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Further Details of the Morphology of the Enigmatic African Fly Mormotomyia hirsuta Austen (Diptera: Mormotomyiidae)
Abstract
Mormotomyia hirsuta Austen, 1936 is one of the most extraordinary and unusual looking Diptera and was placed by E.E. Austen into a family of its own, the Mormotomyiidae, upon its discovery in 1933. Adults superficially resemble small solifugids (sun spiders), having extremely long legs that are clothed, especially in males, in very long, closely-packed brown hair-like setae. The wings are reduced to dysfunctional straps, the halteres to small nodular processes, and the eyes are greatly reduced. M. hirsuta is cavernicolous in all life stages and guanobious at least in the larval stages. The phylogenetic position of the family has long been a subject of much speculation and disagreement. Until recently M. hirsuta had only been collected on two occasions: in May 1933 and December 1948, although there have been numerous unsuccessful rediscovery attempts. The species is apparently confined to the type locality and is, therefore, widely regarded as the “rarest fly in the world”. Here we report the rediscovery of adults, larvae and puparia at the type locality, a cave-like rock fissure at Ukasi Hill, Eastern Province, Kenya, in December 2010. This rediscovery has facilitated a more thorough examination and study of the immature stages using scanning electron microscopy (SEM). This has revealed numerous microstructures not previously described by van Emden in 1950, and the larva and puparium are therefore re-described. An SEM study was conducted of the leg features of adults, specifically the form and structure of the tarsal claw and pulvillus, and these were compared to the same structures in examples of the true bat fly ectoparasitic families Nycteribiidae and Streblidae, and to the phoretic Mystacinobiidae. The basal sclerites of the wing are interpreted for the first time using SEM, the functional morphology of the larva, puparium and adult is discussed and notes are provided on the biology, development and cavernicolous habits of the species. The cuticular parts of the internal female reproductive tract are further described. They comprise a tubular vagina, two sclerotized spermathecae, paired accessory glands, and a small one-chambered sclerotized ventral receptacle. These are compared to species in the Mystacinobiidae, Sphaeroceroidea and Ephydroidea, and it is concluded that the structure of the female reproductive tract lends support to the inclusion of the Mormotomyiidae in the Ephydroidea.
... application of nuclear molecular markers, and through DNA sequencing of the mito chondrial cytochrome oxidase I gene. Details of larval and puparial morphology are presented elsewhere (Kirk-Spriggs et al. 2011 ), and molecular data bearing on the phylogenetic placement of Mormotomyiidae within the Diptera will also be presented elsewhere (Wiegmann et al. ...
... Some specimens were also preserved in 75 % ethanol. Larvae and puparia of M. hirsuta were collected from guano, and their treatment is described in Spriggs et al. (2011). The following morning the expedition returned to the site at ca 10 am, the day remaining overcast until 11 am. ...
... At this time we can only speculate on possible modes of dispersal. Mormotomyia has none of the morphological adaptations for clinging onto bats' fur exhibited in other batfly families (KirkSpriggs et al. 2011). It is unlikely, therefore, that adult Mormoto myia are phoretic on bats or birds, although confirmation of this awaits examination of captured, living bats. ...
Sixty-two years since last observed alive, Mormotomyia hirsuta Austen, the "terrible hairy fly", was found inside and outside a large, cave-like cleft boulder at the summit of Ukasi Hill in eastern Kenya, the type locality of the species. Adults were observed climbing the walls of the boulder and walking on thick layers of bat guano, in which larvae and puparia were also discovered. Large numbers of M. hirsuta were observed on and at the base of the northern side of the boulder, which at the time of capture experienced continuous shade during daylight hours. Only three individuals were observed at the southern opening, ex posed to direct sunlight and hot, dry conditions. A collection of vertebrate bones and skulls from layers of guano both inside and outside the cleft revealed several vertebrate associates, including two species of Chiroptera, Chaerephon cf. bivittatus (Heuglin) and Tadarida aegyptiaca (E. Geoffroy), which are probably the two major guano-producing species responsible for the larval breeding medium. Male-biased sexual size dimorphism was pronounced in adult M. hirsuta, with seven body-part measurements, including legs, larger by 33–61 % in males than females. Males demonstrated isometric growth while female growth was allometric. In contrast to males, female head and thorax lengths did not increase proportionally with leg length. Estimates of genetic diversity in the Ukasi population show higher than expected allelic diversity and indicate possible gene flow and frequent population bottlenecks. To promote the conservation of this endangered species, a joint effort has been initiated between the International Centre of Insect Physiology and Ecology, Nairobi and the National Museums of Kenya, Nairobi, to gazette the Ukasi hill area as a pro tected site.
... Among the Diptera, there are few species as enigmatic as the wingless, solifuge-like "terrible hairy fly", Mormotomyia hirsuta Austen [12] (Fig 1B). Formerly known only from a cave-like rock cleft in Kenya [13,14], this species was rediscovered in 2010 [15,16] at the same locality from which it was originally described and has since been found at other nearby sites [17]. These flies live in and around bats, although they have not yet been observed as attached or riding phoretically on a bat, and they do not seem to be blood or tissue feeders. ...
... McAlpine & Woodley [22], however, found no convincing similarity of Mormotomyia with sphaerocerids and heleomyzids, and these families were combined in D.K. McAlpine's concept of the family Heteromyzidae, although still classified separately by most workers [19,21].As Mormotomyia exhibits characteristics of both calyptrate and acalyptrates [14], Hennig [22] cited instead a possible position as sister group to Calyptratae. In their more recent reevaluation of these flies, Kirk-Spriggs et al. [16] noted features of the female reproductive tract consistent with inclusion in the superfamily Ephydroidea. David K. McAlpine [23] corroborated this placement based on antennal structure. ...
The schizophoran superfamily Ephydroidea (Diptera: Cyclorrhapha) includes eight families, ranging from the well-known vinegar flies (Drosophilidae) and shore flies (Ephydridae), to several small, relatively unusual groups, the phylogenetic placement of which has been particularly challenging for systematists. An extraordinary diversity in life histories, feeding habits and morphology are a hallmark of fly biology, and the Ephydroidea are no exception. Extreme specialization can lead to “orphaned” taxa with no clear evidence for their phylogenetic position. To resolve relationships among a diverse sample of Ephydroidea, including the highly modified flies in the families Braulidae and Mormotomyiidae, we conducted phylogenomic sampling. Using exon capture from Anchored Hybrid Enrichment and transcriptomics to obtain 320 orthologous nuclear genes sampled for 32 species of Ephydroidea and 11 outgroups, we evaluate a new phylogenetic hypothesis for representatives of the superfamily. These data strongly support monophyly of Ephydroidea with Ephydridae as an early branching radiation and the placement of Mormotomyiidae as a family-level lineage sister to all remaining families. We confirm placement of Cryptochetidae as sister taxon to a large clade containing both Drosophilidae and Braulidae–the latter a family of honeybee ectoparasites. Our results reaffirm that sampling of both taxa and characters is critical in hyperdiverse clades and that these factors have a major influence on phylogenomic reconstruction of the history of the schizophoran fly radiation.
... Larvae of Mormotomyiidae were collected from bat guano. It is believed that adult flies feed on the body secretions of bats [89]. ...
... Larvae of Mormotomyiidae were collected from bat guano. It is believed that adult flies feed on the body secretions of bats [89]. ...
... Larvae of Mormotomyiidae were collected from bat guano. It is believed that adult flies feed on the body secretions of bats [89]. ...
... Larvae of Mormotomyiidae were collected from bat guano. It is believed that adult flies feed on the body secretions of bats [89]. ...
