Caccothryptus Ripicola
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''Caccothryptus'' is a
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of minute marsh-loving beetles in the subfamily Limnichinae. The genus was formally established (
circumscribed In geometry, a circumscribed circle for a set of points is a circle passing through each of them. Such a circle is said to ''circumscribe'' the points or a polygon formed from them; such a polygon is said to be ''inscribed'' in the circle. * Circum ...
) by the entomologist David Sharp in 1902, with '' C. compactus'' as the
type species In International_Code_of_Zoological_Nomenclature, zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the spe ...
. In 2014, the entomologists Carles Hernando and Ignacio Ribera published a major revision of the genus, reorganizing it from seven species into twenty species in five species groups, classified mainly by similarities in genital shape. Discoveries in the following years have brought the total number of species up to thirty-four. Due to the relative lack of studies and material on Asian Limnichidae, more species are likely still to be described. Ranging from 2.5 to 5.5 mm in total length, the beetles have ovoid bodies ranging from black to brown in color. Their range extends across
Southeast Asia Southeast Asia is the geographical United Nations geoscheme for Asia#South-eastern Asia, southeastern region of Asia, consisting of the regions that are situated south of China, east of the Indian subcontinent, and northwest of the Mainland Au ...
, alongside portions of
East Asia East Asia is a geocultural region of Asia. It includes China, Japan, Mongolia, North Korea, South Korea, and Taiwan, plus two special administrative regions of China, Hong Kong and Macau. The economies of Economy of China, China, Economy of Ja ...
and
South Asia South Asia is the southern Subregion#Asia, subregion of Asia that is defined in both geographical and Ethnicity, ethnic-Culture, cultural terms. South Asia, with a population of 2.04 billion, contains a quarter (25%) of the world's populatio ...
. They generally live in water-logged dead wood adjacent to small streams.


Taxonomy

''Caccothryptus'' is a genus of Limnichinae, a subfamily of the minute marsh-loving beetles (Limnichidae). The Limnichidae are a diverse family of beetle, absent only from alpine and polar regions, as well as most oceanic islands. They are most commonly
riparian A riparian zone or riparian area is the interface between land and a river or stream. In some regions, the terms riparian woodland, riparian forest, riparian buffer zone, riparian corridor, and riparian strip are used to characterize a ripar ...
, living on the shores of rivers, although some are
intertidal The intertidal zone or foreshore is the area above water level at low tide and underwater at high tide; in other words, it is the part of the littoral zone within the tidal range. This area can include several types of habitats with various sp ...
(living along beaches or coral formations). However, many genera are fully terrestrial, living in forest litter. Little research has been done on most genera, and the
larvae A larva (; : larvae ) is a distinct juvenile form many animals undergo before metamorphosis into their next life stage. Animals with indirect developmental biology, development such as insects, some arachnids, amphibians, or cnidarians typical ...
and
pupae A pupa (; : pupae) is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in their life cycle, the stages th ...
of most are especially poorly documented. Most species are
herbivorous A herbivore is an animal anatomically and physiologically evolved to feed on plants, especially upon vascular tissues such as foliage, fruits or seeds, as the main component of its diet. These more broadly also encompass animals that eat n ...
. Within Limnichinae, ''Caccothryptus'' belongs to the "''Mandersia'' group" of genera, alongside '' Euthryptus'', '' Mandersia'', ''
Pseudothryptus ''Pseudothryptus multiseriatus'' is the only species of ''Pseudothryptus'', a genus of minute marsh-loving beetle in the subfamily Limnichinae. Initially described in 1923 as part of the large genus ''Caccothryptus'', it was redescribed in 2004 ...
'', '' Resachus'', and '' Simplocarina'', all much smaller taxa in terms of species count. These genera share a number of physical features, such as a non-articulated (jointed)
aedeagus An aedeagus ( or aedeagi) is a reproductive organ of male arthropods through which they secrete sperm from the testes during copulation (zoology), copulation with a female. It can be thought of as the insect equivalent of a mammal's penis, th ...
(the male reproductive organ). These lack an articulated spiculum (internal piece) within the median lobe; this distinguishes them from the genus '' Byrrhinus''. The
Afrotropical The Afrotropical realm is one of the Earth's eight biogeographic realms. It includes Sub-Saharan Africa, the southern Arabian Peninsula, the island of Madagascar, and the islands of the western Indian Ocean. It was formerly known as the Ethiopi ...
genera '' Tricholimnichus'' and '' Cyclolimnichus'' are close relatives of the ''Mandersia'' group, but lack some distinguishing characteristics in 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 e ...
(dorsal plate of the thorax).


