Zeugma | |
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Species of Zeugma from Thailand | |
Scientific classification ![]() | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Arthropoda |
Class: | Insecta |
Order: | Hemiptera |
Suborder: | Auchenorrhyncha |
Infraorder: | Fulgoromorpha |
Family: | Derbidae |
Subfamily: | Otiocerinae |
Tribe: | Zoraidini |
Genus: | Zeugma Westwood, 1840 [1] |
Zeugma is a small genus of planthoppers from the family Derbidae, tribe Zoraidini, currently (2024) with 9 species. [2] The distribution of the species is restricted to western and central parts of Indonesia, as well as to tropical parts of southern Asia, mainly in and around Malaysia, southern Thailand and southern Cambodia. [3] [4] [5] However, some species are also found in the Philippines, Taiwan and in Assam (India). [6] [7] [8] The species of the genus Zeugma differ from the typical genera in the tribe Zoraidini by a number of unusual features. They lack the long and narrow forewings and the very short hindwings. In addition, the media vein on the forewing has more than 6 branches. These characters are found among genera in the tribe Derbini, to where Zeugma was assigned before the 1950s. [9] The genus was transferred to the Zoraidini by Fennah, mainly based on the venation of the forewings, specifically that of the cubitus and the clavus. [10] Further, when at rest, the live insects do not raise their wings above the body like in most other species of the tribe Zoraidini, but hold them in a roof-like position. [5] Because of these atypical features, the genus Zeugma is placed in a separate subtribe. [2] The colouration of the body is typically straw-coloured to light brown with some parts dark brown, reddish, black or sometimes yellowish. The forewings are translucent, often with brown stripes, especially on or along the veins. [6] [4] [5]
Species of the genus Zeugma are found in central and western parts of Indonesia, in Malaysia, Thailand, Cambodia, the Philippines, Taiwan and in Assam (India). The description of the type species of Zeugma, Zeugma vittata, was based on a specimen of unknown origin. [11] In 1913, Muir described Zeugma monticola which he later regarded as a synonym name for Zeugma vittata, giving Parit Buntar (peninsular Malaysia), Mowong (Borneo) and Pekalongan (Central Java) as localities. [3] Four other species of Zeugma have been described by Muir from Indonesia, Zeugma elegans and Zeugma karnyi from the Mentawai Islands, off West Sumatra, Zeugma corporaali from Medan in northern Sumatra, and Zeugma javana from Java. [12] [13] [4] In addition, Zeugma fuscinervis has been described from Assam (India), Zeugma valdezi from the Philippines, as well as two species from Taiwan (Zeugma maesta and Zeugma makii). [6] [7] [8] Several unidentified species of Zeugma have been reported from southern Thailand, southern Cambodia and from Sulawesi Island in Indonesia. [5]
Typically, species in the tribe Zoraidini have forewings which are at least 3½ times longer than wide, hind wings which are less than half as long as the forewings and a media vein on the forewings which has 5 or 6 branches. [10] In the genus Zeugma the above features do not apply. The forewings are broader, only about 2½ times longer than wide, the hind wings are longer than half the forewing length, usually around 60% of the forewing length, and the media vein on the forewing has 8 or more branches, usually with none of them branching further into sub branches. Such features are found in the tribe Derbini and Zeugma has been placed into that tribe before the 1950s. [9] However, Zeugma was transferred to the tribe Zoraidini by Fennah after redefining the tribe Zoraidini. This transfer was mainly based on the venation of the forewings, specifically that of the clavus and the cubitus, as well as on the venation of the hind wings. Like in other genera of Zoraidini, the forewings of Zeugma have an open clavus where the two veins of the clavus join, but the combined vein does not reach the hind margin, instead it meets the veins of the cubitus. [10]
When live Zeugma species are at rest, they carry their wings in a roof-like position covering their abdomen. [5] This is also in contrast to most other species of the tribe Zoraidini which raise their wings above the body. [14] Both wing positions are not found in species of the tribe Derbini, which spread out their wings laterally when at rest, holding them parallel to the surface they sit on. [10] [15] Due to the various differences to other genera in the tribe Zoraidini, the genus Zeugma has been placed in a separate subtribe, the Zeugmatina. [16]
Within the tribe Zoraidini, the species of Zeugma are of medium size, the body is around 5 mm long from the tip of the head to the end of the abdomen. The forewings are usually around 9 mm long. [3] [6] Zeugma elegans appears to be the larges species with a body length of 6 mm and a forewing length of 13 mm. [4] The head is broad, as broad as the thorax, another unusual feature in the tribe Zoraidini. The dorsal part of the head between the eyes (the vertex) and the face are also broad. The antennae are elongated, but shorter than the face. [10] The typical colour of Zeugma species is light brown or stramineous (straw-coloured), often with dark brown marks or bands. For example, the forewings or the mesonotum might have dark brown bands. The forewing veins are also often dark brown or are lined with dark brown stripes. Further, the body and legs can have additional marks which may be black, red or yellowish. [3] [4] [5]
There is no information on the biology of the immature stages of Zeugma species. However, it can be assumed that they live in decaying organic matter like other derbids, feeding on fungi. [3] [17] Although there are few host plant records for the adult hoppers in the genus Zeugma, the species seem to feed mainly on palm trees and other monocots like other species in the tribe Zoraidini. For example, Zeugma javana was described from specimens collected from palm trees in Java and Zeugma valdezi has been frequently recorded as feeding on coconut palms in the Philippines. [12] [18]
The following list summarizes the distribution and colouration of the 9 species of Zeugma:
Zeugma corporaali Muir, 1923 [13]
Indonesia, Sumatra, Medan. Similar in size and colouration to Zeugma vittata.