... Мвинги в Кении (рис. 76) [147,254]. По-видимому, хозяевами M. hirsuta являются складчатогубые (Molossidae Gervais, 1856), однако основную часть времени эти мухи -в отличие от никтерибиид и стреблид -проводят на сводах пещеры. Никтерибииды и стреблиды имеют всесветное распространение. ...
Пособие направлено на изучение основ биологии насекомых, паразитирующих на представителях отряда рукокрылых – летучих мышах и крыланах. В пособии содержится теоретический материал по основным вопросам таксономии, морфологии, жизненного цикла и экологии паразитных, поликтенидовых и триатомовых клопов, блох-исхнопсиллид и гептопсилл, мух-гиппобоскоидей и мух-мистацинобиид. Для улучшения усвоения материала пособие сопровождается таблицами и рисунками, даются контрольные вопросы для самоподготовки и проверки знаний студентов. Предназначено для студентов и аспирантов, специализирующихся в области биологии, паразитологии, микробиологии и медицины
... The general structure of the antenna is reminiscent of Leucophenga, a somewhat plesiomorphic example of the Drosophilidae, or could be classed between the more plesiomorphic and more apomorphic taxa of the Ephydridae (Figs 84-93). The three-segmented arista, the location of the preabdominal spiracles in the pleural membrane, and, in the male, the large tergite 6 and asymmetrical sternite 6 place Mormotomyia outside the limits of the Ephydridae, but general antennal structure supports its position in the Ephydroidea suggested by Kirk-Spriggs et al. (2011). ...
. The main features of antennal segments 2 and 3 seen in the higher Diptera are described, including many that are not or inadequately covered in available publications. The following terms are introduced or clarified: for segment 2 or the pedicel—annular ridge, caestus, chin, collar, conus, distal articular surface, encircling furrow, foramen of articulation, foraminal cusp, foraminal ring, pedicellar button, pedicellar cup, rim; for segment 3 or the postpedicel—basal foramen, basal hollow, basal stem, postpedicellar pouch, sacculus, scabrous tongue, sub-basal caecum; for the stylus or arista—stylar goblet. Particular attention is given to the occurrence and position of the pedicellar button. The button is the cuticular component of a chordotonal organ, which perhaps has the role of a baroreceptor. It is present in the majority of families of Diptera, and possibly was present in the ancestral dipteran. Some generalizations about antennal structure are made, and a diagram showing the main trends in antennal evolution in the Eremoneura is provided. The general form of the antenna shows a transition from approximate radial symmetry (e.g., in Empis, Microphor, and Opetia) through to superficial bilateral symmetry (in many taxa of Eumuscomorpha), though there is usually much asymmetry in detail. More detailed descriptions and illustrations are given for selected taxa of Cyclorrhapha. The phenomenon of an additional concealed segment-like structure between segments 2 and 3, found among the Chloropidae, Pyrgotidae, etc., and formed from the basally flexible conus, is described. Some antennal features of the Calyptratae suggest a relationship to the Tephritoidea. Critical comments are made with regard to the recently published phylogenetic association of the Ironomyiidae with the Phoridae and the Pallopteridae with the Neurochaetidae. In discussing relationships of some taxa, a few non-antennal features, some needing further study, are mentioned, e.g., variation in separation of abdominal tergites 1 and 2 in the Opetiidae and other lower cyclorrhaphous families; the presence of supplementary claw-like terminal tarsal processes in the Lonchopteridae; the apparent restriction of the presence of barbed macrotrichia to the Phoridae, among lower cyclorrhaphans; variation in structure of the prelabrum in the Pyrgotidae; the microstructure of the facial cuticle in the Syringogastridae as compared with that of other families; the calyptrate-like development of the squama in some tephritoid taxa; variation in the subscutellum in the Conopidae; a feature of the larval posterior spiracles diagnostic for Coelopidae. Mcalpine, DaviD K. 2011. Observations on antennal morphology in Diptera, with particular reference to the articular surfaces between segments 2 and 3 in the Cyclorrhapha.
... The superfamily Sphaeroceroidea is a medium-sized one, with two families of moderate diversity, Sphaeroceridae (1550 species) and Heleomyzidae (~720 species), and the small family Chyromyidae. The enigmatic afrotropical Mormotomyia hirsuta Austen, 1936 was once placed near Sphaeroceridae but it is now seen as an ephydroid fly (Kirk-Spriggs et al. 2011). McAlpine (2007 has proposed an alternative concept for Sphaeroceroidea with Sphaeroceridae and Heleomyzidae united as a single family called Heteromyzidae. ...
... The superfamily Sphaeroceroidea is a medium-sized one, with two families of moderate diversity, Sphaeroceridae (1550 species) and Heleomyzidae (~720 species), and the small family Chyromyidae. The enigmatic afrotropical Mormotomyia hirsuta Austen, 1936 was once placed near Sphaeroceridae but it is now seen as an ephydroid fly (Kirk-Spriggs et al. 2011). McAlpine (2007) has proposed an alternative concept for Sphaeroceroidea with Sphaeroceridae and Heleomyzidae united as a single family called Heteromyzidae. ...
A Finnish checklist of the sphaeroceroid fly families Chyromyidae and Heleomyzidae is provided.
... Recent molecular studies clearly associate Mystacinobia with the Oestroidea (Gleeson et al., 2000), and morphological analyses based on SEM suggest an affiliation of Mormotomyia with the Ephydroidea (Mormotomyiidae) (Kirk-Spriggs et al., 2011). ...
Host specificity gauges the degree to which a parasite occurs in association with host species and is among the most fundamental properties of parasite–host associations. The degree of specificity is indicative of myriad properties of the host and parasite and of their ecological and evolutionary relationships. Bat flies are highly specialized bloodfeeding ectoparasites of bats worldwide and were historically viewed as unspecific. In the bat fly—bat system, numerous properties actually or potentially interrupt the linkage of parasite to host and should thus decrease specificity. Such properties of bat flies include a life history strategy requiring females to leave the host, an off-host pupal stage, and high dispersal capability of many species. For hosts, properties include high diversity, mobility, sociality, and multispecies roosting environments. These and other biological and ecological characteristics of bats and flies should together facilitate interspecific host transfers and over time lead to nonspecific host–parasite associations. Despite these properties, large and carefully executed biodiversity surveys of mammals and parasites unequivocally demonstrate the high host specificity of many bat flies, and molecular sequence data promise to demonstrate that many cases of lowered specificity are misunderstood due to unresolved parasite species boundaries. On the other hand, experimental approaches have suggested that host specificity is context dependent and may be lessened in cases of ecological disturbance and in particular when novel host–parasite associations are created. Evolution and maintenance of specificity in bat flies depends in part on the encounter and compatibility properties of bats and on the reproductive potential of the flies on available host species. Moreover, the degree to which parasites have coevolved immunological compatibility with their hosts, thereby diminishing immunological surveillance and response, may also serve to maintain high host specificity. Although worldwide bat species on average harbor higher diversity of parasites and pathogens than any other mammalian group, the likelihood of bat flies vectoring disease agents across host species of bats, and particularly to distantly related mammals such as humans, may be relatively small.
The development of molecular tools has dramatically increased our knowledge of parasite diversity and the vectors that transmit them. From viruses and protists to arthropods and helminths, each branch of the Tree of Life offers an insight into significant, yet cryptic, biodiversity. Alongside this, the studies of host-parasite interactions and parasitism have influenced many scientific disciplines, such as biogeography and evolutionary ecology, by using comparative methods based on phylogenetic information to unravel shared evolutionary histories. Parasite Diversity and Diversification brings together two active fields of research, phylogenetics and evolutionary ecology, to reveal and explain the patterns of parasite diversity and the diversification of their hosts. This book will encourage students and researchers in the fields of ecology and evolution of parasitism, as well as animal and human health, to integrate phylogenetics into the investigation of parasitism in evolutionary ecology, health ecology, medicine and conservation.
