taxonID	type	description	language	source
7170D74F6A13FFC6FF4FFA588FF1DB33.taxon	description	This recently described (Carles-Tolrá 2008) species is known only from the male holotype (preserved in ethanol in the Carles-Tolrá collection in Barcelona, Spain) swept from Umbelliferae on 24 June 1995 at Salamanca, northwestern Spain. According to the original description, the species keys out to couplet 10 in Rozkošný & Elberg (1991) and is easily distinguishable by “ Pleurae brown, only propleura and anterior part of anepisternum (between forecoxa and anterior spiracle) yellowish. ” The anterior surstyli are similar to those of C. pectoralis. The cerci are fused basally, elongate apically, and acute.	en	Bratt, Albertus D., Knutson, Lloyd V., Murphy, William L., Daniels, Anthony A. (2020): Biology, immature stages, and systematics of snail-killing flies of the genus Colobaea (Diptera: Sciomyzidae), with overviews of aspects of the tribe Sciomyzini. Zootaxa 4840 (1): 1-64, DOI: 10.11646/zootaxa.4840.1.1
7170D74F6A13FFC4FF4FF8F08CA2D809.taxon	description	(BNN 6512, 6551, 6601, 6608, 6611, 6627)	en	Bratt, Albertus D., Knutson, Lloyd V., Murphy, William L., Daniels, Anthony A. (2020): Biology, immature stages, and systematics of snail-killing flies of the genus Colobaea (Diptera: Sciomyzidae), with overviews of aspects of the tribe Sciomyzini. Zootaxa 4840 (1): 1-64, DOI: 10.11646/zootaxa.4840.1.1
7170D74F6A13FFC4FF4FF8F08CA2D809.taxon	description	Colobaea americana was described from four specimens from Aylmer, Quebec, and Aweme, Manitoba, both in Canada. In New York, USA (at Cobleskill, Geneva, Ithaca, and Spencer), ADB and LVK collected puparia that subsequently produced adult flies. C. O. Berg (in litt.) collected others at Lacombe, Alberta, Canada. The only records of field-collected adult flies are 31 July (Aylmer, Quebec) and 11 August (Aweme, Manitoba) (Steyskal 1954), but Tom Murray (Groton, Massachusetts, USA) posted two photographs that he took in the field on 16 May 2005 in Harvard, Worcester County, Massachusetts, of a live C. americana resting on a leaf. The habitat distribution of C. americana is rather broad, as is that of its most frequent snail prey, Gyraulus parvus (Say). The collecting sites at Cobleskill, Ithaca, and Spencer, New York, and at Lacombe, Alberta, included small, permanent, exposed ponds. A vernal swamp at Geneva, New York, was extremely productive for collecting puparia in floating snail shells. The Geneva locality, Savage Road Swamp, is described in detail in Bratt et al. (1969); all laboratory rearings were initiated with adult flies that emerged from puparia collected there during 1962, 1965, and 1966 (FT 6206, 6501 – 02, 6512, 6551, 6601 – 02, 6606 – 07, 6611). Adult flies usually mated on the day of emergence and continued to mate for as long as 10 days. While mating, the male positioned his foretarsi on the parafrontal areas of the female’s head or on the anterolateral surfaces of her thorax, held his midlegs above her wings, and laterally grasped her postabdomen with his hind tarsi. Adult flies occasionally consumed the mucus of living snails in addition to the mixture of honey, yeast, and powdered milk with which they were provided. The preoviposition period was 4 – 7 days. In the laboratory, adult females oviposited throughout their lives, laying eggs in no apparent pattern on damp moss or cotton, never on living or dead snails. The incubation period at room temperature was about 24 hours. Larvae appeared to prefer the snail G. parvus as prey. Many puparia were found in shells of G. parvus at Geneva, New York, and a few puparia were collected in G. parvus shells at Cobleskill and Spencer, New York, and at Lacombe, Alberta. One puparium was found in the shell of a Physa Dreparnaud sp. (J. L. Bath, collector). During laboratory rearings, larvae killed and consumed G. parvus but did not attack Helisoma trivolvis (Say) or Physa sp. Unlike larvae of many other species of Sciomyzini, newly hatched larvae did not crawl about on the shell or exposed soft parts of the snail and feed on mucus. Instead they immediately inserted themselves between the mantle and shell of the snail, which invariably reacted by retracting into its shell until its soft parts were about one-third of a whorl from the aperture. Each first-instar larva subsequently remained between the mantle and shell, with its posterior spiracles barely exposed. Usually only one first-instar larva, but occasionally as many as three, attacked an individual snail. Each snail infested with one larva lived for about two days after invasion. After the snail had died, the larva continued to feed, consuming almost all of the decomposing snail tissues. A single G. parvus of about 3 mm in diameter appeared to provide enough nourishment for the development of a larva from hatching to pupariation. If a snail penetrated by a newly hatched larva was considerably smaller than 3 mm, the larva subsequently killed and consumed a second snail. During laboratory trials, larvae were unable to subdue snails 5.5 mm or greater in diameter. The total duration of the larval stage was 5 – 6 days. All puparia were formed in snail shells. Before pupariating, larvae pushed any remaining decayed tissue to the aperture of the shell and formed a thin, plate-like encrustation of a small amount of calcareous material at the aperture. The origin and formation of this structure are similar to that involved in the production of “ septa ” by pupariating larvae of certain species of Pherbellia (Knutson et al. 1967, Bratt et al. 1969). As in larvae of some species of Pherbellia, the Malpighian tubules of mature larvae (where the calcareous material apparently is produced / stored) of C. americana appeared very large and white, but, unlike the situation with Pherbellia larvae, those of C. americana formed the septum while still in a feeding position, i. e. with the anterior end deep in the shell and the posterior end near the aperture. The difference in positions of the larvae at the time of excretion of the septum material probably is related to the fact that whereas larvae of Pherbellia produce a relatively large amount of septum material, larvae of C. americana produce only a small amount, retaining most of it. Larvae of Pherbellia can “ afford ” to expend some of the material in displacing it from the inner whorls of the shell to the aperture by peristaltic body movements, but larvae of C. americana apparently cannot. Septa of C. americana were formed at the aperture of the shell, not several millimeters inside the aperture as with species of Pherbellia. Some of the remaining snail tissue that the pupariating C. americana larvae pushed to the aperture usually became incorporated into the septum. The function of the septum is unknown; it might ensure successful overwintering by aiding in flotation of shells containing puparia or it might serve as a physical barrier against potential predators. In the laboratory, many puparia were formed between 20 January and 6 June. The puparial period lasted 9 – 11 days. Adult flies emerged between 29 January and 17 July. Laboratory-reared males lived 11 – 47 days. Females lived 16 – 60 days. Colobaea americana apparently overwinters in the puparium. Puparia were found in the field at Geneva, New York, on 11, 14, 27, and 28 April and on 6 November. The puparia collected during April and held at room temperature produced adult flies 10 – 23 days later. Astonishingly, from puparia collected 6 November 1963 and held at 5 ° C until 20 December 1965, adult flies emerged 9 – 19 days after being returned to room temperature, having spent 775 days in diapause. Seasonal development of the species apparently is limited only by low temperatures. This conclusion also is supported by the fact that five consecutive generations were produced in the laboratory at relatively stable ambient indoor temperatures. Hymenopterous parasitoids of Ichneumonidae and Pteromalidae were reared from puparia collected at Geneva, New York. Seven Orthizema n. sp. (Ichneumonidae) (determined by R. W. Carlson, Systematic Entomology Laboratory, Agricultural Research Service, U. S. Department of Agriculture) emerged between 18 and 21 May, one each from seven puparia collected 27 April. Two Eupteromalus sp. (Pteromalidae) (determined by B. D. Burks, same affiliation as R. W. Carlson) emerged 27 April and 14 May, one each from two puparia collected 14 April, and another emerged 11 May from a puparium collected 28 April.	en	Bratt, Albertus D., Knutson, Lloyd V., Murphy, William L., Daniels, Anthony A. (2020): Biology, immature stages, and systematics of snail-killing flies of the genus Colobaea (Diptera: Sciomyzidae), with overviews of aspects of the tribe Sciomyzini. Zootaxa 4840 (1): 1-64, DOI: 10.11646/zootaxa.4840.1.1
7170D74F6A11FFC4FF4FFBCA8D74D961.taxon	description	Colobaea beckeri is a rare species with entirely black anepisternum and anepimeron. It is distinguished from C. distincta by its longer arista and basal flagellomere, wing veins blackish with infumated crossveins, shining medifacies, and by having only the apical segment of the foretarsus white. Colobaea beckeri was described from a specimen collected in Bad Hall, Austria (19 June); Soós (1958) recorded it from Verebély (= Vráble), Czech Republic (22 May). Nothing is known about its biology or immature stages.	en	Bratt, Albertus D., Knutson, Lloyd V., Murphy, William L., Daniels, Anthony A. (2020): Biology, immature stages, and systematics of snail-killing flies of the genus Colobaea (Diptera: Sciomyzidae), with overviews of aspects of the tribe Sciomyzini. Zootaxa 4840 (1): 1-64, DOI: 10.11646/zootaxa.4840.1.1
7170D74F6A11FFC1FF4FFA628B4ADB1D.taxon	description	(BNN 6426, 6717, 6763)	en	Bratt, Albertus D., Knutson, Lloyd V., Murphy, William L., Daniels, Anthony A. (2020): Biology, immature stages, and systematics of snail-killing flies of the genus Colobaea (Diptera: Sciomyzidae), with overviews of aspects of the tribe Sciomyzini. Zootaxa 4840 (1): 1-64, DOI: 10.11646/zootaxa.4840.1.1