Taxonomic history

In 1902, the entomologist David Sharp described '' Caccothryptus compactus'' (found in Martapura,
Borneo Borneo () is the List of islands by area, third-largest island in the world, with an area of , and population of 23,053,723 (2020 national censuses). Situated at the geographic centre of Maritime Southeast Asia, it is one of the Greater Sunda ...
) in an article in the ''
Entomologist's Monthly Magazine ''Entomologist's Monthly Magazine'' is a British entomological journal, founded by a staff of five editors – T. Blackburn, H. G. Knaggs, M.D., R. McLachlan, F.L.S., E. C. Rye and H. T. Stainton – and first published in 1864. The journal ...
''. He dubbed the genus as the "largest and most remarkable of the Limnichini" and noted its close resemblance to the Central American genus ''Euthryptus''. ''Caccothryptus'' was the first
circumscribed In geometry, a circumscribed circle for a set of points is a circle passing through each of them. Such a circle is said to ''circumscribe'' the points or a polygon formed from them; such a polygon is said to be ''inscribed'' in the circle. * Circum ...
Limnichinae genus native to Asia. Sharp did not list an etymology for the name ''Caccothryptus'' in his description of ''C. compactus''. In 1922 and 1923,
Maurice Pic Maurice Pic (23 March 1866, in Marrigny near Digoin – 29 December 1957, in Les Guerreaux) was a French entomologist who specialised in Coleoptera. He contributed to Mary-Louis Fauconnet's ''Catalogue raisonné des coléoptères de Saône-et- ...
circumscribed a new genus he termed ''Macrobyrrhinus'', and placed in it four species of Limnichinae, with '' M. rouyeri'' as the
type species In International_Code_of_Zoological_Nomenclature, zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the spe ...
. Harry George Champion treated this genus as a
synonym A synonym is a word, morpheme, or phrase that means precisely or nearly the same as another word, morpheme, or phrase in a given language. For example, in the English language, the words ''begin'', ''start'', ''commence'', and ''initiate'' are a ...
of ''Caccothryptus'', in a paper describing three additional species. Pic disregarded Champion's synonymy and placed more species in ''Macrobyrrhinus'' in 1928; however, it is now accepted as a synonym. ''Caccothryptus'' received no further mention in entomological literature until 1998, when it was listed in a checklist of East Asian Limnichidae species. In 2005, the entomologists Carles Hernando and Ignacio Ribera reclassified one species described by Champion, ''C. multiseriatus'', into a new genus dubbed ''
Pseudothryptus ''Pseudothryptus multiseriatus'' is the only species of ''Pseudothryptus'', a genus of minute marsh-loving beetle in the subfamily Limnichinae. Initially described in 1923 as part of the large genus ''Caccothryptus'', it was redescribed in 2004 ...
''. Nine years later, Hernando and Ribera published a major revision of the genus. Previously grouped as seven species, it was reorganized into twenty species in five species groups—''compactus'', ''rouyeri'', ''testudo'', ''jaechi'', and ''zetteli''—mainly classified based on similarities in genital shape, and to a lesser extant certain external features. This 2014 revision also classified one of Champion's species, ''C. laosensis,'' as a synonym of ''C. maculosus''.