The development of molecular tools has dramatically increased our knowledge of parasite diversity and the vectors that transmit them. From viruses and protists to arthropods and helminths, each branch of the Tree of Life offers an insight into significant, yet cryptic, biodiversity. Alongside this, the studies of host-parasite interactions and parasitism have influenced many scientific disciplines, such as biogeography and evolutionary ecology, by using comparative methods based on phylogenetic information to unravel shared evolutionary histories. Parasite Diversity and Diversification brings together two active fields of research, phylogenetics and evolutionary ecology, to reveal and explain the patterns of parasite diversity and the diversification of their hosts. This book will encourage students and researchers in the fields of ecology and evolution of parasitism, as well as animal and human health, to integrate phylogenetics into the investigation of parasitism in evolutionary ecology, health ecology, medicine and conservation.
The development of molecular tools has dramatically increased our knowledge of parasite diversity and the vectors that transmit them. From viruses and protists to arthropods and helminths, each branch of the Tree of Life offers an insight into significant, yet cryptic, biodiversity. Alongside this, the studies of host-parasite interactions and parasitism have influenced many scientific disciplines, such as biogeography and evolutionary ecology, by using comparative methods based on phylogenetic information to unravel shared evolutionary histories. Parasite Diversity and Diversification brings together two active fields of research, phylogenetics and evolutionary ecology, to reveal and explain the patterns of parasite diversity and the diversification of their hosts. This book will encourage students and researchers in the fields of ecology and evolution of parasitism, as well as animal and human health, to integrate phylogenetics into the investigation of parasitism in evolutionary ecology, health ecology, medicine and conservation.
Insect parasites and parasitoids are a major component of terrestrial food webs. For parasitoids, categorization is whether feeding activity is located inside or outside its host, if the host is immobilized or allowed to grow, and if the feeding is done by one or many conspecific or heterospecific individuals, and other features. Fossil evidence for parasitism and parasitoidism consists of taxonomic affiliation, morphology, gut contents, coprolites, tissue damage and trace fossils. Ten hemimetabolous and holometabolous orders of insects developed the parasite condition whereas seven orders of holometabolous insects evolved the parasitoid life habit. Modern terrestrial food webs are important for understanding the Mid Mesozoic Parasitoid Revolution. The MMPR began in late Early Jurassic (Phase 1), in which bottom-to-top regulation of terrestrial food webs dominated by inefficient clades of predators were replaced by top-to-bottom control by trophically more efficient parasitoid clades. The MMPR became consolidated in Phase 2 by the end of the Early Cretaceous. These clades later expanded (phases 3 and 4) as parasitoids became significant ecological elements in terrestrial food webs. Bottom-to-top food webs explained by the resource concentration hypothesis characterize pre-MMPR time. During phases 1 and 2 of MMPR (Middle Jurassic to Early Cretaceous), a shift ensued toward top-to-down food webs, explained by the trophic cascade hypothesis, exemplified by hymenopteran parasitoid clades Stephanoidea and Evanioidea. Clade-specific innovations spurring the MMPR included long, flexible ovipositors (wasps), host seeking, triungulin and planidium larvae (mantispids, beetles, twisted-wing parasites, flies), and extrudable, telescoped ovipositors (flies). After the MMPR, in phases 3 and 4 (Late Cretaceous to Recent), parasitoids increased in taxonomic diversity, becoming integrated into food webs that continue to the present day.
Since the idea of coevolution as a relevant concept for the study of evolutionary ecology of communities was introduced by Ehrlich and Raven (1964), there has been a vast literature around this concept. The first formal definition of coevolution can be attributed to Janzen (1980): coevolution is a change of trait values in a first population as a response to the trait values of a second population, followed by a change of trait value in the second population in response to the new trait value in the first (different modalities of trait dynamics have been described since then – Gandon et al., 2008). Much emphasis is put on the fact that the existence of an interaction is not indicative that the species have coevolved (it can reflect a recent host acquisition, for example). Based on this, Janzen recommends considerable caution when using the word coevolution, and it is worth asking whether, more than 50 years after this word first appeared, we are being cautious enough.
The diversity of organisms is affected by a variety of biotic and abiotic factors. One of the most important forces that affect the diversity of a community is the relationship between this community and communities of higher and/or lower trophic levels. Indeed, a strong link between the diversity of consumers and that of resources is a general characteristic of natural food webs (Polis & Strong, 1996). Top-down effects occur when the diversity of communities at a higher trophic level influences the diversity of communities at lower trophic levels (e.g. Jakobsen et al., 2004), while bottom-up effects occur when the diversity at lower trophic levels controls the diversity at higher levels (e.g. Siemann, 1998; Brandle et al., 2001; Haddad et al., 2009). Moreover, top-down and bottom-up forces can act on communities simultaneously (Hunter & Price, 1992).
The development of molecular tools has dramatically increased our knowledge of parasite diversity and the vectors that transmit them. From viruses and protists to arthropods and helminths, each branch of the Tree of Life offers an insight into significant, yet cryptic, biodiversity. Alongside this, the studies of host-parasite interactions and parasitism have influenced many scientific disciplines, such as biogeography and evolutionary ecology, by using comparative methods based on phylogenetic information to unravel shared evolutionary histories. Parasite Diversity and Diversification brings together two active fields of research, phylogenetics and evolutionary ecology, to reveal and explain the patterns of parasite diversity and the diversification of their hosts. This book will encourage students and researchers in the fields of ecology and evolution of parasitism, as well as animal and human health, to integrate phylogenetics into the investigation of parasitism in evolutionary ecology, health ecology, medicine and conservation.
Species evolved from common ancestors often share many features pertaining to a variety of traits (e.g. Hansen & Martins, 1996; Blomberg & Garland, 2002). The tendency for phylogenetically related species to resemble one another has been labelled variously ‘phylogenetic inertia’ (Wilson, 1975), ‘phylogenetic conservatism’ (Ashton, 2001), ‘phylogenetic correlation’ (Gittleman et al., 1996) and ‘phylogenetic effect’ (Derrickson & Ricklefs, 1988). Recently, Blomberg and Garland (2002) and Blomberg et al. (2003) have argued that the use of some of these terms suggests the action of certain evolutionary mechanisms, although such mechanisms cannot be inferred or estimated from comparative data. Instead, Blomberg and Garland (2002) and Blomberg et al. (2003) have recommended the use of the term ‘phylogenetic signal’ for this pattern because it does not imply any evolutionary mechanism or process that could have caused this resemblance. Indeed, simulations have demonstrated that different evolutionary processes may produce similar phylogenetic signals (Revell et al., 2008).
Parasites are considered to constitute half of the number of living things, and no single species is considered to be parasite-free. Due to important insights gained from clinical, epidemiological or ecological field studies, parasitologists are aware that investigating host–parasite interactions in natural systems may be spurious if not considering the huge diversity of parasites (Petney & Andrews, 1998; Poulin & Morand, 2004; Adams et al., 2010; Steinmann et al., 2010). Multiple infections are the rule because host infection often involves two or more parasite species or genotypes. The number of parasites, their host impacts, their interactions and their circulations in natural and disturbed ecosystems are clearly a knowledge frontier in parasitology and ecology (Telfer et al., 2010; Tompkins et al., 2010; Johnson & Hoverman, 2012).
