Agulla protomaculata
Encyclopedia
Agulla protomaculata is an extinct species
of snakefly
in the raphidiid genus Agulla
. The species is solely known from the Middle Eocene, Lutetian stage
, Parachute Member, part of the Green River Formation
, in the Piceance Creek Basin and Uinta Basin, Garfield County
, northwestern Colorado
, USA.
, three paratype
females, ten male paratypes, and three paratypes of indeterminate sex. The holotype, number "USNM 31487", is a single female specimen consisting of part and counterpart
fossils. All of the type specimens plus an additional male specimen tentatively identified as belonging to A. protomaculata are currently preserved in the Department of Paleobiology collections housed in the National Museum of Natural History
, located in Washington, D.C.
, USA. A. protomaculata was first studied by Michael S. Engel
of the University of Kansas
, Lawrence, Kansas
, USA. His 2009 type description of the new species was published in the journal Transactions of the Kansas Academy of Science
. Engel coined the specific epithet protomaculata as a combination of the Latin
word "protos" meaning "first" and "macula" meaning "mark". At least one other specimen is known and was figured in a 1995 publication by Dayvault et al, however that specimen is though to still be in the private collection of William Hawes. At the time of the species description, Agulla protomaculata was the only member of the order Raphidioptera to be described from the Green River Formation.
in the females. Overall the species was a dark brown to brown coloration, with the legs shading from light brown to a yellow tone. The head is dark brown with extensive yellow coloration including yellow stripes along the sides and to the rear of the eyes, and a central stripe of yellow running down the center of the head behind the eyes. In females the gentility curved ovipositor is light brown and ranges between 3.47 and 3.8 mm (0.136614173228346 and 0.149606299212598 in) in length. The wings are hyaline
in coloration with a slight darkening of the pterostigma
and averaging 7.42 millimetre (0.292125984251969 in) long for the fore-wings
. The vein structure of the fore-wing and notably the length and placement of the M vein in the hind-wing indicate the species to be a member of the genus Agulla.
Species
In biology, a species is one of the basic units of biological classification and a taxonomic rank. A species is often defined as a group of organisms capable of interbreeding and producing fertile offspring. While in many cases this definition is adequate, more precise or differing measures are...
of snakefly
Snakefly
Snakeflies are a group of insects in the order Raphidioptera, consisting of about 210 extant species. Together with the Megaloptera they were formerly placed within the Neuroptera, but now these two are generally regarded as separate orders....
in the raphidiid genus Agulla
Agulla (snakefly)
Agulla is a genus of modern snakeflies in the family Raphidiidae.Agulla species are predatory, both as adults and larvae. In North America occur exclusively in the Western United States, namely in the Rocky Mountains and westward, including the southwestern deserts.-Systematics:At present 31 living...
. The species is solely known from the Middle Eocene, Lutetian stage
Lutetian
The Lutetian is, in the geologic timescale, a stage or age in the Eocene. It spans the time between and . The Lutetian is preceded by the Ypresian and is followed by the Bartonian. Together with the Bartonian it is sometimes referred to as the Middle Eocene subepoch...
, Parachute Member, part of the Green River Formation
Green River Formation
The Green River Formation is an Eocene geologic formation that records the sedimentation in a group of intermountain lakes. The sediments are deposited in very fine layers, a dark layer during the growing season and a light-hue inorganic layer in winter. Each pair of layers is called a varve and...
, in the Piceance Creek Basin and Uinta Basin, Garfield County
Garfield County
Garfield County is the name of several counties in the United States:* Garfield County, Colorado* Garfield County, Montana* Garfield County, Nebraska* Garfield County, Oklahoma* Garfield County, Utah* Garfield County, Washington...
, northwestern Colorado
Colorado
Colorado is a U.S. state that encompasses much of the Rocky Mountains as well as the northeastern portion of the Colorado Plateau and the western edge of the Great Plains...
, USA.
History and classification
The species is known from a series of eighteen fossils, the holotypeHolotype
A holotype is a single physical example of an organism, known to have been used when the species was formally described. It is either the single such physical example or one of several such, but explicitly designated as the holotype...