7170D74F6A11FFC1FF4FFA628B4ADB1D.taxon	materials_examined	Type locality. “ In Kjellunge Gothlandiae [Sweden] ” (Fallén, 1820). 1 ♀, NHRS, Stockholm. Its known geographical distribution — from Counties Mayo and Wexford, Ireland, across northern and central Europe to Hungary (Kalocsa) and Moscow, northward to Vuollerim, Sweden and northernmost Siberia, and east to the South Maritime Territory of Russia — extends farther north and east than that of any other species of Colobaea. Colobaea bifasciella appears to be able to exploit its hosts nearly to the northern, most rigorous limits of their distributions, but C. bifasciella apparently has not expanded significantly into the warmer, southern parts of the host snails’ ranges. Capture records of adult flies range from 10 May (Strachotin, Czech Republic) to 12 September (Gyón, Hungary). Adult flies of C. bifasciella are found in various macrohabitats, as are those of other species of Sciomyzidae whose immature stages occur only in restricted microhabitats. However, because of their obligatory trophic relationship with G. truncatula and S. palustris, we know that the larval microhabitat must fall somewhere within those occupied by the host snails. The larval microhabitats are further limited to areas in which G. truncatula or S. palustris are stranded, aestivating, foraging out of the water, or otherwise exposed. Submerged snails cannot serve as oviposition sites nor as hosts for larvae of Colobaea. Where a large, dense population of C. bifasciella was encountered at Frederikslund, near Holte, Denmark, C. bifasciella eggs were found on snails among moist, dead leaves near a small permanent pond in a heavily shaded vernal swamp. Infested snails also were collected from muddy banks bordered by Carex spp. in exposed marshy areas along rivers in northern Sweden. A few puparia were found in shells floating among emergent grasses at the edge of a small, unshaded vernal marsh during spring at Suserup, near Sorø, Denmark. Laboratory rearings and observations were initiated with many eggs, larvae, pupae, and adult flies collected 7 May – 22 August 1964 (FT 6436, 6451 – 6452, 64100) near Sorø and Holte, Zealand, Denmark; 8 – 28 July 1967 (FT 6721 – 6731) at Brattby, near Umeå, Sweden; and with 1 ♂ collected 14 June 1967 (FT 6705) and 1 ♀ collected 8 July 1967 (FT 6721) near Vuollerim, Sweden. Adult flies mated readily in the laboratory between 27 May and 7 August. Females copulated for the first time 9 – 12 days after emergence. Females collected on 23 May in Denmark and on 8 July in northern Sweden and then held without males laid viable eggs. The copulatory postures were similar to those of most other species of Sciomyzidae. The male placed his foretarsi on the dorsomesal margins of the female’s compound eyes, grasped the basiventral surfaces of her half-outstretched wings or the sides of her thorax with his midtarsi, and held the sides of her postabdomen with his hind tarsi. Copulation commenced within 10 minutes after males and females had been placed in the same jar. Preoviposition periods of laboratory-reared females ranged from 12 to 22 days. Females laid eggs from 24 May to 16 August. A female collected on 23 May laid 72 eggs between 24 and 29 May, and a female collected 8 July laid 343 eggs between 9 July and 19 August. Field-collected S. palustris, usually 10 snails of the same sizes as found infested by C. bifasciella in nature, were added to the chamber with the latter female for 24 - hour periods during the first half of her oviposition period and for longer periods thereafter. Some snails died in the oviposition chamber; she laid no eggs on them. Daily egg production ranged from 3 to 15 and averaged 8.3 eggs per day. Egg production did not decrease markedly when snails were left in the chamber with the female for several days; under these conditions, she simply laid additional eggs on each snail. One female that emerged in the laboratory laid 55 eggs between 16 and 21 July, and another laid 18 eggs between 9 and 16 August. Unhatched viable eggs were found on snails in the field between 25 May and 23 July. Almost all eggs found in nature, as well as those laid in the laboratory, had been placed across the sutures of living S. palustris (Figs 2 – 3). However, a viable egg was found on the shell of a living terrestrial snail, Balea perversa (L.), and another was found on an empty shell of S. palustris during a period when the population of C. bifasciella at Frederikslund, Denmark, was very dense. Other species of aquatic and terrestrial snails occurring with the infested S. palustris were offered as hosts to adult female C. bifasciella but seldom were accepted as oviposition sites. On a few occasions, in the laboratory, female C. bifasciella laid eggs on Aplexa hypnorum (L.), young Lymnaea stagnalis (L.) (4.5 – 5.0 mm in length), and Radix peregra (O. F. Müller). Lundbeck (1923) stated that all puparia he found were in shells of G. truncatula; Dr. G. Mandahl-Barth (in litt.) confirmed Lundbeck’s determination of the host snail. In rearing jars crowded with adult flies, as many as 29 eggs were laid on each snail, but most shells bore only one or two eggs, whereas with snails collected in nature only one egg (occasionally two eggs) were found on each infested S. palustris. Although Lundbeck (1923) made no mention of eggs of C. bifasciella, many of the snail shells he studied and which ADB and LVK examined at ZMUC still bear one or two collapsed eggshells across the sutures. The incubation period at room temperature was 3 – 5 days. Eggs usually hatched at the end nearest the aperture of the snail shell. The feeding behavior of C. bifasciella larvae is one of the most specialized of all Sciomyzidae. It seems to be identical with that of its Nearctic ecological / behavioral equivalent, Sciomyza varia (Berg 1964, Barnes 1990). According to the definition of Reuter (1913), C. bifasciella should be classified as a parasitoid. That is, the larva feeds carefully within the host in such a way that the host is not killed until the larva is relatively well developed, at which time the larva consumes most of the remaining tissue. Such an insidious manner of feeding during early larval life, with a transition to predaceous habits and finally to saprophagous habits, indicates the evolution of a series of adaptations that are finely adjusted to the activity, defensive reactions, and physiological conditions of the host snail. The occurrence of these three trophic stages (parasitoid, predaceous, and saprophagous) in the development of each larva is reminiscent of the feeding behavior of some individuals of Atrichomelina pubera (Loew) (Foote et al. 1960) and of some species of Pherbellia (Bratt et al. 1969). However, the behavior of larval A. pubera is much more variable than that of C. bifasciella in that larval A. pubera also can develop entirely in a parasitoid / predaceous or saprophagous manner. A simple but effective technique used during the 1964 and 1967 rearings by ADB and LVK permitted exceptionally close observation of the development of C. bifasciella larvae; see also under Materials and Methods, above, the technique used by Barnes (1990) to study Sciomyza varia. In the laboratory, some snails bearing C. bifasciella eggs were removed from oviposition jars and placed separately in 1 x 5 - cm glass vials containing a wad of damp cotton and plugged with dry cotton. Each snail soon affixed its aperture to the side of the tube. Development of C. bifasciella larvae was then observed through the walls of the tube under 50 X magnification by use of a stereoscopic microscope. As soon as the egg hatched, the larva crawled down the side of the shell and penetrated between the mantle and foot until only its posterior spiracles remained exposed. Larvae were very sensitive to sudden movement. When disturbed, they pulled their posterior end below the surface of the snail tissues and remained so for long periods of time. None of the viscous, colorless mucus normally produced by S. palustris when irritated was secreted when the tiny (about 1.0 mm in length) C. bifasciella larvae attacked the snail. When more than one egg had been laid on a snail shell, all larvae that emerged entered the snail, but then all except one of them left the snail within about one day, even though the established larva did not overtly attack the other larvae. Individual snails remained active for as long as 10 days after larval invasion, but it usually produced an epiphragm within a few hours and thereafter remained motionless. The substance consumed by the newly established larva was not identified during this study, but it seems likely that the youngest larvae feed on mucus and extrapallial fluid. Sometime during the first or second stadium, larvae began consuming nerve tissue between the eye stalks. This subdued the snails but did not kill them immediately. The posterior spiracles of newly molted second-instar C. bifasciella larvae became clearly visible between the retracted foot and the mantle of the still-living snails 7 – 10 days after the initial attack. The larvae then began to consume all snail tissue within reach. Infested snails usually died and began to decay shortly before the larvae completed the second stadium, but sometimes a pin prick elicited slight muscular movements of the foot and mantle even after a larva had reached the third stadium. Larvae continued to feed on decaying snail tissue for the remaining 1 – 2 days of the second stadium and for the entire 4 – 6 days of the third stadium. In the laboratory, the total duration of larval life ranged from 20 to 31 days (first stadium, 10 – 12; second, 6 – 13; and third, 4 – 6). When the host snail of a first- or young second-instar larva died, the larva remained in the dead snail shell but died after a few days. Each larva developed through all three stadia in a single snail. Additional snails placed in the rearing containers never were attacked, even when the first host snail was very small. The range in width / length measurements of 19 S. palustris attacked in nature was 2.2 – 3.8 mm wide and 4.8 – 11.0 mm long; the range for seven hosts attacked in the laboratory was 3.4 – 5.5 mm wide and 4.8 – 10.1 mm long. Adult flies that emerged from puparia in small shells were, in some instances, only half as large as those that developed in larger hosts. Larvae consumed all or almost all of the mostly decayed snail tissues before pupariating, at which time they pushed any remaining tissues and debris out of the snail shell. Often the muscular anterior part of the foot, the mantle, and the ovotestis could be seen among this material. Such emptying of the shell seems to have two advantages: it reduces the danger of mold growth that might kill the pupa or attract enemies, and it makes the shell somewhat more buoyant. No septum material such as that excreted by pupariating larvae of C. americana and by several species of Pherbellia was produced by any pupariating C. bifasciella larvae. The anterior portion of the puparium was characteristically expanded to close off the shell whorl (Fig. 4). Lundbeck (1923) reported obtaining adult flies during spring from puparia found in flood debris. A puparium that ADB and LVK collected on 7 May 1964 at Suserup Skov, near Sorø, Denmark, produced a female on 16 May. In the laboratory, 81 puparia were formed by field-collected larvae, from larvae that had developed from field-collected eggs, or from larvae that had developed