Rickettsia is a genus of alpha-proteobacteria in the order Rickettsiales. All known members of the genus are obligate intracellular endosymbionts, unable to survive outside the host cell environment. However, this fundamental similarity belies the extraordinary diversity of the group. For example, known hosts of Rickettsia are found in freshwater, marine and terrestrial habitats, and include protozoa, arthropods, vertebrates, photosynthetic algae and plants (Perlman et al., 2006; Weinert et al., 2009b). This cosmopolitan host range is mirrored by the range of transmission strategies employed by the bacteria (Table 8.1). Rickettsia strains vary widely in their dependence on horizontal versus vertical transmission, and in their effects on their hosts (which range from mutualism to parasitism, and include a remarkable array of reproductive manipulations).
The majority of animals on this planet are invertebrates, and a great number of them are found in aquatic habitats including freshwater, brackish or marine environments. It is likely that they also harbour a significant fraction of all parasite biodiversity.
Studies of the evolution of virulence aim at understanding how and why certain parasite strains have evolved to cause morbidity and mortality to their hosts, while others have remained benign. In parasitism, according to definition, a parasite benefits at the expense of its host. These benefits are usually realized upon harming the host following infection. Conventional wisdom holds that damaging the host is detrimental to the interests of the invading parasites. Therefore, parasites should have evolved to become avirulent to their hosts as they otherwise risk driving their hosts, and therefore themselves, to extinction. This view, which has been criticized for its reliance on group selection, no longer prevails (Lenski & May, 1994). Instead, the evolutionary theory of virulence strives to elucidate the underlying selective forces that lead to increased or reduced virulence by examining the costs and benefits of virulence to both parasite and host. Ultimately, this theory attempts to identify which expressions of virulence are adaptive in the long-run, and which are non-adaptive, i.e. coincidental or short-sighted (Levin & Bull, 1994; Levin, 1996).
Acanthocephalans are an enigmatic group of endoparasites with complex life-cycles that involve vertebrates as final definitive hosts and invertebrates as intermediate hosts. The name of the phylum refers to the attachment organ commonly known as a proboscis. The proboscis is variable in shape and is covered by a tegument within which are embedded the roots of recurved, sclerotized hooks. Hooks are the structures that allow these parasites to attach to the intestinal wall, causing on some occasions severe damage to their definitive hosts. The proboscis shape, and the number, size and arrangement of hooks, are taxonomic traits that have been traditionally used to classify approximately 1300 described species distributed worldwide (Amin, 1987; Poulin & Morand, 2004; Kennedy, 2006; Amin, 2013; Figure 9.1).
Parasites live in an intimately close relationship with their hosts, either inserted within their anatomic structures or securely attached onto their surface. Therefore, we expect biotic effects – such as host individual, population and community characters – to influence all features of parasite communities, including richness (Poulin, 2008). In contrast, physical features of the abiotic environment, such as temperature, humidity, etc. are not usually expected to influence communities of parasites directly. The direct influence of the physical environment may be even less pronounced in terrestrial than in aquatic habitats because the terrestrial environment is less habitable for the free-living stages of parasites. Not surprisingly, major textbooks on parasite ecology rarely discuss the effects of abiotic environmental factors on terrestrial parasite assemblages (but see Bordes et al., 2010).
Bats (Chiroptera) represent more than 20% of the known mammalian diversity, making them second only to Rodentia (Teeling et al., 2005; Wilson & Reeder, 2005). As bats were carving out their predominantly aerial and nocturnal niche, Diptera had long prevailed on Earth (origins: 260 mya) (Wiegmann et al., 2011). Among these were the ecologically highly diverse Calyptratae. Presumably, at some point a lineage of these flies began to associate with this newly available mammalian niche, which subsequently gave rise to the extant obligatory parasitic bat flies, currently encompassing ~500 described species (Dick & Patterson, 2006).
This paper presents the results of investigations conducted between 2011 and 2013 to discover additional populations of Mormotomyia hirsuta Austen. These investigations were conducted primarily in the relatively dry savanna of eastern Kenya, focusing on small hills and rocky outcrops resembling that of Ukasi Hill, the type locality of the “terrible hairy fly”. Investigations were conducted at 144 caves and at ground level, directly below 104 above-ground, narrow, horizontally-oriented fissures, often on near-vertical rock faces. Evidence of Mormotomyia was not found in any of the caves investigated. During the dry season, however, desiccated corpses of Mormotomyia were discovered embedded in a matrix of dried bat guano adhering to the rock face directly below fissures at Ngauluka and Makilu Hills, also located in the Ukasi area. Later, rainy season visits to these two hills revealed populations of living Mormotomyia while, contemporaneously, flies were absent from the type locality. Like the type locality, the rock face directly below the fissures on Ngauluka and Makilu was discolored with pink and purple vertical streaking, presumably stained by bat urine and guano. Using the characteristically stained rock face as a search image, expeditions were expanded to include areas further afield and living flies were found at a third site 187 km to the south. Formerly considered “the rarest fly in the world”, the conservation status of Mormotomyia appears robust. Mormotomyia was actively preyed upon in the field by two species of lizards and remains of the fly were found in a jumping-spider nest. During laboratory observations of five live flies, the single male exhibited lengthy periods of female-guarding, with females being enclosed within the span of the much longer and setulose legs of males for more than 10 minutes.
Tapeworms (cestodes) and flukes (trematodes) comprise approximately 56% of the currently known ~30 000 species of platyhelminths (Caira & Littlewood, 2013). These flatworms are considered to be among the most successful of metazoan parasites, as measured by their numerical abundance, their geographical reach and host (habitat) diversity (e.g. Poulin & Morand, 2004). Their complex life-cycles involve one or more intermediate host(s) from molluscs (particularly for trematodes), arthropods (particularly for cestodes), annelids, ctenophores, echinoderms, hexapods and vertebrates, before reaching sexual maturity in a definitive (almost exclusively vertebrate) host.
Curtonotidae are a small, mostly tropical family of acalyptrate flies in the Ephydroidea, a superfamily that also includes the Drosophilidae. Two species of Curtonotidae are described from Miocene amber of the Dominican Republic, 17–20 Ma: Curtonotum †electrodominicum Grimaldi and Kirk-Spriggs, sp. n., and Depressonotum †priscum Grimaldi and Kirk-Spriggs, gen. et sp. n. The fossil Curtonotum is based on a female specimen and has some features of Neotropical species and a small clade of African species. Depressonotum †priscum, gen. et sp. n., is based on a male and female and has a unique combination of features that are plesiomorphic and derived for the Curtonotidae. These are the only definitive fossils of the family and the only Curtonotidae known from the Caribbean. The fossil Curtonotidae provide rare data on the geological occurrence of ephydroid flies, essential for estimates of divergence times.
Abstract: A review of specific publications dealing with Baltic amber Diptera, Acalyptratae, from the years 1822 until 2008 includes
38 articles. H. LOEW was the first entomologist searching systematically for Diptera in amber. Two of his three articles are
discussed. Parts of his first one (1850) are translated from German because of its extreme rarity in libraries. From Eocene Baltic
amber 35 families of acalyptrates are known now, a further four from British Eocene sediments. Natalimyzidae, ?Piophilidae and
Pyrgotidae are recorded for the first time, Natalimyza was known only from the recent Afrotropical fauna. Different counts of the
percentage of acalyptrates among insect-, Diptera- and “true fly“-inclusions, respectively, are compared. Less than 1% of all flies
belong here. Reasons for this rarity are discussed together with an overview of the rare aggregation of acalyptrates in single amber
pieces. Peculiar morphological apomorphies, which enable family identification worldwide because of their singularity, were
already present during the Tertiary. Three such examples are discussed. The dispute about the doubted synchroneous genesis of
German Bitterfeld amber is solved by demonstrating 15 conspecific acalyptrate species in both deposits. Intraspecific variability
of an amber acalyptrate has never been studied before. On the basis of 45 specimens and the holotype of Protoscinella electrica
(Chloropidae) the slowness of evolutionary transformations is exemplified with the background of the phylogenetic value of 16
selected characters. A spotlight is thrown on the polyphyletic reduction process in wing venation and bristle equipment observed
in several families of acalyptrates.