Species and species groups

All ''Caccothryptus'' species have a non-articulated aedeagus, while their
parameres Parameres ('side parts') are part of the external reproductive organs of male insects and the term was first used by Verhoeff in 1893 for the lateral genital lobes in Coleoptera. The primary phallic lobes which appear in the nymph or larval stag ...
(side lobes of the external genitalia) can be either separated or fused. This can vary even within a species group, although all species of ''compactus'' and ''testudo'' have separated parameres. A lobe along the center of the aedeagus has a sulcus (groove), which varies between species groups. ''C. rouyeri'', the only species of the ''rouyeri'' group, can be distinguished by lateral expansions on the median lobe of its aedeagus, while the ''jaechi'' group can be recognized by a denticled (serrated) ventral portion of this lobe. All species have microscopic glandular pores on the middle portion of the fourth
sternite The sternum (: sterna) is the ventral portion of a segment of an arthropod thorax or abdomen. In insects, the sterna are usually single, large sclerites, and external. However, they can sometimes be divided in two or more, in which case the su ...
(one of the rear segments of the abdomen), while the ''jaechi'' and ''zetteli'' groups also have these on the sides of the fifth sternite. Males of ''compactus'' group species have dissimilar front claws, with the internal claw longer and more indented than the external. Fifteen new species of ''Caccothryptus'' have been identified since the 2014 redescription, all within the ''compactus'' and ''testudo'' groups. The current number of 34 species is likely to increase further, due to a relative lack of studies and material on Asian Limnichidae. Many species are only known from a tiny number of specimens within a small area. In terms of species count, ''Caccothryptus'' is among the largest genera of minute marsh-loving beetles, behind fellow Limnichidae genera '' Byrrhinus'' and '' Limnichus''.


Description

''Caccothryptus'' beetles range from in length. They have ovoid bodies, ranging from brown to black in color, covered in both long and short layers of
setae In biology, setae (; seta ; ) are any of a number of different bristle- or hair-like structures on living organisms. Animal setae Protostomes Depending partly on their form and function, protostome setae may be called macrotrichia, chaetae ...
; the shorter layer lies flat against the surface of the body, while the longer layer stands erect. The color of the setae varies depending on the angle of illumination. The head is slightly retracted into the pronotum, with long and narrow eleven-segment antennae covered in short setae. The underside of the abdomen (
ventrum The abdomen (colloquially called the gut, belly, tummy, midriff, tucky, or stomach) is the front part of the torso between the thorax (chest) and pelvis in humans and in other vertebrates. The area occupied by the abdomen is called the abdominal ...
) has deep depressions where the long, slender legs connect with the body. The smooth and shiny surface of their
elytra An elytron (; ; : elytra, ) is a modified, hardened forewing of beetles (Coleoptera), though a few of the true bugs (Hemiptera) such as the family Schizopteridae are extremely similar; in true bugs, the forewings are called hemelytra (sometime ...
are covered in dense rows of punctures. Sexual dimorphism is relatively minimal. Females have long
ovipositors The ovipositor is a tube-like organ (anatomy), organ used by some animals, especially insects, for the laying of Egg (biology), eggs. In insects, an ovipositor consists of a maximum of three pairs of appendages. The details and morphology of t ...
and gonocoxites (structures covering the underside of the genitals) which taper to a sharp point. ''Caccothryptus'' species can be distinguished from most other genera in the ''Mandersia'' group due to their lack of slanted frontal sulci on their heads. They can be distinguished from the genus ''Simplocarina'' (which also lacks these sulci) by their long antennae, which extend beyond the posterior end of their pronotum. Irregular punctures of different sizes form stripes on the surface of their
elytra An elytron (; ; : elytra, ) is a modified, hardened forewing of beetles (Coleoptera), though a few of the true bugs (Hemiptera) such as the family Schizopteridae are extremely similar; in true bugs, the forewings are called hemelytra (sometime ...
. Additionally, all ''Caccothryptus'' species have a longitudinally divided region at the apex of their aedeagus, which forms two distinct lobes. They generally live in forests, inside piles of water-logged dead wood adjacent to small streams. They
fly Flies are insects of the order Diptera, the name being derived from the Greek δι- ''di-'' "two", and πτερόν ''pteron'' "wing". Insects of this order use only a single pair of wings to fly, the hindwings having evolved into advanced ...
readily and are most often collected using light traps.