from eggs laid in the laboratory. Fourteen adult flies emerged between 10 January and 12 February 1968 from the 57 puparia formed between 26 July and 21 August 1967 from the rearing (BNN 6763) initiated with a female collected 8 June 1967 at Brattby, near Vuollerim, Sweden. Twenty-four puparia formed between 14 June and 28 August 1964 were held at 76 % relative humidity at room temperature; 18 adult flies emerged between 2 July and 19 December. The puparial periods for 10 of these adult flies (7 ♀, 3 ♂) were not of exceptional duration, ranging from 14 to 21 days (average 17.4 days), in contrast to the duration of six other puparia formed at the same time by both laboratory-reared and field-collected larvae; these required 114 – 154 days (average 131.8 days) to produce adult flies (1 ♀, 5 ♂). Also recorded were two puparial periods of intermediate (63 and 81 days) duration. In this initial study of the basic life cycle of C. bifasciella, ADB and LVK were able to show only that the duration of the puparial period is variable; it may be very short and similar to summer generational periods of other species of Sciomyzidae, or it may be as much as 10 X longer than those and include a quiescent stage. Because both quiescent and nonquiescent pupae were obtained from the same group of eggs laid in the laboratory and held at the same temperature and humidity, it seems likely that the occurrence of puparial periods of two distinctly different durations is controlled genetically and is not solely a function of environmental conditions. Similar behavior was observed during rearings of Pherbellia similis (Cresson) (Bratt et al. 1969). Regarding longevity, in the laboratory, four field-collected adult flies lived 4, 7, 42, and 50 days, and six laboratory-reared adult flies lived 6 – 32 days. The fact that adult flies emerge soon after puparia are collected in the spring, that eggs in nature are laid as early as 25 May, and that puparia formed in late summer and held at room temperature may require as long as 154 days before emergence of adult flies indicates that C. bifasciella overwinters in the puparium. Regarding natural enemies, Lundbeck (1923) obtained “ a Cryptine and a Chalcidid ” from puparia collected during spring; although thought to be deposited in the ZMUC, Thomas Pape informed LVK (in litt. March 2015) that these two parasitoids could not be located there. A puparium collected at Sorø, Denmark on 9 June 1964 and held continuously at room temperature and 76 % relative humidity produced an ichneumonid wasp on 10 July 1965.	en	Bratt, Albertus D., Knutson, Lloyd V., Murphy, William L., Daniels, Anthony A. (2020): Biology, immature stages, and systematics of snail-killing flies of the genus Colobaea (Diptera: Sciomyzidae), with overviews of aspects of the tribe Sciomyzini. Zootaxa 4840 (1): 1-64, DOI: 10.11646/zootaxa.4840.1.1
7170D74F6A14FFC0FF4FF8FE8F7FDD59.taxon	description	Map 5 Colobaea canadensis has been found only in the province of Manitoba, Canada. It most closely resembles C. bifasciella in that both species have a dull, tomentose, setose, yellow frons; a similarly shaped head; two pairs of postalar setae; and R 1 extending to the level of the r-m crossvein. These two species can be readily distinguished from each other, however, as C. canadensis has a mainly shiny black body and unpatterned wings, whereas C. bifasciella has a yellow body with black markings and patterned wings. Nothing is known about the biology or immature stages of C. canadensis.	en	Bratt, Albertus D., Knutson, Lloyd V., Murphy, William L., Daniels, Anthony A. (2020): Biology, immature stages, and systematics of snail-killing flies of the genus Colobaea (Diptera: Sciomyzidae), with overviews of aspects of the tribe Sciomyzini. Zootaxa 4840 (1): 1-64, DOI: 10.11646/zootaxa.4840.1.1
7170D74F6A15FFC0FF4FFE3A8DEADB8D.taxon	description	(BNN 7105 – 07, 7317 – 18, 7343) This species, from Iran and Pakistan, is similar in color pattern to the African C. occidentalis, known from Niger, northern Nigeria, and Ethiopia, and to C. punctata, a widespread Palearctic species found from Ireland to Israel (northern Negev), northeastern Iran (Mazanderan), and east to Kazakhstan and eastern Siberia. The biology and immature stages of C. deemingi are known only from progeny reared from adult flies collected by LVK in southwestern Iran (five localities near Dezful, one locality near Shooshtar) during November 1971 (FT 7110) and April 1973 (FT 7320, 7334). During November 1971, adult flies were found at only two localities, but during April 1973 they were found at six localities. Most adult flies were swept from hygrophilous grasses and Carex emerging from a few inches of water at the margins of marshes near the Persian Gulf. The 0.5 – 20 hectare marshes were rimmed by extensive growths of Typha angustifolia. At Boneh Javaz, 41 km SE of Dezful, in November 1971, adult flies were swept from wet, grassy areas between patches of Typha growing in unshaded, shallow depressions receiving water from a nearby irrigation channel. Adult flies were especially common at that location; in fact, during April 1973, C. deemingi was one of the most abundant species of flies at a site in Bayza, 610 km SE of Dezful. No C. deemingi were collected in a Malaise trap placed where adult flies had been collected with a sweep net during November 1971 at Boneh Sayyed Moosa, 16 km W of Dezful. No larvae or puparia of C. deemingi have been found in nature. For detailed descriptions of typical aquatic habitats in southwestern Iran, with discussion of the ecology of freshwater snails, see Chu et al. (1968). Other species of Sciomyzidae found with C. deemingi in Iran and Pakistan included Ditaeniella grisescens (Meigen), Hydromya dorsalis (Fabricius), Ilione turcestanica (Hendel), Pherbellia cinerella (Fallén), P. fuscipes (Macquart), Sepedon sphegea (Fabricius), and S. spinipes (Scopoli). Aquatic snails present at the collecting sites included Bulinus truncatus (Audouin), Gyraulus intermixtus (Mousson), Melanoides tuberculata (O. F. Müller), Planorbis planorbis, and Radix gedrosiana (Say). Adult flies collected on 4 November 1971 mated the same day and continued to mate until 14 November; adult flies collected on 16 April 1973 were observed mating between 17 and 29 April; and a pair of adult flies were collected in copula on 25 April. In the laboratory, adult flies that emerged between 25 and 28 November first mated eight days later and produced viable eggs during December 1971 and January 1972. Adult flies collected 16 April 1973 were first seen to oviposit on 21 April. Females scattered eggs over the damp moss or cotton in the bottom of the breeding jar and did not oviposit on a living snail. The incubation period was about 48 hours. Larvae killed and consumed G. intermixtus of 2.5 – 6.5 mm in greatest diameter. Mature larvae also killed and fed on Radix gedrosiana of about 4.0 mm in length, but first-instar larvae did not attack small Lymnaea sp. or Helisoma trivolvis. Each larva consumed one or two G. intermixtus or R. gedrosiana. The larval feeding behavior was very similar to that of C. punctata. Several larvae often fed together in a single snail. They continued to feed on black, putrid tissues long after the snail prey had died. The total duration of larval life was 8 – 10 days. Puparia were formed in shells of G. intermixtus and occasionally in shells of R. gedrosiana. Pupariating larvae produced a mass of frothy septum material in the aperture of the shell. In Iran, larvae pupariated between 13 November and 16 December 1971 and between 23 April and 3 May 1973. Twenty-five adult flies emerged between 25 November and 8 December 1971, and many more emerged between 5 and 15 May 1973. A complete second generation was not reared from these adult flies because of the authors’ travel requirements. The puparial period lasted about 12 days. The total duration of the immature stages was 22 – 24 days.	en	Bratt, Albertus D., Knutson, Lloyd V., Murphy, William L., Daniels, Anthony A. (2020): Biology, immature stages, and systematics of snail-killing flies of the genus Colobaea (Diptera: Sciomyzidae), with overviews of aspects of the tribe Sciomyzini. Zootaxa 4840 (1): 1-64, DOI: 10.11646/zootaxa.4840.1.1
7170D74F6A1AFFCFFF4FFF3F8D66DF21.taxon	description	Colobaea distincta is a rather common species that may be identified by the entirely black anepisternum and anepimeron, arista shorter than face, basal flagellomere only twice as long as its width at base, two apical segments of the foretarsus white, all wing veins yellow and infumated, and middle of face tomentose. The adult male was figured beautifully by Hendel (1901). Colobaea distincta ranges from northwestern Ireland (County Mayo) and the UK (Porthcawl, Wales and Devonshire, England) across central Europe to St. Petersburg, Russia (at the eastern end of the Gulf of Finland) in the north and to Bulgaria (Bagau) and southern Russia (Sarepta, near Volgograd) in the south. The habitat distribution of C. distincta is quite broad. Verbeke (1948) cited it as being found in “ bois ” (= forest) at Melle and Schelderode, Belgium. P. Skidmore (in litt. to LVK) collected the species in a marsh by the River Avon in southern England. Stora (1956) collected adult flies on the seashore at Lillsanviken, in Faboda, Finland. Stackelberg (1958) recorded it from marshy margins of lakes near Leningrad, Russia. Colobaea distincta seems to have a relatively long flight period, having been collected between 5 May and 12 September in southern England and on similar dates on the European continent. Rozkošný (1967) obtained 1 ♀ on 16 April from a puparium in a shell of Anisus spirorbis (L.) collected 3 April 1965 at Vranovice, Czech Republic. Beuk (1993) reared 3 ♂ collected 24 April 1986 from “ flood debris ” (including snail shells) from backwaters of the Rhine River on 14 April.	en	Bratt, Albertus D., Knutson, Lloyd V., Murphy, William L., Daniels, Anthony A. (2020): Biology, immature stages, and systematics of snail-killing flies of the genus Colobaea (Diptera: Sciomyzidae), with overviews of aspects of the tribe Sciomyzini. Zootaxa 4840 (1): 1-64, DOI: 10.11646/zootaxa.4840.1.1
7170D74F6A1AFFCEFF4FFC238C4ADCCD.taxon	description	As noted above, Li et al. (2019) recorded Colobaea in China for the first time during their review of sciomyzid genera known from China. Their specimen data are as follow: 2 ♂, CHINA, Inner Mongolia, Genhe, 835 m, 2016. VII. 30, Li Shi (IMAU); 2 ♂, CHINA, Inner Mongolia, Genhe, 870 m, 2016. IX. 27, Li Shi (IMAU). Colobaea sp.: 1 ♀, CHINA, Xinjiang, Tacheng, 1955. VII. 24, Shijun Ma (IZCAS). Nothing is known about the biology or immature stages of this species. As noted by Sueyoshi (2001), C. eos and C. flavipleura can be distinguished from their congeners by the presence of a single pair of dorsocentral setae.	en	Bratt, Albertus D., Knutson, Lloyd V., Murphy, William L., Daniels, Anthony A. (2020): Biology, immature stages, and systematics of snail-killing flies of the genus Colobaea (Diptera: Sciomyzidae), with overviews of aspects of the tribe Sciomyzini. Zootaxa 4840 (1): 1-64, DOI: 10.11646/zootaxa.4840.1.1