The systematic part presents overviews of a large number of morphological details used for taxonomy. Two tables enable the easy
check and documentation of each detected specimen, as well as an understanding of the puzzling termini and abbreviation systems
in subsequent periods of dipterology. Two differently organized identification keys are presented: one listing 97 exceptional,
rare, or newly detected morphological pecularities, a second one is a trial to key out all 56 described and all 161 detected undescribed
species. The well known terminology of HENNIG‘s publication series on amber is used in order to enable an easy cross-reference
to his partly complicated descriptions, to his plentity of figures, and to the dipterous literature until the year 1981 when a
new terminology was proposed. All family- and generic transfers since HENNIG‘s times are listed with their references, as well a
species breakdown of parts of the identified 1,141 inclusions.
A review of specific publications dealing with Baltic amber Diptera, Acalyptratae, from the years 1822 until 2008 in-cludes 38 articles. H. LOEW was the first entomologist searching systematically for Diptera in amber. Two of his three articles are discussed. Parts of his first one (1850) are translated from German because of its extreme rarity in libraries. From Eocene Baltic amber 35 families of acalyptrates are known now, a further four from British Eocene sediments. Natalimyzidae, ?Piophilidae and Pyrgotidae are recorded for the first time, Natalimyza was known only from the recent Afrotropical fauna. Different counts of the percentage of acalyptrates among insect-, Diptera-and " true fly " -inclusions, respectively, are compared. Less than 1% of all flies belong here. Reasons for this rarity are discussed together with an overview of the rare aggregation of acalyptrates in single am-ber pieces. Peculiar morphological apomorphies, which enable family identification worldwide because of their singularity, were already present during the Tertiary. Three such examples are discussed. The dispute about the doubted synchroneous genesis of German Bitterfeld amber is solved by demonstrating 15 conspecific acalyptrate species in both deposits. Intraspecific variability of an amber acalyptrate has never been studied before. On the basis of 45 specimens and the holotype of Protoscinella electrica (Chloropidae) the slowness of evolutionary transformations is exemplified with the background of the phylogenetic value of 16 selected characters. A spotlight is thrown on the polyphyletic reduction process in wing venation and bristle equipment observed in several families of acalyptrates. The systematic part presents overviews of a large number of morphological details used for taxonomy. Two tables enable the easy check and documentation of each detected specimen, as well as an understanding of the puzzling termini and abbreviation sys-tems in subsequent periods of dipterology. Two differently organized identification keys are presented: one listing 97 exceptional, rare, or newly detected morphological pecularities, a second one is a trial to key out all 56 described and all 161 detected unde-scribed species. The well known terminology of HENNIG's publication series on amber is used in order to enable an easy cross-ref-erence to his partly complicated descriptions, to his plentity of figures, and to the dipterous literature until the year 1981 when a new terminology was proposed. All family-and generic transfers since HENNIG's times are listed with their references, as well a species breakdown of parts of the identified 1,141 inclusions. Santrauka: 1822-2008 metais paskelbtos 38 moksline . s publikacijos, skirtos Baltijos gintare rastiems Diptera, Acalyptratae vabzdžiams. H. LOEWAS buvo pirmasis entomologas, sistematiškai tyrine . je ˛s dvisparnius gintare. Aptariami du iš triju˛ jo straipsniu˛. Atskiros jo pirmojo darbo (1850) dalys yra išverstos iš vokiečiu˛ kalbos, nes publikacija yra tapusi didele bibliografine retenybe. Iš eoceninio Baltijos gintaro dabartiniu metu žinomos 35 akaliptratiniu˛ dvisparniu˛ (Acalyptratae) šeimos. Dar trys rastos eocenine . se nuogulose Didžiojoje Britanijos. Agromyzidae, Natalimyzidae, Piophilidae, Pyrgotidae ir Sphaeroceridae yra aprašomos pirma˛ kar-ta˛, Natalimyza buvo žinoma tik iš dabartine . s Afrikos tropiku˛ faunos. Palyginami skirtingu˛ autoriu˛ pateikiami akaliptratu˛ gausu-mo paskaičiavimai tarp visu˛ vabzdžiu˛, dvisparniu˛ ir " tikru˛ju˛ musiu˛ " inkliuzu˛. Šioms muse . ms priklauso mažiau negu vienas pro-centas gintare randamu˛ inkliuzu˛. Šio retumo priežastys aptariamos kartu apžvelgiant retai pasitaikančias akaliptratu˛ sankaupas viename gintaro gabale. Ypatingos ir kartu unikalios morfologine . s apomorfijos, kuriomis pasižymi visos Žeme . s rutulyje sutinkamos akaliptratine . s muse . s, egzistavo jau terciare. Aptariami trys tokie pavyzdžiai. Diskusija apie abejotina˛ ta˛ pačia˛ vokiškojo Biterfel-do ir Baltijos gintaru˛ kilme ˛ išsprendžiama parodant, kad abiejuose gintaruose randama 15 vienodu˛ Acaliptratae ru – šiu˛. Vidinis ru – šinis variavimas tarp gintaruose randamu˛ akaliptratu˛ anksčiau nebuvo tirtas. Remiantis 45 pavyzdžiais ir Protoscinella electrica (Chloropidae) holotipu demonstruojamas evoliuciniu˛ transformaciju˛ le . tumas kartu pagrindžiant 16 parinktu˛ požymiu˛ filogeneti-ni ˛ reikšminguma˛. Nušviečiamas polifiletinis sparno gyslotumo ir šeriuotumo redukcijos procesas, stebimas keliose akaliptratu˛ šei-mose. Denisia 26, zugleich Kataloge der oberösterreichischen Landesmuseen Neue Serie 86 (2009): 171–212
Katacamilla cavernicola Papp, collected from caves at four localities in northern and central Namibia, is here shown to exhibit both parietal and troglophilic habitat preferences, and the larval stage is trophically guanobious. Katacamilla cavernicola was reared in the laboratory from the guano of the rock pigeon Columba guinea L. (parietal), and of the common slit-faced bat Nycteris thebaica Geoffrey (troglophilic) and larvae and puparia were obtained. The duration of the larval stage of K. cavernicola was found to be 7 days and the pupal stage 6–7 days, eggs remaining dormant within guano, and larval development being apparently triggered by periodic moistening, either by urine, or other liquids. Descriptions of the third instar larva and puparium are presented and are illustrated with line drawings and stereoscan micrographs. These descriptions of the immature stages are the first known for the Camillidae. Larval features are compared to the three closely related families in the Ephydroidea, for which the immature stages are known: Drosophilidae, Curtonotidae and Ephydridae. The larval morphology of Katacamilla is found to be generally close to the putative groundplan for the Ephydroidea. However, the basal sclerite has very large open windows with the ventral arm of the dorsal cornu being very long and needle-like. The thoracic segments are very long and slender and unusually well-separated from one another, and the abdominal segments and the prothoracic Keilin organs are very close together, a trait that is within the Ephydroidea currently only known in the Ephydridae.