Zeugma elegans Muir, 1926 [4]
Described from Indonesia, Mentawai Islands (Se Pora, off West Sumatra), also reported from Singapore and mainland Malaysia. [19] Body light brown, a dark brown mark down the frons and clypeus, a broad dark mark down middle of mesonotum; abdomen dark brown, forewings light brown with dark veins.
Zeugma fuscinervis Muir, 1922 [7]
In Assam (India). Head and pronotum light brown, darker over vertex and down the middle of pronotum; mesonotum and tegulae dark brown or black; abdomen dark brown; legs light brown; forewings translucent, slightly yellowish, veins dark spreading into membrane.
Zeugma javana Muir, 1915 [12]
In Java, Indonesia. Body straw-coloured or light brown; dark brown or black between the keels of the vertex, the face and the clypeus; forewings straw-coloured, veins yellow with brown borders.
Zeugma karnyi Muir, 1926 [4]
In Indonesia, Mentawai Islands, off West Sumatra. Body straw-coloured, some parts red, clypeus black, middle of pronotum and mesonotum dark brown; forewings translucent and straw-coloured, veins yellow or reddish.
Zeugma maesta Yang & Wu, 1993 [8]
In Taiwan. General colour pale yellow; red in a rather broad band on pro- and mesonotum, on the vertex and on the ridges of face and postclypeus, also on the upper portion of the tegula; forewings pale yellow with black spots and lines.
Zeugma makii Muir, 1914 [20]
In Taiwan. Body straw-coloured, vertex, face, clypeus, middle of pronotum and scutellum red; front legs brown; forewings translucent, pale dirty yellow with lighter veins, media vein and bases of median branches as well as cross-veins brown.
Zeugma valdezi Muir, 1917 [6]
In the Philippines. Body brown or reddish brown, a dark longitudinal mark passing over vertex face and clypeus, five dark marks on mesonotum, abdomen darker; forewings very light brown or reddish yellow, darker along veins, veins reddish brown.
Zeugma vittata Westwood, 1840 TYPE SPECIES [1]
Synonym: Zeugma monticola Muir 1913 [3]
In peninsular Malaysia, Singapore, Borneo, Java and Bali. [3] [5] Head, pronotum, sterna and legs orange-yellow, some parts blackish or brown; forewings translucent, tinted with grey, a brown line covering cross veins of cubitus and media branches, another over the media vein and a third through the subcostal cell to the wing tip, a black line over apical cross veins.
The family Derbidae is a large and diverse group of planthoppers, widely distributed in the tropics and also found in subtropical and temperate regions. It includes more than 150 genera and more than 1500 species. The adults suck the sap of plants while the nymphs live mainly in decaying organic matter, for example under the bark of dead tree trunks, feeding on fungi. In some groups of species the general structure of the insects is largely uniform, whereas in others like in the subfamily Otiocerinae various forms can be found. For example, the shape of the head, the antennae or the wings can differ considerably among genera and species.
Otiocerinae is a subfamily of planthoppers in the family Derbidae with more than 100 genera and more than 1000 species. The species are widely distributed, especially in the Americas, Sub-Saharan Africa, as well as parts of Asia, Australia and the Pacific. Like in other Derbidae, the adults suck the sap of plants while the nymphs live mainly in decaying organic matter, for example under the bark of dead tree trunks, feeding on fungi.