7170D74F6A1BFFCEFF4FFE8E8FF5DE25.taxon	description	Colobaea flavipleura was described from a male collected in “ USSR, South Maritime Territory, Kamenushka, 27. vii. 1983. ” The holotype is deposited in the ZMUM (male genitalia on slide). The type series included 7 ♂ and 6 ♀, the deposition of which was not noted in the original description. This species has been found only in Russia (Amur area: Zeya. Novosibirskaya Oblast: Kamenushka. Primorsky Krai: Ussuriysk). Nothing is known about its biology or immature stages.	en	Bratt, Albertus D., Knutson, Lloyd V., Murphy, William L., Daniels, Anthony A. (2020): Biology, immature stages, and systematics of snail-killing flies of the genus Colobaea (Diptera: Sciomyzidae), with overviews of aspects of the tribe Sciomyzini. Zootaxa 4840 (1): 1-64, DOI: 10.11646/zootaxa.4840.1.1
7170D74F6A1BFFCEFF4FFD268AE9DFD1.taxon	description	Adult flies of C. limbata appear rather similar to those of C. pectoralis and C. punctata, from which they may be distinguished by the presence of two black spots below the notopleural suture; the apically narrowed basal flagellomere, which is 3 X as long as the width at base; and the dark, infuscated crossveins. The type specimens in the ZIN (1 ♀, 1 ♂) were collected at “ Chiva ” [Khiva], Uzbekistan, on 21 April (Hendel 1933). Colobaea limbata also is known from four specimens (3 ♀, 1 ♂) collected 24, 27, and 28 July 1965 at three localities in nearby Tajikistan (Elberg & Remm 1974). Nothing is known about the biology or immature stages of this species.	en	Bratt, Albertus D., Knutson, Lloyd V., Murphy, William L., Daniels, Anthony A. (2020): Biology, immature stages, and systematics of snail-killing flies of the genus Colobaea (Diptera: Sciomyzidae), with overviews of aspects of the tribe Sciomyzini. Zootaxa 4840 (1): 1-64, DOI: 10.11646/zootaxa.4840.1.1
7170D74F6A1BFFCEFF4FFBB28C12D985.taxon	description	Colobaea montana has been found only in the state of Montana, USA. With a mostly matt black frons and R 1 terminating considerably basad of the level of crossvein r-m, C. montana is more similar to C. americana, C. nigroaristata, C. punctata, and related species than it is to C. bifasciella and C. canadensis. Colobaea montana is most similar to C. nigroaristata, males of which can be separated readily by the internal structures of their terminalia and most notably the shape of the posterior surstylus (see Rozkošný 1984 a). The shape of the posterior surstylus, short rather than long dorsobasal setulae on the arista, and lack of silvery tomentum along the margin of the eye distinguish C. montana from C. americana, the only other species of Colobaea likely to occur with C. montana. Nothing is known about its biology or immature stages.	en	Bratt, Albertus D., Knutson, Lloyd V., Murphy, William L., Daniels, Anthony A. (2020): Biology, immature stages, and systematics of snail-killing flies of the genus Colobaea (Diptera: Sciomyzidae), with overviews of aspects of the tribe Sciomyzini. Zootaxa 4840 (1): 1-64, DOI: 10.11646/zootaxa.4840.1.1
7170D74F6A1BFFCEFF4FFA468F51DB1D.taxon	description	Colobaea nigroaristata has been found only in Sweden (Norrbotten Co., Gällivare; Västerbotten Co., Sorsele) and Finland (Lapland: Inari). It is most closely related to C. distincta but differs from the latter species in having a yellowish anterior margin of the frons, wholly black postpedicel and arista, and wider gena; the male posterior surstylus is heart shaped and lacks the slender posterior process of C. distincta. Nothing is known about its biology or immature stages.	en	Bratt, Albertus D., Knutson, Lloyd V., Murphy, William L., Daniels, Anthony A. (2020): Biology, immature stages, and systematics of snail-killing flies of the genus Colobaea (Diptera: Sciomyzidae), with overviews of aspects of the tribe Sciomyzini. Zootaxa 4840 (1): 1-64, DOI: 10.11646/zootaxa.4840.1.1
7170D74F6A1BFFCDFF4FF8FE8DFCDD05.taxon	description	Colobaea occidentalis has been found in only Niger, Nigeria, and Ethiopia. Knutson et al. (2018) described the adult of C. occidentalis, the first species of Colobaea described from sub-Saharan Africa. Colobaea occidentalis is most similar to C. pectoralis and C. punctata, having an unpatterned wing, frons deeply matt black, postpedicel mainly black and twice as long as wide at the base, anepisternum mainly yellow, and male anterior surstylus with peglike processes. Nothing is known about its biology or immature stages.	en	Bratt, Albertus D., Knutson, Lloyd V., Murphy, William L., Daniels, Anthony A. (2020): Biology, immature stages, and systematics of snail-killing flies of the genus Colobaea (Diptera: Sciomyzidae), with overviews of aspects of the tribe Sciomyzini. Zootaxa 4840 (1): 1-64, DOI: 10.11646/zootaxa.4840.1.1
7170D74F6A18FFCBFF4FFEC58FACDCCB.taxon	description	(BNN 6035; FT 6012 – 15, 6447) Colobaea pectoralis and all other species of Colobaea except C. bifasciella have color patterns similar to those of many species of Pteromicra. Colobaea pectoralis is distinguished from other species of Colobaea by the yellow anepisternum and anepimeron, elongate black stripe along the upper margin of the anepisternum, and slightly infumated crossveins. Type locality: Kallunge (Gotland, Sweden), lectotype in the MZLU. Colobaea pectoralis is known from northern and central Europe, extending from Ireland (County Clare) and Finland south throughout southern England, France, and Belgium to the Czech Republic (Pieštany) and east to the St. Petersburg area, the Russian Upland Area (Black Soil Region south of Moscow), and northern Kazakhstan. Becker’s (1903) record from Cairo, Egypt, may pertain to C. punctata, which was not recognized as a distinct species until Lundbeck (1923), or to a related species. Dates of capture records of adult flies range from 19 April (Jakobstad, Finland) to 17 September (Copenhagen, Denmark). Laboratory rearings and observations are based on puparia and adult flies, all collected in Zealand, Denmark: at Snøgedam, Hillerød (FT 6012 – 15) (5 July – 1 September 1960) and at Vollerup Mose, near Sorø (FT 6447) (18 May 1964). Adult flies were swept from low, emergent vegetation around a small, densely shaded pond (Snøgedam). Newly formed puparia were found in shells of Anisus vortex lying on a mat of Lemna and at the bases of Sparganium simplex. A few puparia were found in shells of A. vortex attached to cases of Limnephilus Leach sp. (Trichoptera) at Snøgedam. Shells of Bathyomphalus contortus (L.) containing puparia were found among emergent vegetation in an exposed marsh (Vollerup Mose). Lundbeck (1923), Rozkošný (1967), Nienhuis (1970), and Przhiboro (2001) reared adult flies from puparia found in shells of A. vortex in Denmark, Czech Republic, the Netherlands, and northeastern Russia, respectively. Adult flies collected during the summer mated between 5 July and 28 August. Four pairs that emerged between 11 and 18 January 1961 from puparia formed in the laboratory during the previous autumn were placed in separate breeding vials. All four pairs copulated within 48 hours after emergence, and one pair mated within 24 hours; Nienhuis (1970) also noted that adult flies mated on the day of emergence. Each mating lasted 10 – 60 minutes, with pairs often copulating several times daily. A distinct difference was noted in the copulatory posture of C. pectoralis from that of all other species of Sciomyzidae in which the copulatory posture is known except Ectinocera borealis Zetterstedt (Tetanocerini). Males of most other species place their foretarsi on the head of the female, along the inner margins of her compound eyes. Males of C. pectoralis, however, almost always place their foretarsi on the anterolateral angles of the female’s thorax. The midlegs of the male hold the basicostal margins of her wings, while his hind tarsi grasp the sides of her postabdomen. Females collected during early July oviposited between 7 July and 2 September. Those that emerged in the laboratory during January laid viable eggs four days after emergence. Although sprigs of fresh moss, strips of fresh Typha leaves, and living A. vortex snails were present, females oviposited mainly on wet peat moss at the bottom of the breeding jars. Females frequently were observed to wriggle down through a tangle of dry moss to oviposit onto the wet moss. Females always laid eggs singly and distributed them at random. After incubation periods of 2 – 5 days at 24 ° C, the eggs hatched between 12 July and 3 September and during January. Lundbeck (1923) evidently thought that C. pectoralis was host specific. After stating that he had reared adult flies only from puparia in shells of A. vortex (as “ Planorbis vortex ”), he added that C. bifasciella and C. pectoralis “ are each found only in 1 species of snail though in the places where I collected them also other species of small snails were present. ” ADB and LVK documented larvae of C. pectoralis successfully attacking two additional species of snails: 1) puparia were found in nature in shells of B. contortus that the larvae had evidently killed and consumed, and 2) larvae killed and consumed small individuals of Planorbis planorbis during laboratory rearings in addition to A. vortex. The larvae never attacked larger individuals of P. planorbis, nor did they consume freshly crushed aquatic snails. Most laboratory rearings were conducted with A. vortex. Newly hatched C. pectoralis larvae readily attacked living A. vortex. The larvae penetrated between the mantle and shell, adjacent to the upper side of the aperture, until only their posterior spiracles remained exposed. When reared individually with one snail per larva, the snail usually did not die until after the first larval molt; the larva then continued to feed in the decaying snail tissues. Larvae that attacked full-grown snails completed their development in those individuals, whereas larvae that attacked smaller snails left the emptied shell of the first host and attacked a second and sometimes even a third snail before pupariating. Under crowded rearing conditions, several first-instar larvae would attack and feed within a single snail; such snails usually died within 24 hours. After consuming the tissues of the first snail, these larvae then attacked another snail, either individually or together. Second-instar larvae that were removed from the liquefied tissues of a dead