The dipteran clade Calyptratae is comprised of approximately 18 000 described species (12% of the known dipteran diversity) and includes well-known taxa such as houseflies, tsetse flies, blowflies and botflies, which have a close association with humans. However, the phylogenetic relationships within this insect radiation are very poorly understood and controversial. Here we propose a higher-level phylogenetic hypothesis for the Calyptratae based on an extensive DNA sequence dataset for 11 noncalyptrate outgroups and 247 calyptrate species representing all commonly accepted families in the Oestroidea and Hippoboscoidea, as well as those of the muscoid grade. DNA sequences for genes in the mitochondrial (12S, 16S, cytochrome c oxidase subunit I and cytochrome b) and nuclear genome [18S, 28S, the carbamoyl phosphate synthetase region of CAD (rudimentary), Elongation factor one alpha] were used to reconstruct the relationships. We discuss problems relating to the alignment and analysis of large datasets and emphasize the advantages of utilizing a guide tree-based approach for the alignment of the DNA sequences and using the leaf stability index to identify ‘wildcard’ taxa whose excessive instability obscures the phylogenetic signal. Our analyses support the monophyly of the Calyptratae and demonstrate that the superfamily Oestroidea is nested within the muscoid grade. We confirm that the monotypic family Mystacinobiidae is an oestroid and further revise the composition of the Oestroidea by demonstrating that the previously unplaced and still undescribed ‘McAlpine’s fly’ is nested within this superfamily as a probable sister group to Mystacinobiidae. Within the Oestroidea we confirm with molecular data that the Calliphoridae are a paraphyletic grade of lineages. The families Sarcophagidae and Rhiniidae are monophyletic, but support for the monophyly of Tachinidae and Rhinophoridae depends on analytical technique (e.g. parsimony or maximum likelihood). The superfamilies Hippoboscoidea and Oestroidea are consistently found to be monophyletic, and the paraphyly of the muscoid grade is confirmed. In the overall relationships for the calyptrates, the Hippoboscoidea are sister group to the remaining Calyptratae, and the Fanniidae are sister group to the nonhippoboscoid calyptrates, whose relationships can be summarized as (Muscidae (Oestroidea (Scathophagidae, Anthomyiidae))).
Flies are one of four superradiations of insects (along with beetles, wasps, and moths) that account for the majority of animal life on Earth. Diptera includes species known for their ubiquity (Musca domestica house fly), their role as pests (Anopheles gambiae malaria mosquito), and their value as model organisms across the biological sciences (Drosophila melanogaster). A resolved phylogeny for flies provides a framework for genomic, developmental, and evolutionary studies by facilitating comparisons across model organisms, yet recent research has suggested that fly relationships have been obscured by multiple episodes of rapid diversification. We provide a phylogenomic estimate of fly relationships based on molecules and morphology from 149 of 157 families, including 30 kb from 14 nuclear loci and complete mitochondrial genomes combined with 371 morphological characters. Multiple analyses show support for traditional groups (Brachycera, Cyclorrhapha, and Schizophora) and corroborate contentious findings, such as the anomalous Deuterophlebiidae as the sister group to all remaining Diptera. Our findings reveal that the closest relatives of the Drosophilidae are highly modified parasites (including the wingless Braulidae) of bees and other insects. Furthermore, we use micro-RNAs to resolve a node with implications for the evolution of embryonic development in Diptera. We demonstrate that flies experienced three episodes of rapid radiation--lower Diptera (220 Ma), lower Brachycera (180 Ma), and Schizophora (65 Ma)--and a number of life history transitions to hematophagy, phytophagy, and parasitism in the history of fly evolution over 260 million y.
Hippoboscoidea is a superfamily of Diptera that contains the Glossinidae or tsetse flies, the Hippoboscidae or louse flies, and two families of bat flies, the Streblidae and the Nycteribiidae. We reconstruct the phylogenetic relationships within Hippoboscoidea using maximum parsimony and Bayesian methods based on nucleotide sequences from fragments of four genes: nuclear 28S ribosomal DNA and the CPSase domain of CAD, and mitochondrial 16S rDNA and cytochrome oxidase I. We recover monophyly for most of the presently recognized groups within Hippoboscoidea including the superfamily as a whole, the Hippoboscidae, the Nycteribiidae, the bat flies, and the Pupipara (=Hippoboscidae+Nycteribiidae+Streblidae), as well as several subfamilies within the constituent families. Streblidae appear to be paraphyletic. Our phylogenetic hypothesis is well supported and decisive in that most competing topological hypotheses for the Hippoboscoidea require significantly longer trees. We confirm a single shift from a free-living fly to a blood-feeding ectoparasite of vertebrates and demonstrate that at least two host shifts from mammals to birds have occurred. Wings have been repeatedly lost, but never regained. The hippoboscoid ancestor also evolved adenotrophic viviparity and our cladogram is consistent with a gradual reduction in the motility of the deposited final instar larvae from active burrowing in the soil to true pupiparity where adult females glue the puparium within the confines of bat roosts.
The basic segmentation of the abdomen in Cyclorrhapha has been well discussed by Crampton (1942), van Emden & Hennig (1956), Steyskal (1957a) and Hennig (1958). If the andrium represents the 9th segment (as now seems almost universally accepted), the numbering of the preceding segments can be established from considerations of comparative morphology. The Cyclorrhapha contain some member groups in which all eight preceding segments are clearly defined and bear discrete sclerites. The correct numbering for groups in which sclerites have been fused or lost can be established by analysis of the sequence of morphological changes which have occurred and by ontogenetic evidence, when this is available. The interpretation of the various conditions found in particular groups is discussed below in my treatment of the groups concerned. One remaining source of dispute affecting the interpretation of the postabdominal structure of all Cyclorrhapha is whether or to what extent the 8th segment is rotated. This question is discussed in detail below in section 3.2, where I present what seems to me conclusive evidence that this segment is inverted (rotated through 180°). I therefore follow Crampton in calling the dorsal sclerite of this segment the 8th sternum, and the reduced ventral sclerite the 8th tergum. Reduction of the 8th tergum to a narrow band is probably a groundplan condition of the Cyclorrhapha (see section 4).
The internal female reproductive tract of Risa is characterized by a pair of short spermathecal ducts with no spermathecae, paired short accessory glands, and a large thimble-shaped strongly sclerotized ventral receptacle. This condition is identical with that in the ground plan of Ephydridae but differs from that of their next relatives, i.e., Camillidae and Diastatidae, regarding the shape of the ventral receptacle. It therefore corroborates the close relationship of Risa with Ephydridae either as its sister taxon or within that family.
The surface morphology of a dome-shaped genital chamber in the female Musca domestica L., where the sperm and egg meet following ovulation, was examined to determine its role in fertilization. The inner surface of the chamber was found to be lined with 3 types of nonarticulated cuticular spines. Examination of eggs removed from the chamber indicated that the distinctly robust spines at the apex were involved with the removal of a mucoid secretion which occludes the micropyle opening. The spines lining the rest of the chamber were more slender and flexible than those at the apex, and may function in ensuring that sperm remain at the fertilization site when the egg is placed into the chamber.
Mystacinobia zelandica n.sp. is described. It is the sole member of Mystacinobia new genus and of Mystacinobiidae new family, and belongs to the superfamily Drosophiloidea. The species lives in large communities in roosts of the New Zealand short‐tailed bat, Mystacina tuberculata, and requires temperatures around 30°c for development and survival. Adults are physogastric, apterous, and have reduced eyes. The claws are adapted for movement over bat fur, but the mouthparts are not modified for blood‐feeding. Adults and larvae feed on guano. Eggs are laid in clusters in roost wood, and have non‐functional respiratory horns. Larvae have elongate anterior spiracles, tubular posterior spiracles, and 5 pairs of anal papillae. The puparium has a reduced operculum. Dispersal to new roosts depends entirely on transport by Mystacina, and as many as 10 phoretic flies have been found embedded in fur of individual bats leaving a roost to feed at night. The species has reached a degree of sociality which includes group oviposition, partial overlapping of generations, clustering of all stages, mutual grooming, male polymorphism, and extension of the males’ life‐span beyond the reproductive phase to form a sound‐producing guard caste which probably prevents the bats from interfering with the bat‐fly community. Mystacinobia zelandica is part of the New Zealand Endemic (Archaic) Element, which also includes Mystacina tuberculata.
