253-268. doi : 10.1111 / j.1365-3113.1992.tb00336.x
  20. ^ Müller: Textbook of palaeontology. Volume II, part 3, Jena 1978.
  21. ↑ C. Gröhn: Inclusions in Baltic amber. Kiel / Hamburg 2015. ISBN 978-3-529-05457-0 .
  22. ↑ Heinrich Kemper: The animal pests in language use. Duncker & Humblot, 1959. Chap. Motte und Schabe, p. 66 ff. Limited preview on Google Books