A. vortex and then placed with a living snail attacked it immediately. Upon contact, these larger larvae would slash at the exposed foot or mantle of the snail, causing the snail to retract into its shell. The larvae would continue to slash and hook at the exposed tissues. With each contact, the snail would retract farther. The larvae fed voraciously as they penetrated into the shell. By the time the larvae were in the third stadium, the snail tissues had become a black, putrid, viscous mass. Each larva remained immersed in the liquefied tissue up to its posterior spiracles and consumed most of this material before pupariating. Total duration through all three larval stadia required 8 – 10 days. Larvae of C. pectoralis moved about less actively than did larvae of other sciomyzid species studied by the authors. They were never seen crawling more than a few millimeters up the dry sides of the rearing containers, even when they had not eaten for a day or more. Larvae readily attacked snails lying on wet moss, but larvae apparently did not encounter snails held only a few millimeters above the surface of the wet moss by sprigs of drier moss. When dead snails containing actively feeding larvae were submerged, the larvae quickly wriggled out of the shells but did not float to the surface, remaining instead at the bottom, defecating and extending / retracting their posterior spiracles. Puparia were formed in the laboratory between 26 July and 10 August. Most larvae pupariated in snail shells, with one puparium being formed in each shell. When as much as half of the snail shell had been chipped away to allow observation of a larva throughout its development, the larva eventually left the shell and pupariated on wet filter paper on the bottom of the rearing container or on the dry sides of the container. Some puparia that were formed outside of the shell retained the typical curved shape, but others, especially those formed in a corner of a rearing box or under wet paper, were distorted. Puparia were formed within shells at various distances from the aperture. For example, the anterior end of one puparium was 1 ½ whorls in from the aperture, while the anterior third of another puparium extended beyond the aperture. Usually the anterior end of the puparium was situated at a distance of about one whorl inside the edge of the aperture. Pupariating larvae produced no septum material such as that excreted and formed into an operculum-like structure by C. americana. Pupae became visible through the translucent integument 2 – 3 days after the puparia were formed. Dates of emergence of adult flies indicate that C. pectoralis overwinters within the puparium and that a facultative diapause or period of quiescence may develop in pupae formed during late summer and autumn. Lundbeck (1923) reared adult flies from puparia collected during spring. ADB and LVK examined Lundbeck’s specimens in the ZMUC, where 21 adult flies are pinned with their puparia and with the A. vortex shells in which those puparia had been formed. The shells are labeled as having been collected 12 April 1906 at Utterslev Mose (a bog about 6 km northwest of Copenhagen), but no dates of emergence of the adult flies are noted on the labels. Rozkošný (1967) obtained three adult flies (1 ♀, 2 ♂) on 9 and 13 April from puparia collected 25 March 1965 at Musov, Moravia. Nienhuis (1970) obtained seven adult flies between 10 and 27 April from puparia collected 4 April 1969 in Herweg, The Netherlands. Przhiboro (2001) obtained 1 ♂ on 20 May 1998 from a puparium in a shell of A. vortex collected 7 May 1998 on the shore of Anninskoe Lake near St. Petersburg, Russia. ADB and LVK found puparia in shells of B. contortus on 18 May 1964; adult flies (5 ♀, 3 ♂) emerged between 23 and 30 May. Puparia found in shells of A. vortex between 22 June and 8 August 1960 and held at room temperature produced adult flies (4 ♀, 5 ♂) 1 – 11 days after they had been collected. Other puparia found in shells of A. vortex at the same place and at about the same time (19 July – 8 August 1960) and exposed to 4 ° C and 76 % relative humidity from 23 November to 31 December 1960 produced adult flies (2 ♀, 3 ♂) between 12 and 18 January 1961. These puparial periods ranged from 159 to 183 days. Similarly, puparia formed in the laboratory between 26 July and 10 August 1960 and exposed to 4 ° C and 76 % relative humidity from 23 November to 31 December 1960 produced adult flies (2 ♀, 5 ♂) between 11 and 18 January 1961. The longest puparial period, 263 days, was that of a puparium formed in the laboratory on 28 July 1960; it produced 1 ♀ on 17 April 1961. Duration of the nonquiescent puparial period of summer generations of C. pectoralis was about 11 days. Among field-collected adult flies held in the laboratory, 1 ♂ lived 68 days and 1 ♀ lived 70 days, but the others lived only 6 – 28 days.	en	Bratt, Albertus D., Knutson, Lloyd V., Murphy, William L., Daniels, Anthony A. (2020): Biology, immature stages, and systematics of snail-killing flies of the genus Colobaea (Diptera: Sciomyzidae), with overviews of aspects of the tribe Sciomyzini. Zootaxa 4840 (1): 1-64, DOI: 10.11646/zootaxa.4840.1.1
7170D74F6A1EFFCAFF4FFE898FFCDA79.taxon	description	Map 15 (BNN 6411; FT 6411, 6416, 6419 – 6420) Adult flies of Colobaea punctata are very similar to those of C. pectoralis but differ noticeably in the extent of black pigmentation on the pleura and in characters of the male postabdomen. Adult flies of C. punctata have a discrete black spot below the anterior notopleural seta, whereas those of C. pectoralis have a black stripe extending along the upper margin of the mesopleuron to the base of the wing (see photograph at https: // upload. wikimedia. org / wikipedia / commons / c / c 7 / Colobaea _ punctata. jpg). Rozkošný (1959) and Rivosecchi (1992) both provided a figure of an adult female.	en	Bratt, Albertus D., Knutson, Lloyd V., Murphy, William L., Daniels, Anthony A. (2020): Biology, immature stages, and systematics of snail-killing flies of the genus Colobaea (Diptera: Sciomyzidae), with overviews of aspects of the tribe Sciomyzini. Zootaxa 4840 (1): 1-64, DOI: 10.11646/zootaxa.4840.1.1
7170D74F6A1EFFCAFF4FFE898FFCDA79.taxon	materials_examined	Type locality. Rozkošný (1984 b) noted, “ The type-series [in MZLU] consists of 10 males and 10 females, but only four of them have locality labels (Donse and Damhusmosen in Denmark) ” [bogs near Hillerød and Copenhagen, Zealand, Denmark]. Although C. punctata seems to have about the same northern limits (Jakobstad, Finland) as the closely related C. pectoralis, it extends much farther south, with a southeastern extension that is fairly continuous to Lake Ohrid, which lies between North Macedonia and eastern Albania. We have seen 1 ♂ of C. punctata from Turkey (Finike, Antalya); 1 ♀ from the northern Negev, Israel; 1 ♂ from Iran (Mazanderan, 25 km west of Gorgan); 2 ♀ and 5 ♂ from Atvasar, Kazakhstan; and 1 ♂ from Bala Murghab Herat, northeastern Afghanistan (Rozkošný & Knutson 2006). Rozkošný & Elberg (1991) recorded the species from “ eastern Siberia ” without providing a specific locality. LVK reared C. punctata in southeastern Spain (Adra, near Almeria). The record from Almeria (37 ° N) on the Mediterranean Sea is about 1100 km south of the previously known southernmost locality (Montpellier, France; 43.5 ° N) in western Europe. Becker (1903) recorded C. pectoralis from Cairo, Egypt, but that was before C. punctata had been described; thus his record may have been based on a specimen of the latter species.	en	Bratt, Albertus D., Knutson, Lloyd V., Murphy, William L., Daniels, Anthony A. (2020): Biology, immature stages, and systematics of snail-killing flies of the genus Colobaea (Diptera: Sciomyzidae), with overviews of aspects of the tribe Sciomyzini. Zootaxa 4840 (1): 1-64, DOI: 10.11646/zootaxa.4840.1.1
7170D74F6A1EFFCAFF4FFE898FFCDA79.taxon	discussion	Laboratory rearings and observations are based on adult flies reared from puparia found in shells of P. planorbis collected between 15 February and 20 March 1964 at Adra, Spain.	en	Bratt, Albertus D., Knutson, Lloyd V., Murphy, William L., Daniels, Anthony A. (2020): Biology, immature stages, and systematics of snail-killing flies of the genus Colobaea (Diptera: Sciomyzidae), with overviews of aspects of the tribe Sciomyzini. Zootaxa 4840 (1): 1-64, DOI: 10.11646/zootaxa.4840.1.1
7170D74F6A1EFFCAFF4FFE898FFCDA79.taxon	distribution	The habitat distribution of the adult flies seems to be quite broad. Adult flies were collected along the partly shaded margin of a small lake (Wienerwald See) near Vienna, Austria (C. O. Berg in litt. to LVK). Coe (1958) recorded 1 ♂ “ swept from vegetation by lake ” at Lake Ohrid. Specimens in the ZMUC were collected in bogs in Zealand, Denmark, and at a millrace pond in Jutland, Denmark. Lundbeck (1923) found puparia in shells of several freshwater snails floating in spring flood refuse in Danish ponds and bogs. ADB and LVK discovered puparia in many shells of P. planorbis found floating among Carex, Juncus, and hygrophilous grasses emerging through 10 – 50 cm of water along the exposed shore of a small lake in the Albufera de Adra at Adra, Spain.	en	Bratt, Albertus D., Knutson, Lloyd V., Murphy, William L., Daniels, Anthony A. (2020): Biology, immature stages, and systematics of snail-killing flies of the genus Colobaea (Diptera: Sciomyzidae), with overviews of aspects of the tribe Sciomyzini. Zootaxa 4840 (1): 1-64, DOI: 10.11646/zootaxa.4840.1.1
7170D74F6A1EFFCAFF4FFE898FFCDA79.taxon	materials_examined	Adult flies mated within 24 hours after emergence from puparia and continued to mate from 9 March until 14 May, throughout the entire time during which the cultures of flies were maintained. The copulatory posture of the adult flies was found to be typical of that of most Sciomyzidae [described above under “ Colobaea bifasciella (Fallén), 1820 ”] and not unusual as in the closely related C. pectoralis. Even very gravid females were observed to copulate for long periods of time. Females that emerged between 29 February and 18 April first oviposited as early as four days after emergence and at the latest within 17 days after emergence. They laid eggs between 14 March and 26 May on damp or wet vegetation or cotton in the breeding jars but laid no eggs on living P. planorbis that were also kept in the jars. Eggs hatched between 16 March and 30 May after incubation periods of 2 – 4 days at 19 – 21 ° C.	en	Bratt, Albertus D., Knutson, Lloyd V., Murphy, William L., Daniels, Anthony A. (2020): Biology, immature stages, and systematics of snail-killing flies of the genus Colobaea (Diptera: Sciomyzidae), with overviews of aspects of the tribe Sciomyzini. Zootaxa 4840 (1): 1-64, DOI: 10.11646/zootaxa.4840.1.1