The New Zealand batfly, Mystacinobia zelandica, is unique in its phoretic association with the endemic short‐tailed bat, Mystacina tuberculata However, the origins of Mystacinobia, and its systematic position within the Diptera, have long been unclear We have investigated the phylogenetic position of Mystacinobia using sequence data derived from a 51 lbp region of the mitochondnal 16S rDNA gene A total of 17 taxa, representing 11 families and 14 genera within the Diptera, were included in the phylogenetic analysis The resulting phylogenetic trees support morphological revisions which place Mystacinobia within the Oestroidea, most closely associated with the Calhphondae
The morphology of the puparia of Curtonotum saheliense Tsacas and C. simile Tsacas are described or redescribed, based on material reared from locust egg pods in Mali and Oman, using SEM techniques and light microscopy. The original specimens on which Greathead based his 1958 larval description of C. cuthbertsoni Duda, have been located and examined and are here ascribed to C. simile, rather than C. saheliense as previously assumed, based on examination of the type specimens of both. An additional larva, also reared from locust egg pods in northern Nigeria, is tentatively ascribed to C. saheliense and is described using SEM techniques, despite its relatively poor state of preservation. The two species are shown to be separable based on the relative lengths and widths of the mouthooks and intermediate sclerite of the cephaloskeleton and in the degree of expansion of semi-circular cuticular folding adjacent to the anal opening on the anal pad. A diagnosis of the immature stages of the genus Curtonotum Macquart is provided and the larval and puparial features of C. saheliense and C. simile are compared to the puparium of the Nearctic species C. helvum (Loew), the other species of the genus for which the immature stages are partially known. The composition and homologies of caudal tubules are discussed, as are features in common between the three species under discussion, and tentative comparison is made of the known immature stages of Curtonotum to those of the known larvae of other families of the Ephydroidea, as additional larval features are now discernible for the Curtonotidae. It is concluded that the larval morphology of the Curtonotidae is generally close to the putative groundplan of the Ephydroidea and are probably most closely allied to the Drosophilidae and Camillidae. The relative length of abdominal compartments six and seven is probably an autapomorphy of the family.
The internal female reproductive tract of Risa is characterized by a pair of short spermathecal ducts with no spermathecae, paired short accessory glands, and a large thimble-shaped strongly sclerotized ventral receptacle. This condition is identical with that in the ground plan of Ephydridae but differs from that of their next relatives, i.e., Camillidae and Diastatidae, regarding the shape of the ventral receptacle. It therefore corroborates the close relationship of Risa with Ephydridae either as its sister taxon or within that family.
SYNOPSIS
An account is given of the uterine anatomy of virgin Glossina austeni Newstead and this is compared with the anatomy of the female tract both during mating and after the mating act has been completed. Notes on the functional morphology of the male genitalia are included.
Bat flies are a small but diverse group of highly specialized ectoparasitic, obligatory bloodsucking Diptera. For the first time, the phylogenetic relationships of 26 species and five subfamilies were investigated using four genes (18S rDNA, 16S rDNA, COII, and cytB) under three optimality criteria (maximum parsimony (MP), maximum likelihood (ML), and Bayesian inference). Tree topology tests of previous hypotheses were conducted under likelihood (Shimodaira-Hasegawa test). Major findings include the non-monophyly of the Streblidae and the recovery of an Old World- and a New World-Clade of bat flies. These data ambiguously resolve basal relationships between Hippoboscidae, Glossinidae, and bat flies. Recovered phylogenies resulted in either monophyly (Bayesian approach) or paraphyly (MP/ML topologies) of the bat flies, thus obscuring the potential number of possible associations with bats throughout the history of this group. Dispersal-vicariance analysis suggested the Neotropical region as the possible ancestral distribution area of the New World Streblidae and the Oriental region for the Old World bat flies. The genes examined show conflicting support across the nodes of the tree, particularly in the basal positions. Additionally, there is poor character support among all genes for the nodes associated with early hippoboscoid diversification. This results in extremely short basal branches, adding support to the idea of a rapid radiation among the four major groups of Hippoboscoidea.
Female yellow dung flies, Scathophaga stercoraria, can influence the traffic of sperm stored in their spermathecae to the site of fertilization in the bursa copulatrix. However, the anatomical mechanisms employed are largely unknown. We investigated the anatomy of the female genital tract, seeking structures involved in sperm transfer and egg fertilization. We found a membranous structure descending from the ends of the spermathecal and accessory gland ducts into the bursa copulatrix. We call this the prolatus. Sperm accumulate in the prolatus during oviposition. When an egg is in the bursa the egg micropyle, rather than being aligned towards the dorsal openings of the spermathecal ducts, lies on the opposite, ventral side. We also confirm the presence, and suggest a function for, a cuticularized pouch on the ventral wall of the anterior bursa copulatrix. This pouch, plus a previously undescribed chamber, may be homologous to the ventral receptacle/fertilization chamber found in other dipterans. Further, we describe a translucent cap, apparently transversed by channels, covering the micropyle. Sperm were observed to aggregate on and in the micropyle cap, which appears to attract and hold sperm. We interpret the prolatus as a structure that allows an ovipositing female to transfer a few sperm onto the ventral bursal wall and thus, indirectly, onto the micropyle cap. Such anatomy potentially gives the female a large degree of control over sperm traffic from storage to the site of fertilization.
De Meyer, dated 5 September14), dorso-sublateral pits (Fig. 13, dslp), with three overlapping leaf-shaped projections arranged in a spiral
- G R Letter
- Cunningham-Van Someren
Letter: G.R. Cunningham-van Someren, to M. De Meyer, dated 5 September14), dorso-sublateral pits (Fig. 13, dslp), with three overlapping leaf-shaped projections arranged in a spiral (Fig. 15);
Anal pad (Fig. 22, apWUDQVYHUVHHORQJDWHVXEUHFWDQJXODUDQXVVXUURXQGHGE\ODUJHUDLVHGÀHVK\ Muscidae (Musca domestica L.) (43), Camillidae (Katacamilla sp
- Fig
Fig. 23). Anal pad (Fig. 22, apWUDQVYHUVHHORQJDWHVXEUHFWDQJXODUDQXVVXUURXQGHGE\ODUJHUDLVHGÀHVK\ Muscidae (Musca domestica L.) (43), Camillidae (Katacamilla sp. n.) (19), Fanniidae (Fannia sp.) (3) and Chyromyidae (1).
Speomyia absoloni, n.g., sp.n. (Dipt.), eine degenerierte Hhlengliege aus dem herzegovinisch-montenegrinosohen Hochgebirge.
- M. Bezzi
BEZZI, M. 1914. Speomyia absoloni
Zoologischer Anzeiger 44: 504-507.
A fossil tsetse fly and other Diptera from Florissant, Colorado.
- T.D.A. Cockerell
COCKERELL, T.D.A. 1917. A fossil tsetse
Proceedings of the
Biological Society of Washington 30: 19-22.
7KHSK\ORJHQHWLFFODVVL¿FDWLRQRI'LSWHUD&\FORUUKDSKDZLWKVSHFLDOUHIHUHQFHWR the structure of the male postabdomen7KH+DJXH+ROODQG Evolution of the insects Die Familien der Diptera Schizophora und ihre phylogenetischen Verwandtschaftsbeziehungen
- G C D Griffiths
- D Xqn Grimaldi
- M S Engel
- W Hennig
GRIFFITHS, G.C.D. 1972. 7KHSK\ORJHQHWLFFODVVL¿FDWLRQRI'LSWHUD&\FORUUKDSKDZLWKVSHFLDOUHIHUHQFHWR the structure of the male postabdomen7KH+DJXH+ROODQG:-XQN GRIMALDI, D. & ENGEL, M.S. 2005. Evolution of the insects. Cambridge: Cambridge University Press. HENNIG, W. 1958. Die Familien der Diptera Schizophora und ihre phylogenetischen Verwandtschaftsbeziehungen. Beiträge zur Entomologie 8: 505–688.