Proutista moesta is a common species of planthopper from the family Derbidae, tribe Zoraidini. It is found in tropical parts of Asia from India to southern parts of China, some islands in the western Pacific, as well as in the Indian Ocean. There is also a record from eastern Africa, a possible recent introduction. P. moesta can be common on crops like oil palms, coconut palms, sugarcane and several cereals, where it sucks the sap from the plants. It is often regarded as a pest of these crops, a vector of phytoplasma diseases and a biosecurity risk in countries outside its distribution range. Males of P. moesta are smaller than females when measured from the tip of the head to the end of the forewings. The forewings alone have a length of around 6.1 mm for males or around 6.6 mm for females. The colour is predominantly black, but the legs, antennae, facial ridges and the rostrum are yellowish to light orange. The ridges on the mesonotum and the clypeus are whitish to light yellow and the body has scattered white dots. The black forewings have some clear patches, mainly along the costal margin and in the distal half between the hind margin and the media vein. When at rest, the insect raises its wings above the body and spreads them out at an angle of about 60°.
The Rhotanini is a group (tribe) of planthoppers in the family Derbidae. They are found mainly in countries of south-eastern Asia like Malaysia, the Philippines, Indonesia and Papua New Guinea. They are characterized by their small size, their broad forewings, the vein pattern on their wings, and a simple head structure. Around 300 species have been described. Like in other derbids, the adults suck the sap of plants while the nymphs live mainly in decaying organic matter.
Levu is a genus of small planthoppers from the family Derbidae, tribe Rhotanini, with more than 30 species. The species are found mainly on the islands of the western Pacific, particularly Melanesia. A few species have been also reported from eastern Australia and mainland Asia. The adult hoppers are around 4–5 mm long from head to the tip of the forewings. They lack the bright colours or conspicuous wing patters of other derbids, but a few species have red or orange markings. The wings may be carried in a roof-like (tectiform) position when the insects are at rest or they form a common plane like in house flies.
Rhotana is a large genus of planthoppers from the family Derbidae, tribe Rhotanini, with more than 70 species. Most species have been reported from the rainforests of Borneo, Indonesia and New Guinea, but the distribution of the genus extends over a much wider area in southern and south-eastern Asia, as far north as southern Japan and including parts of the Pacific islands as well as Australia. The species are the largest in the tribe Rhotanini, typically between 5–6 mm from head to the tip of the wings. The forewings frequently have striking patterns and colours, some species have also marks on the hind wings.
Saccharodite is a large genus of planthoppers from the family Derbidae, tribe Rhotanini, with more than 100 species. The largest number of species have been reported from Borneo, Indonesia, New Guinea and the Philippines, but the genus has a much wider distribution, ranging from Sri Lanka in the West, over southern parts of mainland Asia up to southern Japan in the North. Saccharodite is also found in other parts of the western Pacific like Micronesia, Fiji and New Caledonia, up to eastern parts of Australia, in the South as far as Tasmania. The adult hoppers are around 5 mm long from head to the tip of the forewings. They are characterized by their forewing venation. The forewings are usually glassy and bright red marks are common on the head and body, up to most parts except the wings being coloured bright red. When the insects are at rest, the wing surfaces form a common plane like in house flies.
Alara is a genus of small planthoppers from the family Derbidae, tribe Rhotanini, with more than 30 species. The species are found mainly in New Guinea, other parts of Indonesia, Borneo and the Philippines. However, some species have been also reported from Sri Lanka, parts of mainland Asia, Taiwan and Australia. The adult hoppers are around 4 mm long from head to the end of the wings. The forewings are nearly oval in outline with 5 broad costal cells that gradually decrease in length from the base to the tip of the wings. The antennae and the rostrum are longer than in other genera of Rhotanini. The forewings are often dark brown and lack any bright colours. However, there may be red marks on the head. When the insects are at rest, the wing surfaces form a common plane like in house flies.
Dichotropis is a small genus of planthoppers from the family Derbidae, tribe Rhotanini, with 9 species. The species are found on the island of New Guinea, including the Indonesian part and the neighbouring islands of New Britain and New Ireland. The distribution range of several species extends to other countries, as far as West Malaysia, the Philippines, the Solomon Islands and northern Australia. The adult hoppers are around 5–6 mm long from head to the tip of the wings. The species can be recognized by their white and powdered forewings which are carried in a roof-like position when the insects are at rest. The ridges on the sides of the face are separated and do not meet in front of the eyes. The body is typically straw-coloured or light brown. The ridges on the head may be orange or the whole head and body may be orange, but bright red marks are missing.