7170D74F6A1EFFCAFF4FFE898FFCDA79.taxon	discussion	Of the five reared species of Colobaea, C. punctata appears to have the broadest food range. Lundbeck (1923) wrote, “ This species I have bred from several snails, most often from young specimens of Planorbis planorbis L., and further from P. [Planorbis] albus Müll. and from youngs (sic) of P. [Planorbarius] corneus L. and also from Lymnaea ovata Drap. var. peregra [L. ovata = Radix balthica (L.)]. ” In contrast to Lundberg’s experience, all puparia found at Adra were in shells of P. planorbis. Small individuals of other species of aquatic snails [Galba truncatula, Physella acuta (Draparnaud), and R. peregra] were abundant at Adra, but no puparia of C. punctata were found in shells of those species. During laboratory rearings, however, many C. punctata larvae killed, consumed, and developed to pupariation solely on those species of snails. Larvae also killed and consumed A. vortex, B. contortus, P. planorbis, Segmentina nitida (O. F. Müller), and S. palustris. A few larvae attacked but did not kill small specimens of Succinea putris L. from Denmark. They did not attack “ Phystra myosotis ” [= Myosotella myosotis (Draparnaud)] from Valencia, Spain, nor did any first-instar larvae initially feed on dead snails. Newly hatched larvae crawled about actively and readily attacked snails. The methods of attack and feeding were similar for all species of snails consumed, but most of the detailed observations were made with small (about 5 - mm-diam.) P. planorbis and small (about 4 mm long) P. acuta. Each snail observed during the rearings was attacked by a single first-instar larva; in crowded rearing containers, as many as 15 first-instar larvae would attack a single snail. It appeared that the young larvae fed on mucus, hemolymph, or extrapallial fluid rather than on snail tissues. An hour or so after the initial contact, depending upon the size of the host and the number of larvae that attacked it, the snail gradually began to retreat into its shell. The larvae followed, feeding at the edges of the mantle and foot. By the second day of feeding, the quiescent snail had withdrawn completely into its shell, and the larvae had penetrated between the mantle and shell until only their posterior spiracles remained exposed. A 5 - mm-diam. P. planorbis that had been attacked by 10 first-instar larvae lived for three days after the initial attack. A 3 - mm-long P. acuta lived for five days after being attacked by three first-instar larvae. Larvae continued to feed in the decaying tissues even after the tissues had become black and putrid until most of the contents of the shells had been consumed. Larval feeding appeared to be unaffected by the large populations of nematodes that often developed in the putrid snails. Older larvae attacked vigorously, immediately penetrated between the shell and the mantle in the acute corner of the aperture of P. acuta, and killed the snail within a few hours. The larvae killed and partially consumed a second and even a third snail if the first snail had not provided sufficient nourishment for completion of larval development. The total duration of larval life for 19 individuals was 9 – 16 days (average 10.3 days) at 19 – 22 ° C. Larvae of C. punctata were reared to pupariation on P. planorbis, P. acuta, and S. palustris. Most larvae pupariated within shells even if copious liquefied tissue remained in the aperture. Only one puparium was formed in each shell. A few larvae pupariated on or under damp cotton or moss in the rearing containers. Shells of P. planorbis collected at Adra that contained puparia were 3.3 – 7.2 mm in diameter. Pupariating larvae excreted a small quantity of septum material, but on no occasion was the material formed into even a rudimentary septum as with C. americana and some species of Pherbellia. In the laboratory, 24 puparia were formed between 28 March and 25 April 1964, from which 19 adult flies (10 ♀, 9 ♂) emerged between 12 April and 12 May after puparial periods of 15 – 20 days (average 17.4 days) at 19 – 22 ° C. Colobaea punctata apparently overwinters in the puparium. Lundbeck (1923) reared adult flies from puparia collected in the spring in Denmark. From 152 puparia that LVK collected between 15 February and 20 March 1964 in southern Spain in shells of P. planorbis, 99 adult flies (46 ♀, 53 ♂) emerged between 29 February and 17 June. Females emerged between 9 March and 18 April, 5 – 45 days (average 16.3 days) after the puparia had been collected, and males emerged between 29 February and 17 July, 6 – 89 days (average 18.2 days) after the puparia had been collected. LVK encountered neither adult flies nor other immature stages at Adra even when great numbers of puparia were present.	en	Bratt, Albertus D., Knutson, Lloyd V., Murphy, William L., Daniels, Anthony A. (2020): Biology, immature stages, and systematics of snail-killing flies of the genus Colobaea (Diptera: Sciomyzidae), with overviews of aspects of the tribe Sciomyzini. Zootaxa 4840 (1): 1-64, DOI: 10.11646/zootaxa.4840.1.1
7170D74F6A1EFFCAFF4FFE898FFCDA79.taxon	description	Of 40 laboratory-reared adult flies, 19 ♂ lived 4 – 26 days (average 14.0 days) and 21 ♀ lived 7 – 31 days (average 15.0 days).	en	Bratt, Albertus D., Knutson, Lloyd V., Murphy, William L., Daniels, Anthony A. (2020): Biology, immature stages, and systematics of snail-killing flies of the genus Colobaea (Diptera: Sciomyzidae), with overviews of aspects of the tribe Sciomyzini. Zootaxa 4840 (1): 1-64, DOI: 10.11646/zootaxa.4840.1.1
7170D74F6A1EFFCAFF4FFE898FFCDA79.taxon	discussion	Data from capture records of adult flies extend from 8 April (Finike, Antalya, Turkey) to 26 September (Berlin area; Kassebeer 2001). In his study of the flight periods and voltinism of the Sciomyzidae as based on museum material, Soós (1958) recorded 18 specimens of C. punctata collected between 15 July and 17 September. He concluded that C. punctata is a species “ die sicher nur einer Generation besitzt ” [“ which surely has only one generation ”]. Lundbeck (1923) wrote that he “ reared a Cryptine and a Chalcidid from Ctenulus punctatus, in all cases a single parasite from each pupa. ” Sixteen of 152 puparia collected at Adra produced a single parasitic wasp (Hymenoptera: Chalcididae). The wasps emerged between 19 March and 26 May, 15 – 67 days after the C. punctata puparia harboring them had been collected.	en	Bratt, Albertus D., Knutson, Lloyd V., Murphy, William L., Daniels, Anthony A. (2020): Biology, immature stages, and systematics of snail-killing flies of the genus Colobaea (Diptera: Sciomyzidae), with overviews of aspects of the tribe Sciomyzini. Zootaxa 4840 (1): 1-64, DOI: 10.11646/zootaxa.4840.1.1
7170D74F6A1FFFCAFF4FF9078AD2DBA3.taxon	discussion	1. Colobaea sp. near C. canadensis. The DEBU collection contains 1 ♂ in perfect condition (entire specimen removed from point and stored in genitalia vial, with postabdomen in separate genitalia vial, in litt. 22 Oct 2019, Stephen A. Marshall to WLM) of an undescribed species labeled “ USSR, Siberia, Novosibirsk Region, Cherny Mts., ca. 20, flt / pans lakeshore, 20 – 29 June 1991, S. A. Marshall. ” 2. Colobaea sp. near C. eos. The ZMUC contains 1 ♀ in perfect condition, labeled “ India (Uttar Pradesh), Mussoorie [northeast of Delhi], c. 1500 – 2300 m., 3 – 14 August 1978, Copenhagen Zool. Mus. Exp. ”	en	Bratt, Albertus D., Knutson, Lloyd V., Murphy, William L., Daniels, Anthony A. (2020): Biology, immature stages, and systematics of snail-killing flies of the genus Colobaea (Diptera: Sciomyzidae), with overviews of aspects of the tribe Sciomyzini. Zootaxa 4840 (1): 1-64, DOI: 10.11646/zootaxa.4840.1.1
7170D74F6A27FFF1FF4FFA158DCBD9A8.taxon	description	First-instar larva (based on 10 specimens from Geneva, New York, USA). Length 0.70 – 1.00 mm; greatest width 0.15 – 0.20 mm. Basal rings of sensory papillae large, lightly pigmented; post-oral and segmental spinules lightly pigmented. Segmental spinules and spinules on posterior end small and sparse. Cephalopharyngeal skeleton light brown, cornua slender, mouthhooks long, slender. Length 0.08 – 0.09 mm. Indentation index 72 – 81. Posterior end with three pairs of marginal lobes; spiracular tubes 0.015 mm long. Float hairs long, slender. Second-instar larva (based on five specimens from Geneva, New York, USA). Length 1.2 – 2.0 mm long; greatest width 0.2 – 0.4 mm. Anterior spiracles with 12 – 14 marginal papillae, bifid, papillae tightly covered by integument. Cephalopharyngeal skeleton 0.11 mm long. Mouthhooks contiguous apically, widened to articulation with hypostomal sclerite. Indentation index 86. No spinule band on segment III. Lateral pads with spinules. Posterior spiracular disc with three pairs of peripheral lobes (Fig. 13); dorsal lobes very small, ventral and ventrolateral lobes appear as welts, subequal; lateral protuberances small. No mammillae visible above ventral lobes. Three groups of hyaline float hairs at margin of spiracular plates. Third-instar larva (Fig. 31) (based on 10 specimens from Geneva, New York, USA). Length 1.5 – 3.0 mm; greatest width 0.3 – 1.0 mm. Body tapering, anterior half slender, posterior part larger, dorsoventrally compressed, narrowing abruptly to small posterior spiracular disc (Fig. 49). Lateral welts large, obscuring small lateral pads. Anterior spiracles bifid, no prismatic spots, 14 – 16 marginal papillae, integument not tightly enclosing each papilla (Fig. 8). Cephalopharyngeal skeleton (Fig. 38) 0.26 mm long. Indentation index 80. Dorsal and ventral cornua slender. Mouthhooks long, slender, ventrally directed. Ventral arch with 25 – 30 teeth. Dorsal cornu long, slender with elongate, complete window; ventral cornu narrow with small, complete window. Margins of cornua entire. Spinules very sparse and lightly pigmented, segment III without ventral spinule patch, segment IV with undivided ventral spinule patch, segment V with two-part patch, segments VI – XII with three-part patches. No dorsal spinule patches or scales. Lateral pads with spinules. Posterior end with four pairs of lobes (Fig. 15). Ventral and ventrolateral marginal lobes large, subequal; lateral protuberances large, rounded; dorsal lobes very small. Mammillae at bases of ventral lobes. Small, sparse spinules on lobes and outer part of disc. Peritremes, slit margins, and stigmatic scars lightly pigmented; margin of peritreme at outer end of each spiracular slit with low, spinelike process (Fig. 49). Puparium (Figs 25, 44) (based on 10 specimens from Geneva, New York, USA). Length 2.0 – 3.0 mm; greatest width 0.7 – 1.1 mm. Formed to fit inside snail shell. Body form robust. First three segments not contracted but attenuated and laterally expanded to occlude snail shell aperture; next five segments contracted, dorsally depressed by shell whorl; last segments abruptly narrowed to small posterior spiracular disc. Integument subshiny, finely wrinkled with minute punctations, thick. Anterior spiracles and dorsal cephalic cap blackish brown; remainder dark reddish brown. Primary integumentary folds apparent; secondary integumentary folds obscure; lateral intersegmental pads visible. Ventral spinule patches apparent, spinules dark. Anterior spiracles bifid, strongly protrudent from anterolateral angles of cephalic cap. Posterior spiracles protrudent, dorsally directed. Peripheral lobes of posterior end slightly reduced.	en	Bratt, Albertus D., Knutson, Lloyd V., Murphy, William L., Daniels, Anthony A. (2020): Biology, immature stages, and systematics of snail-killing flies of the genus Colobaea (Diptera: Sciomyzidae), with overviews of aspects of the tribe Sciomyzini. Zootaxa 4840 (1): 1-64, DOI: 10.11646/zootaxa.4840.1.1