, three paratype
Paratype
Paratype is a technical term used in the scientific naming of species and other taxa of organisms. The exact meaning of the term paratype when it is used in zoology is not the same as the meaning when it is used in botany...
females, ten male paratypes, and three paratypes of indeterminate sex. The holotype, number "USNM 31487", is a single female specimen consisting of part and counterpart
Counter slab
Counter slab and slab are the matching halves of a fossil-bearing matrix formed in sedimentary deposits. When excavated the matrix may be split along the natural grain or cleavage of the rock. A fossil embedded in the sediment may then also split down the middle, with fossil remains sticking to...
fossils. All of the type specimens plus an additional male specimen tentatively identified as belonging to A. protomaculata are currently preserved in the Department of Paleobiology collections housed in the National Museum of Natural History
National Museum of Natural History
The National Museum of Natural History is a natural history museum administered by the Smithsonian Institution, located on the National Mall in Washington, D.C., United States. Admission is free and the museum is open 364 days a year....
, located in Washington, D.C.
Washington, D.C.
Washington, D.C., formally the District of Columbia and commonly referred to as Washington, "the District", or simply D.C., is the capital of the United States. On July 16, 1790, the United States Congress approved the creation of a permanent national capital as permitted by the U.S. Constitution....
, USA. A. protomaculata was first studied by Michael S. Engel
Michael S. Engel
Michael S. Engel is an American paleontologist and entomologist. He has undertaken field work in Central Asia, Asia Minor, and the Western Hemisphere, and published more than 300 papers in scientific journals. He was trained at the University of Kansas where in 1993 he received a B.S. in Cellular...
of the University of Kansas
University of Kansas
The University of Kansas is a public research university and the largest university in the state of Kansas. KU campuses are located in Lawrence, Wichita, Overland Park, and Kansas City, Kansas with the main campus being located in Lawrence on Mount Oread, the highest point in Lawrence. The...
, Lawrence, Kansas
Lawrence, Kansas
Lawrence is the sixth largest city in the U.S. State of Kansas and the county seat of Douglas County. Located in northeastern Kansas, Lawrence is the anchor city of the Lawrence, Kansas, Metropolitan Statistical Area, which encompasses all of Douglas County...
, USA. His 2009 type description of the new species was published in the journal Transactions of the Kansas Academy of Science
Transactions of the Kansas Academy of Science
Transactions of the Kansas Academy of Science is a biannual peer-reviewed academic journal published by the Kansas Academy of Science. The journal covers biological and physical sciences, mathematics and computer science, history, culture, and philosophy of science, and science education....
. Engel coined the specific epithet protomaculata as a combination of the Latin
Latin
Latin is an Italic language originally spoken in Latium and Ancient Rome. It, along with most European languages, is a descendant of the ancient Proto-Indo-European language. Although it is considered a dead language, a number of scholars and members of the Christian clergy speak it fluently, and...
word "protos" meaning "first" and "macula" meaning "mark". At least one other specimen is known and was figured in a 1995 publication by Dayvault et al, however that specimen is though to still be in the private collection of William Hawes. At the time of the species description, Agulla protomaculata was the only member of the order Raphidioptera to be described from the Green River Formation.
Description
Specimens of Agulla protomaculata average 8.33 millimetre (0.327952755905512 in) in length not including the ovipositorOvipositor
The ovipositor is an organ used by some animals for oviposition, i.e., the laying of eggs. It consists of a maximum of three pairs of appendages formed to transmit the egg, to prepare a place for it, and to place it properly...
in the females. Overall the species was a dark brown to brown coloration, with the legs shading from light brown to a yellow tone. The head is dark brown with extensive yellow coloration including yellow stripes along the sides and to the rear of the eyes, and a central stripe of yellow running down the center of the head behind the eyes. In females the gentility curved ovipositor is light brown and ranges between 3.47 and 3.8 mm (0.136614173228346 and 0.149606299212598 in) in length. The wings are hyaline
Hyaline
The term hyaline denotes a substance with a glass-like appearance.-Histopathology:In histopathological medical usage, a hyaline substance appears glassy and pink after being stained with haematoxylin and eosin — usually it is an acellular, proteinaceous material...
in coloration with a slight darkening of the pterostigma
Pterostigma
The pterostigma is a cell in the outer wing of insects which is often thickened or coloured and so stands out from other cells. It is particularly noticeable in dragonflies, but present also in other insect groups, such as snakeflies, hymenopterans and megalopterans.The purpose of the pterostigma,...
and averaging 7.42 millimetre (0.292125984251969 in) long for the fore-wings
Insect wing
Insects are the only group of invertebrates known to have evolved flight. Insects possess some remarkable flight characteristics and abilities, still far superior to attempts by humans to replicate their capabilities. Even our understanding of the aerodynamics of flexible, flapping wings and how...
. The vein structure of the fore-wing and notably the length and placement of the M vein in the hind-wing indicate the species to be a member of the genus Agulla.