Rhotanella is a small genus of planthoppers from the family Derbidae, tribe Rhotanini, with 8 species. The species are found in the Philippines, Indonesia, New Guinea, the Solomon Islands and southern China. The adult hoppers are between 4–6 mm long from head to the tip of the wings. They can be recognized by the shape and venation of their forewings which have a broad base and 5 very wide costal cells. Especially the first (basal) costal cell is very broad and has a width of about one quarter to one third of the width of the whole wing. The ridges on the sides of the face meet in front of the eyes. The profile of the head is also unusual for the tribe Rhotanini and has either an approximately rectangular or a somewhat triangular outline. The colour of the body is straw-coloured to light brown. The legs may be partly red or have red stripes. The forewings are brownish, but may have red marks in the costal cells. The forewings are carried in a roof-like position when the insects are at rest.
Sumangala is a genus of planthoppers from the family Derbidae, tribe Rhotanini. The 13 species are found in southern Asia from southern India and Sri Lanka in the West over parts of mainland Asia to south-east Asian islands like Taiwan and the Philippines and including parts of Indonesia. The adult hoppers are around 5 mm long from head to the tip of the wings. They can be recognized by their forewing venation where the branch of the cubitus joins the first media branch to form the basal median cell, unlike in other genera of Rhotanini where the basal median cell is formed by a crossvein between the first media branch and the cubitus branch or by the first branch of the media splitting again close to its base and the inner sub branch joining the cubitus branch. In Sumangala, the first media branch does not split and as a consequence only 6 veins of the cubitus-media complex reach the margin at the end of the wing. The forewings of Sumangala are mainly whitish and most of them lack larger black marks.
Muiralevu is a genus of planthoppers from the family Derbidae, tribe Rhotanini, with 19 species. While the species of all other genera in the tribe Rhotanini are only found in southern Asia and the Pacific, the species of Muiralevu also occur in western Africa. The distribution of the type species covers several West African countries, ranging from Sierra Leone to the Central African Republic. Four other, very similar species are also found in western Africa, while 14 other species occur in south-eastern Asia. The species of Muiralevu can be recognized by their forewing venation, especially the 4 costal cells and the venation around the basal median cell. Apart from the basal median cell, there is also a characteristic trapezoid cell at the base of the first media branch. On average, the forewings are more elongated compared to most other species of Rhotanini. The adult hoppers are between 4 and 6 mm long from head to the tip of the wings. The head is evenly rounded and the ridges on the sides of the face meet in front of the eyes. The body is usually straw-coloured to light brown. The forewings are mostly whitish and powdered, but in a few species they have large black marks.
The Zoraidini is a group (tribe) of planthoppers in the family Derbidae. They are found in tropical and subtropical parts of Africa, Asia, Australia and in parts of the western Pacific. Almost 300 species in 27 genera are recognized. They are characterized by their long and narrow forewings and the short hind wings, usually not longer than half the length of the forewings and often even shorter. The forewings are typically around 8–10 mm long and the body is usually not more than half that length. In most species, the wings are raised when the insects are at rest, and are often spread out at an angle of about 45–90°. The head is compressed and the face is narrow in frontal view. The antennae are often long and the eyes cover about half the face. Like in other derbids, the adults suck the sap of plants while the nymphs live mainly in decaying organic matter. A few species are regarded as minor agricultural pests.
Zoraida is a large genus of planthoppers from the family Derbidae, tribe Zoraidini, with more than 100 species. These are widely distributed in the Old World tropics and in some subtropical parts of eastern Asia. In Africa, they are found mainly in the humid tropics of West Africa like Sierra Leone, the Ivory Coast, the Central African Republic, Nigeria or the Democratic Republic of Congo. However, Zoraida species have been also described from eastern Africa as far north as South Sudan and as far south as parts of South Africa. In Asia, the largest number of species have been described/reported from Indonesia, the Philippines, Taiwan and Sri Lanka, but species are also known as far south as northern Australia and as far north as the southern tip of far-eastern Russia. Like other species of the tribe Zoraidini, Zoraida species have long and narrow forewings and short hind wings. They can be identifies by the forewing venation and the structure of the head.
Proutista is a small genus of planthoppers from the family Derbidae, tribe Zoraidini, with 10 species, as of 2024. The type species, Proutista moesta is widely distributed and often very common, its distribution ranging from Tanzania in Africa, over some Indian Ocean islands and southern parts of western Asia to far eastern Asia and parts of the western Pacific. Seven of the remaining species are also found in tropical parts of Asia and only two additional species have been recorded from Africa, where Proutista fritillaris is the most common one. The species of Proutista are characterized by a combination of different features, mainly the shape and venation of the forewings, the size of the hind wings and the structure of the head and the antennae. On the forewings the media vein has 6 branches, none of them branching further into sub branches. The hind wings are about half as long as the forewings and have a rounded tip. The head has a narrow face (frons) and the antennae are rather short, much shorter than the face.