7170D74F6A24FFF0FF4FFA5F8C25DAC4.taxon	description	First-instar larva (based on three specimens, ex egg from Brattby, Sweden). Length 0.65 – 1.00 mm; greatest width 0.16 – 0.30 mm. Basal ring of sensory papillae darkly pigmented; spinules of post-oral spinule patch large, numerous, darkly pigmented. Three large, unicuspid spinules on each lateral surface of mouth opening, contiguous to paired sclerites below mouthhooks. Mouthhooks long, slender; cephalopharyngeal skeleton light brown, 0.17 – 0.20 mm long. Indentation index 87. Posterior end with small ventral and ventrolateral lobes. Spiracular tubes 0.02 mm long, protrudent, robustly cylindrical. Hyaline float hairs short. Third-instar larva (Fig. 21) (based on 10 specimens from Hillerød, Denmark, and Brattby, Sweden). Length 3.0 – 6.5 mm; greatest width 1.0 – 2.5 mm. Body form dorsoventrally compressed on posterior half; lateral welts apparent, obscuring lateral pads. Anterior spiracles large (Fig. 50), 14 – 20 marginal papillae, integument closely enclosing each papilla; disc of spiracle with scattered prismatic spots (Fig. 35). Cephalopharyngeal skeleton (Fig. 42) 0.50 mm long. Indentation index 90. Ventral cornu large, with irregularly pigmented posterior margin; dorsal cornu with elongate, incomplete window; dorsal bridge small. Hypostomal sclerite small. Mouthhooks large, anteriorly directed; ventral arch small, reduced to lightly pigmented strap, with 20 – 22 long, slender teeth. Spinules more numerous and more darkly pigmented than in other species of Colobaea. Segment III without ventral spinule patch, segment IV with single ventral spinule patch, segments V and XII with divided patches, segments VI – XI with three-part patches. No spinules on lateral pads. Posterior third of body with fine scales dorsally and laterally; segments XI and XII with dense scales and spinules within dorsal pigmented areas, integument in those areas thickened and tanned (Fig. 22). Posterior spiracles strongly protrudent from small posterior spiracular disc. Two pairs of peripheral lobes (ventral and ventrolateral) on disc, both small, not acute, appearing as welts; no mammillae at bases of ventral lobes (Fig. 16); 3 – 4 rows of small, dark spinules on lobes and adjacent areas. Puparium (Figs 4, 24, 27) (based on 10 specimens from Hillerød, Denmark). Length 4.3 – 5.0 mm; greatest width 2.0 – 2.5 mm. Shaped to fit inside snail shell, dorsal part of first six segments occluding aperture of shell, posterior segments twisted. Robustly spindle shaped in lateral aspect, first six segments strongly convex with longitudinal groove formed by pressure from the columella of the snail shell, segments XI and XII upturned. Anterior half dark reddish brown, integument strongly sclerotized; posterior half yellowish brown, integument thin. Integument subshiny, with fine punctations and wrinkles. Segmentation conspicuous; lateral intersegmental pads obscure; lateral longitudinal welts apparent. Spinule patches obscure. Anterior spiracles large, projecting dorsally from angles of cephalic cap; posterior spiracles protruding weakly. Lobes on posterior end reduced.	en	Bratt, Albertus D., Knutson, Lloyd V., Murphy, William L., Daniels, Anthony A. (2020): Biology, immature stages, and systematics of snail-killing flies of the genus Colobaea (Diptera: Sciomyzidae), with overviews of aspects of the tribe Sciomyzini. Zootaxa 4840 (1): 1-64, DOI: 10.11646/zootaxa.4840.1.1
7170D74F6A25FFFFFF4FF8838D7DDA79.taxon	description	First-instar larva (based on 10 specimens from Dezful, Iran). Length 0.50 – 0.70 mm; greatest width 0.07 – 0.14 mm. Post-oral spinules very long, slender, recurved, lightly pigmented. Cephalopharyngeal skeleton light brown, 0.09 – 0.10 mm long. Indentation index 82 – 88. Mouthhooks long, slender; pharyngeal sclerites strongly sclerotized. Posterior end with three pairs of large marginal lobes plus lateral protuberances. Large, darkly pigmented spinules below spiracular plates (Fig. 29). Spiracular tubes 0.017 mm long, slender. Float hairs long, slender. Peri-anal pad large, protrudent, subovate, with apices attenuated. Second-instar larva (based on four specimens from Dezful, Iran). Length 1.6 – 1.8 mm; greatest width 0.2 – 0.3 mm. Body form slender, spindle shaped. Post-oral spinules numerous, large, sharp; cephalopharyngeal skeleton (Fig. 12) dark brown; 0.20 mm long. Indentation index 76 – 80. Pharyngeal sclerites short; dorsal and ventral cornua with incomplete windows; mouthhooks long, with small, recurved hook part; ventral arch small, straplike, with 18 – 20 small, slender teeth. Anterior spiracles small, with 12 – 15 marginal papillae. Spinules present on lateral pads; segmental spinules sparse, lightly pigmented. Posterior end with three pairs of large marginal lobes plus lateral protuberances. Mammillae present at bases of ventral lobes. Peritremes lightly pigmented, three spiracular slits in central, pigmented area; no hyaline float hairs (Fig. 14). Third-instar larva (based on 10 specimens from Dezful, Iran). Length 2.2 – 3.5 mm; greatest width 0.4 – 0.9 mm. Body form spindle shaped, lateral welts not apparent, posterior half of body not expanded, lateral pads visible. Anterior spiracles (Fig. 33) with expanded apical part, 12 – 14 marginal papillae tightly enclosed by integument. Cephalopharyngeal skeleton (Fig. 43) dark brown, 0.34 mm long. Indentation index 80. Mouthhooks large, hook part directed ventrally. Ventral arch slender, with 18 – 20 teeth. Several rows of dark spinules by oral opening, lateral to mouthhooks. Dorsal cornu with small, complete window; ventral cornu with large, elongate window; apparent margins of cornua entire. Spinules present on lateral pads; segments III and IV with ventral spinule patch, segment V with two-part patch, segments VI – XII with three-part patches. Segmental spinules small, lightly pigmented. Posterior end (Fig. 19) with four pairs of lobes; dorsal pair small, ventral and ventrolateral pairs large, conical; lateral protuberances large, less acute than lobes. Mammillae present at bases of ventral lobes. Peritremes, slit margins, and stigmatic scars lightly pigmented (Fig. 30). Peri-anal pad with lateral margins attenuated into points. Puparium (Figs 45 – 46) (based on six specimens from Dezful, Iran). Length 4.1 – 5.0 mm; greatest width 1.0 – 1.1 mm. Typically distorted to fit planorbiform snail host shell. Dorsal surface of segments II – IV dark, reddish brown if host shell is planorbiform or else all exposed surfaces darkly pigmented; other integument tan. Integument subshiny, finely punctate. Segmentation, secondary integumentary folds, and spinules obscure. Lateral pads and lateral longitudinal welts visible on segments IX – XI. In dorsal aspect, dorsal cephalic cap flattened and expanded to occlude whorl of host shell; widest at posterior margin of segment V and then narrowed abruptly and continued with margins parallel to those of segment XI, which narrows to rounded end of posterior spiracular disc. Crescent shaped in lateral aspect, segments VI – IX depressed by inner whorl of host shell. Anterior spiracles small, projecting dorsally from anterolateral angles of cephalic cap; papillae very small. Posterior spiracles large, protrudent from dome-shaped disc; marginal lobes small, pointed. Posterior spiracular disc not darkened. Peri-anal pad suboval, concolorous, not invaginated.	en	Bratt, Albertus D., Knutson, Lloyd V., Murphy, William L., Daniels, Anthony A. (2020): Biology, immature stages, and systematics of snail-killing flies of the genus Colobaea (Diptera: Sciomyzidae), with overviews of aspects of the tribe Sciomyzini. Zootaxa 4840 (1): 1-64, DOI: 10.11646/zootaxa.4840.1.1
7170D74F6A2AFFFEFF4FF91D8B2DDCE1.taxon	discussion	We cite here an English translation (from the original German) of Rozkošný’s (1967) description of the puparium and third-instar cephalopharyngeal skeleton based in part on a puparium found in a shell of Anisus spirorbis collected 3 April 1965 in Vranovice, southern Czech Republic, from which a female emerged on 16 April:	en	Bratt, Albertus D., Knutson, Lloyd V., Murphy, William L., Daniels, Anthony A. (2020): Biology, immature stages, and systematics of snail-killing flies of the genus Colobaea (Diptera: Sciomyzidae), with overviews of aspects of the tribe Sciomyzini. Zootaxa 4840 (1): 1-64, DOI: 10.11646/zootaxa.4840.1.1
7170D74F6A2AFFFEFF4FF91D8B2DDCE1.taxon	description	“ Puparium (length 3.6 – 4.2 mm, width 0.8 – 1.0 mm) red-brown, anterior end blackish. Surface smooth. Original segmentation especially dorsally very indistinct. Cuticular spinules on the ventral side very tiny, only indistinctly visible through strong magnifying glass. Posterior spiracles on narrow stigmatic tubes. Remains of anterior spiracles at the sides of bilobed anterior cornua. Number of the spiracular slits on the anterior spiracles 15 – 17. CP skeleton (length 0.39 mm) with relatively massive mouthhooks, ventral arch with 22 – 24 long teeth, pharyngeal sclerite with divergent cornua and large, longitudinal oval windows in each cornu. ” The descriptions and figures were reproduced in Rozkošný (2002) and the figures in Vala (1989).	en	Bratt, Albertus D., Knutson, Lloyd V., Murphy, William L., Daniels, Anthony A. (2020): Biology, immature stages, and systematics of snail-killing flies of the genus Colobaea (Diptera: Sciomyzidae), with overviews of aspects of the tribe Sciomyzini. Zootaxa 4840 (1): 1-64, DOI: 10.11646/zootaxa.4840.1.1