Range

''Caccothryptus'' species are mainly within the
Indomalayan The Indomalayan realm is one of the eight biogeographic realms. It extends across most of South and Southeast Asia and into the southern parts of East Asia. Also called the Oriental realm by biogeographers, Indomalaya spreads all over the Ind ...
biogeographic realm A biogeographic realm is the broadest biogeography, biogeographic division of Earth's land surface, based on distributional patterns of terrestrial animal, terrestrial organisms. They are subdivided into bioregions, which are further subdivid ...
, although some reach the southern frontier of the eastern
Palearctic The Palearctic or Palaearctic is a biogeographic realm of the Earth, the largest of eight. Confined almost entirely to the Eastern Hemisphere, it stretches across Europe and Asia, north of the foothills of the Himalayas, and North Africa. Th ...
. They have been found across
Southeast Asia Southeast Asia is the geographical United Nations geoscheme for Asia#South-eastern Asia, southeastern region of Asia, consisting of the regions that are situated south of China, east of the Indian subcontinent, and northwest of the Mainland Au ...
, and portions of southern China and the northern portions of the
Indian subcontinent The Indian subcontinent is a physiographic region of Asia below the Himalayas which projects into the Indian Ocean between the Bay of Bengal to the east and the Arabian Sea to the west. It is now divided between Bangladesh, India, and Pakista ...
. The
Sunda Islands The Sunda Islands (; Tetun: ''Illa Sunda'') are a group of islands in the Indonesian Archipelago. They consist of the Greater Sunda Islands and the Lesser Sunda Islands. Etymology "Sunda" denotes the continental shelves or landmasses: the Sun ...
and
Philippines The Philippines, officially the Republic of the Philippines, is an Archipelagic state, archipelagic country in Southeast Asia. Located in the western Pacific Ocean, it consists of List of islands of the Philippines, 7,641 islands, with a tot ...
are particular centers of diversity within the genus. Many species are known from a single location, although some have been found across a wide swath of territory. Most notable among these species is ''C. maculosus'', which has a range that encompasses
Mainland Southeast Asia Mainland Southeast Asia (historically known as Indochina and the Indochinese Peninsula) is the continental portion of Southeast Asia. It lies east of the Indian subcontinent and south of Mainland China and is bordered by the Indian Ocean to th ...
plus
Hainan Hainan is an island provinces of China, province and the southernmost province of China. It consists of the eponymous Hainan Island and various smaller islands in the South China Sea under the province's administration. The name literally mean ...
, portions of Indonesia, and the
Andaman Islands The Andaman Islands () are an archipelago, made up of 200 islands, in the northeastern Indian Ocean about southwest off the coasts of Myanmar's Ayeyarwady Region. Together with the Nicobar Islands to their south, the Andamans serve as a mari ...
. The ''jaechi, zetelli'', and ''rouyeri'' species groups are only found in
Maritime Southeast Asia Maritime Southeast Asia comprises the Southeast Asian countries of Brunei, Indonesia, Malaysia, the Philippines, Singapore, and East Timor. The terms Island Southeast Asia and Insular Southeast Asia are sometimes given the same meaning as ...
, while the ''testudo'' group is limited to the mainland. Only the ''compactus'' group stretches across both.


References


Bibliography

* * * * * * * * * * {{Taxonbar, from=Q14932675 Byrrhoidea genera Limnichidae Beetles of Asia Taxa described in 1902