Diostrombus is a large genus of planthoppers from the family Derbidae, tribe Zoraidini, with more than 40 species. The large majority of species have been described from Africa and the Near East. However, a few species, including the type species, are found in southern and eastern parts of Asia. In Africa, the distribution range of Diostrombus species extends from the humid tropics of western Africa to many parts of eastern Africa, as far north as parts of Sudan and as far south as South Africa. Three species have been described from Yemen and the Iran. Like other species of the tribe Zoraidini, species of Diostrombus have long and narrow forewings and short hind wings. The main characteristic of Diostrombus is the forewing venation with the media vein having 6 branches which do not split further into sub branches. The hind wings are much shorter than half the forewing length and have a pointed tip. In profile the head is usually flat, but in some African species, the ridges on the sides of the face are extended in front of the antennae. The face is narrow. The antennae are short compared to other genera of the tribe Zoraidini, about half as long as the face.
Pamendanga is a genus of planthoppers from the family Derbidae, tribe Zoraidini, with 37 species, as of 2024. About two-thirds of the species, including the type species, are found in tropical and some subtropical parts of Asia, from Sri Lanka in the West to parts of Japan in the North-East and including the Philippines, Indonesia and New Guinea. The remaining species have been described from Africa, mainly from the humid tropics of West Africa, ranging from Sierra Leone and the Ivory Coast in the North to the Democratic Republic of Congo in the South. Some species are also known from eastern Africa. Like other species of the tribe Zoraidini, Pamendanga species have long and narrow forewings and short hind wings. They can be recognized by the forewing venation and the shape of the head and the antennae. On the forewing, the media vein has 6 branches and the 3rd branch forks further into 2 sub branches. The head has a flat profile and a narrow face with cylindrical antennae which are elongated but not longer than the face.
Lydda is a genus of planthoppers from the family Derbidae, tribe Zoraidini, with 21 species, as of 2024. The known distribution range of its species is restricted to parts of Queensland and the Northern Territory in Australia, to the island of New Guinea and to the island of Larat in south-eastern Indonesia. The type species, Lydda elongata, is mainly found in parts of Queensland and in New Guinea. In the past, the genus was often confused with the genus Proutista and a number of Lydda species have been originally described under Proutista. While species of Lydda and Proutista share a similar club-shaped outline of their forewings and a similar wing venation, there are significant differences between both genera in the forewing venation and the structure of the aedeagus in males. On the forewings, the media vein has only 5 branches, while Proutista species have 6 branches. In the type species, Lydda elongata, the eyes are unusually large and the head is very broad. The hind wings are about half as long as the forewings. Some species of Lydda have elaborate patterns on their body and forewings. For example, in live specimens of Lydda elongata, the body has a mixture of reddish brown and bluish patterns, while the forewings are brownish with darker spots. Species that have been previously described under the genus Lydda in Africa are now assigned to the genus Lyddastrombus.
Lyddastrombus is a small genus of planthoppers from the family Derbidae, tribe Zoraidini, currently (2024) with 8 African species. The morphology of the species is similar to those in the genus Lydda and most species have been originally described as part of Lydda. Lyddastrombus species can be recognized by the venation of their forewings and other features. The genus also shows similarities to the genera Diostrombus and Proutista, sharing with them the short antennae and a related forewing venation, where the media vein has 6 branches with none of them forking further into sub branches. However, Lyddastrombus species differ from both genera by their much broader head. Lyddastrombus was original described as a subgenus of Diostrombus, but it is now regarded as a separate genus, due to the broader head. The species of Lyddastrombus are found mainly in the tropical rainforests of West Africa from Sierra Leone over Nigeria to the Democratic Republic of the Congo, but have been also reported from parts of eastern Africa. Lyddastrombus hoppers have been frequently reported feeding on coconut and oil palms. In Mozambique, they were found to be carriers of a phytoplasma disease of coconut palms.
Helcita is a small genus of planthoppers from the family Derbidae, tribe Zoraidini, currently (2024) with only 4 species. All of the species are uncommon. They have rather restricted and widely separated distribution ranges. Two species are found in tropical parts of Africa, one in Sri Lanka and southern India, and one in Taiwan and some neighbouring islands of Japan. The species of Helcita can be distinguished from other similar genera by a combination of features related to the venation of the forewings and the shape of the head. On the forewing, the media vein has 6 branches with the 3rd branch forking further into 2 sub branches. This venation is similar to that of the genus Pamendanga from which Helcita species can be distinguished by the profile of the head. In Helcita species, the head extends conically in front of the eyes, whereas Pamendanga species have a flat profile.