7170D74F6A2BFFFDFF4FFEE28ECCDADD.taxon	description	First-instar larva (based on five specimens from Hillerød, Denmark). Length 1.00 – 1.20 mm; greatest width 0.10 – 0.20 mm. Basal rings of sensory papillae dark, large; post-oral spinules long, slender, lightly pigmented. Cephalopharyngeal skeleton (Fig. 9) brown, 0.10 – 0.11 mm long. Indentation index 80; mouthhooks short, broad. Posterior end with three pairs of marginal lobes and lateral protuberances; spinules sparse, lightly pigmented. Spiracular tubes long, cylindrical, 0.14 mm long. Float hairs long, slender. Second-instar larva (based on three specimens from Hillerød, Denmark). Length 1.5 – 2.5 mm; greatest width 0.3 – 0.5 mm. Anterior spiracles small, stalk narrow, 16 – 18 marginal papillae tightly enclosed by integument. Cephalopharyngeal skeleton (Fig. 10) 0.14 mm long. Indentation index 87. Ventral arch with 18 – 20 teeth. Spinules present on lateral pads. Segment III without ventral patch. Posterior end with four pairs of lobes; dorsal pair very small; lateral protuberances large, blunt, anterior to disc; ventral and ventrolateral pairs about half the size of lateral pair, rounded at apices. Mammillae present at bases of ventral lobes. No float hairs. Third-instar larva (based on 10 specimens from Hillerød, Denmark). Length 4.2 – 5.5 mm; greatest width 0.8 – 1.2 mm. Body form slender, spindle shaped, circular in cross section, not flattened and expanded posteriorly. Anterior spiracles with 14 – 16 marginal papillae, without prismatic spots, papillae not tightly covered by integument (Fig. 32). Cephalopharyngeal skeleton 0.35 mm long (Figs 36 – 37, 41). Dorsal cornu large, with small, incomplete window; ventral cornu slender, elongate, with small, incomplete window; dorsal bridge prominent. Indentation index 75. Mouthhooks long, anteroventrally directed; ventral arch large, with 20 – 22 teeth. Spinules sparse, lightly pigmented. Segment III with no spinule patch, segment IV with single ventral patch, segment V with two-part patch, segments VI – XII with three-part patches. Lateral pads with spinules. No dorsal spinules. Posterior end with four pairs of lobes (Fig. 17). Ventral and ventrolateral pairs large, subequal; lateral protuberances at anterior margin of disc large, dorsal pair very small. Mammillae at bases of ventral lobes. Ventral and ventro-lateral lobes and lateral protuberances with 3 – 4 rows of encircling spinules. Peritremes, slit margins, and stigmatic scars darkly pigmented. Puparium (Figs 23, 26) (based on 10 specimens from Hillerød, Denmark). Length 4.9 – 6.2 mm; greatest width 1.4 – 1.6 mm. Shaped to fit inside snail shell. Crescent shaped in lateral aspect. In dorsal aspect, first three segments widen from apex of cephalic caps, next six segments parallel sided, last segments narrowed abruptly to small posterior spiracular disc. Dorsal cephalic cap concave, next two segments convex, next four segments dorsally depressed by shell, last segments convex. First four segments dark reddish brown, last two segments lighter, middle segments lightly pigmented; dorsal surface lighter than ventral. Integument thick, finely wrinkled, with light punctations. Segmentation conspicuous, lateral intersegmental pads apparent, secondary integumentary folds not apparent. Anterior spiracles strongly protrudent anterodorsally from angles of cephalic cap. Posterior spiracular tubes protrudent. Posterior spiracular disc distorted, reduced; ventral and ventrolateral lobes visible as small welts, other lobes not apparent. Lundbeck (1923) illustrated and provided a brief description of the puparium. For comparison with our descriptions, we cite here an English translation (from the original German) of Rozkošný’s (1967) description of the puparium and third-instar cephalopharyngeal skeleton of C. pectoralis, based on three puparia found in shells of A. vortex collected 25 March 1965, from which 2 ♂ and 1 ♀ emerged on 13 April: “ Puparium (Dimensions: 2.8 – 3.3 x 0.7 – 1.0 mm) red brown-brown. Anterior end of puparium is wide and expanded, differing from Colobaea distincta. The anterior spiracles lie at the sides of the anterior end, shaped longitudinally oval with tiny spiracular slits (about 15). Under the posterior spiracles, the remains of 2 pairs of marginal lobes are easily visible. Shape of the puparium longitudinal cylindrical adapted to the shell shape of Anisus vortex. CP skeleton (length 0.35 mm) very similar to the one of Colobaea distincta. Mouthhooks not so large. Both cornua of the pharyngeal sclerites a little less divergent, anterior bridge well developed. ” Rozkošný’s (1967) descriptions and figures were reproduced in Rozkošný (2002) and the figures in Vala (1989).	en	Bratt, Albertus D., Knutson, Lloyd V., Murphy, William L., Daniels, Anthony A. (2020): Biology, immature stages, and systematics of snail-killing flies of the genus Colobaea (Diptera: Sciomyzidae), with overviews of aspects of the tribe Sciomyzini. Zootaxa 4840 (1): 1-64, DOI: 10.11646/zootaxa.4840.1.1
7170D74F6A28FFF9FF4FF8BE8CD7DF0D.taxon	description	Egg (based on four specimens from Adra, Spain). Length 0.60 – 0.80 mm; greatest width 0.20 – 0.30 mm. Robust, anterior end smoothly rounded, dorsal surface slightly convex in lateral aspect, ventral surface strongly convex. Chorion finely punctate with coarse, reticulate pattern. First-instar larva (based on 10 specimens from Adra, Spain). Length 0.60 – 0.96 mm; greatest width 0.12 – 0.20 mm. Basal rings of sensory papillae lightly pigmented; post-oral spinules long, slender, lightly pigmented. Segmental spinules sparse, lightly pigmented. Cephalopharyngeal skeleton brown, 0.08 – 0.09 mm long. Indentation index 72 – 81. Mouthhooks short, broad. Posterior end with three pairs of marginal lobes and lateral protuberances; spinules on lobes small, lightly pigmented. Spiracular tubes 0.014 mm long. Float hairs short, slender. Second-instar larva (based on 10 specimens from Adra, Spain). Length 1.0 – 2.0 mm; greatest width 0.3 – 0.5 mm. Anterior spiracles small, apical part slightly expanded, subcircular, 14 – 16 marginal papillae. Cephalopharyngeal skeleton (Fig. 11) 0.17 mm long. Indentation index 90. Mouthhooks long, slender; hook part large, sharp. Ventral arch with 18 – 20 teeth. Dorsal cornu with small, incomplete window; ventral cornu with large, complete window. Segments III and IV with undivided ventral spinule patch. No spinules on lateral pads. Posterior end with four pairs of lobes. Lateral pair largest, apices rounded, projecting anterior to margin of disc; ventral and ventrolateral pairs subequal, apices rounded, about half the size of lateral pair; dorsal pair very small. Mammillae present above bases of ventral lobes. No float hairs at margins of spiracular plates. Third-instar larva (based on five specimens from Adra, Spain). Length 2.8 – 5.0 mm; greatest width 0.8 – 1.2 mm. Body form slender, spindle shaped, circular in cross section, not expanded or flattened posteriorly. Anterior spiracles with 14 – 16 marginal papillae, no prismatic spots, integument not closely enclosing each papilla (Fig. 34). Cephalopharyngeal skeleton (Fig. 39 – 40) 0.38 mm long. Indentation index 83. Dorsal cornu of pharyngeal sclerite with large, elongate, complete window; ventral cornu with large, complete window; dorsal bridge prominent; mouthhooks large, ventrally directed. Ventral arch with 18 – 20 teeth. Spinules sparse, not pigmented. Segments III and IV with ventral spinule patch, segment V with two-part patch, segments VI – XII with three-part patches. Lateral pads with spinules. No spinules or scales dorsally. Posterior end with four pairs of lobes (Fig. 18). Ventral and ventrolateral lobes large, subequal; lateral protuberances at anterior margin of disc (Fig. 20); dorsal lobes very small. Mammillae at bases of ventral and ventrolateral lobes. Ventral and ventrolateral lobes and lateral protuberances with 3 – 4 rings of spinules. Peritremes, slit margins, and stigmatic scars darkly pigmented. Puparium (Figs 47 – 48) (based on 10 specimens from Adra, Spain). Length 2.7 – 3.7 mm; greatest width 1.3 – 1.5 mm Shaped to fit inside snail shells. Dorsal surface of first six segments flat and occluding shell, ventral surface of first six segments convex and appressed to outer surface of shell whorl, dorsal surface of last segments concave and appressed to inner surface of shell whorl, ventral surface of last segments not contacting shell. Body form robustly spindle shaped, widest in one-third to one-half distance from anterior end, narrowed to small posterior spiracular disc. Anterior half of dorsal surface darkly pigmented, dark reddish brown; posterior half of dorsal surface light reddish brown. Integument of dorsal surface of first six segments thick, that of posterior surface thin, fragile, finely granular, wrinkled at segmental folds. Secondary integumentary folds apparent dorsally. Primary integumentary folds conspicuous on anterior half, less obvious on posterior half. Lateral intersegmental pads obscure, lateral longitudinal welts apparent. Anterior spiracles small, protruding from dorsal surface of cephalic cap near angles; posterior spiracles lightly pigmented, protrudent. Lobes of posterior end slightly reduced and distorted. Lundbeck (1923) described but did not figure the puparium of his new species C. punctata; however, he emphasized the differences in shape of puparia found in Planorbarius corneus, Planorbis albus, Planorbis planorbis, and “ L. ovata var. peregra ” [= Radix peregra] from the shape of puparia of C. pectoralis found only in Planorbis vortex. He then expanded upon his 13 - line original description of the adult flies of C. pectoralis and C. punctata, noting that, “ the specimens in our old collection of C. pectoralis, which Zetterstedt had seen, I found them to be the present species — I think he has had both species [Denmark, Sweden] before him. ” It is interesting to note that Lundbeck was motivated to describe his new species C. punctata (based on adult flies) because of results of his initial comparisons of immature stages of C. pectoralis and C. punctata, which reinforces our temerity in describing a new species of Colobaea based partly on immature stages.	en	Bratt, Albertus D., Knutson, Lloyd V., Murphy, William L., Daniels, Anthony A. (2020): Biology, immature stages, and systematics of snail-killing flies of the genus Colobaea (Diptera: Sciomyzidae), with overviews of aspects of the tribe Sciomyzini. Zootaxa 4840 (1): 1-64, DOI: 10.11646/zootaxa.4840.1.1
7170D74F6A34FFE1FF4FFF3F8BB8D945.taxon	description	The only keys to genera of larvae and puparia of Palearctic Sciomyzidae are those of Rozkošný (2002) and Knutson & Vala (2011). Rozkošný (2002) provided a key to the puparia of two species of Colobaea (C. distincta and C. pectoralis).	en	Bratt, Albertus D., Knutson, Lloyd V., Murphy, William L., Daniels, Anthony A. (2020): Biology, immature stages, and systematics of snail-killing flies of the genus Colobaea (Diptera: Sciomyzidae), with overviews of aspects of the tribe Sciomyzini. Zootaxa 4840 (1): 1-64, DOI: 10.11646/zootaxa.4840.1.1