Catalogue of the Diptera of the Afrotropical RegionDiptera: Curtonotidae), from Africa and the Middle East, with a discussion of relationships to other known Ephydroidea larvae
- Family Nycteribiidae
Family Nycteribiidae. In: Crosskey, R.W., ed., Catalogue of the Diptera of the Afrotropical Region/RQGRQ%ULWLVK0XVHXP1DWXUDO+LVWRU\SS± KIRK-SPRIGGS$+$FRQWULEXWLRQWRWKHNQRZOHGJHRIWKHLPPDWXUHVWDJHVRICurtonotum (Diptera: Curtonotidae), from Africa and the Middle East, with a discussion of relationships to other known Ephydroidea larvae. African Entomology 16: 226–243.
The immature stages of Katacamilla cavernicola 3DSSWKH¿UVWGHVFULEHGIRUWKH&DPLOOLGDH'LSWHUD6FKL]RSKRUDZLWKFRPSDULVRQWRRWKHU known Ephydroidea larvae, and notes on biology
- Kirk-Spriggs +barraclough
- D A Meier
KIRK-SPRIGGS$+BARRACLOUGH, D.A. & MEIER, R. 2002. The immature stages of Katacamilla cavernicola 3DSSWKH¿UVWGHVFULEHGIRUWKH&DPLOOLGDH'LSWHUD6FKL]RSKRUDZLWKFRPSDULVRQWRRWKHU known Ephydroidea larvae, and notes on biology. Journal of Natural History 36: 1105–1128.
1.3. Morphology and terminology of the female postabdomen. In
- M. Kotrba
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KOTRBA, M. 2000. 1.3. Morphology and terminology of the female postabdomen. In: Papp, L. & Darvas,
B., eds, Contributions to a Manual of Palaearctic Diptera.
pp. 75-84.
The internal female reproductive tract of Opomyzidae (Diptera, Schizophora).
- M. Kotrba
- A. Baptista
KOTRBA, M. & BAPTISTA, A. 2002. The internal female reproductive tract of Opomyzidae (Diptera, Schizophora). Studia dipterologica 9: 57-71.
The ecology of ectoparasitic insects Les Dipteres cavernicoles
- A G Marshall
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MARSHALL, A.G. 1981. The ecology of ectoparasitic insects. London: Academic Press. MATILE, L. 1994. Les Dipteres cavernicoles. In: Juberthie, C. & Decu, V., eds, Encycloépaedia biospeologica.
LSWHUD6FKL]RSKRUDDQGDUHFODVVL¿FDWLRQ of the family into tribes
- Mcalpine ' 7kh
- Xvwudoldqjhqhudri+hohrp
- Lgdh
MCALPINE'.7KH$XVWUDOLDQJHQHUDRI+HOHRP\]LGDH'LSWHUD6FKL]RSKRUDDQGDUHFODVVL¿FDWLRQ of the family into tribes. Records of the Australian Museum 36: 203–251.
Chapter 100 Cryptochaetidae
- J F Mcalpine
MCALPINE, J.F. 1987. Chapter 100. Cryptochaetidae. In: McAlpine, J.F. et al., eds, Manual of Nearctic Diptera. Vol.
2QWKHQDWXUDOKLVWRU\DQGPRUSKRORJ\RIWKHHJJ¿UVWLQVWDUODUYD puparium, and female reproductive system of Curtonotum helvum (Curtonotidae; Ephydroidea
- R Meier
- M Kotrba
- Barber
MEIER, R., KOTRBA, M. & BARBER.2QWKHQDWXUDOKLVWRU\DQGPRUSKRORJ\RIWKHHJJ¿UVWLQVWDUODUYD puparium, and female reproductive system of Curtonotum helvum (Curtonotidae; Ephydroidea;
Chapter 81. Braulidae. In
- B.V. Peterson
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PETERSON, B.V. 1987. Chapter 81. Braulidae. In: McAlpine, J.F. et al., eds, Manual of Nearctic Diptera. Vol.
Superfamily Muscoidea. 81. Mormotomyiidae Catalogue of the Diptera of the Afrotropical Region The female terminalia of the Agromyzidae, with description of a new genus (I)
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PONT, A.C. 1980. Superfamily Muscoidea. 81. Mormotomyiidae. In: Crosskey, R.W., ed., Catalogue of the Diptera of the Afrotropical Region/RQGRQ%ULWLVK0XVHXP1DWXUDO+LVWRU\S SASAKAWA, M. 1958. The female terminalia of the Agromyzidae, with description of a new genus (I). 6FLHQWL¿F Reports of the Saikyo University, Agriculture, Kyoto 10: 133–150.
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STEYSKAL, G.C. 1987a. Chapter 58. Tanypezidae. In: McAlpine, J.F. et al., eds, Manual of Nearctic Di-ptera. Vol.
The anatomy of fertilization in the Scathophaga stercoraria AUSTEN a new family, genus, and species
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ARTHUR, B.I. Jr., SBILORDOPEMBERTON, A.J. & WARD, P.I. 2008. The anatomy of fertilization in the
Scathophaga stercoraria. Journal of Morphology 269: 630–637.
AUSTEN
a new family, genus, and species. Proceedings of the Zoological Society of London [1936]:
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The immature stages of Katacamilla cavernicola known Ephydroidea larvae, and notes on biology
- Kirk-Spriggs Barraclough
- D A Meier
- R Kirk-Spriggs
- B R Stuckenberg
KIRK-SPRIGGS
BARRACLOUGH, D.A. & MEIER, R. 2002. The immature stages of Katacamilla cavernicola
known Ephydroidea larvae, and notes on biology. Journal of Natural History 36: 1105-1128.
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STUCKENBERG, B.R. 2009. Afrotropical Diptera -rich savannas, poor rainforests. In:
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88. Family Streblidae Catalogue of the Diptera of the Afrotropical Region Family Nycteribiidae Catalogue of the Diptera of the Afrotropical Region KIRK-SPRIGGSCurtonotum (Diptera: Curtonotidae), from Africa and the Middle East, with a discussion of relationships to other known Ephydroidea larvae
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HUTSON, A.M. & OLDROYDa. 88. Family Streblidae. In: Crosskey, R.W., ed., Catalogue of the Diptera
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KIRK-SPRIGGSCurtonotum (Diptera:
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Ephydroidea larvae. African Entomology 16: 226–243.
1.4. Morphology and terminology of Diptera larvae Contribution to a manual of Palaearctic Diptera (with special economic importance General and applied dipterology Cave life evolution and ecology Molecular phylogenetic analysis of nycteribiid phylogeographic origins
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economic importance), Volume 1. General and applied dipterology. Budapest:
CULVER, D.C. 1982. Cave life evolution and ecology
DITTMAR, K., PORTER, M.L., MURRAY, S. & WHITING, M.F. 2006. Molecular phylogenetic analysis of nycteribiid
phylogeographic origins. Molecular Phylogenetics and Evolution 38: 155–170.
MCALPINE of the family into tribes
- L Matile
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MCALPINE
of the family into tribes. Records of the Australian Museum 36: 203-251.
Molecular phylogenetic analysis of nycteribiid phylogeographic origins
- D C Culver
- K Dittmar
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- M F Whiting
CULVER, D.C. 1982. Cave life evolution and ecology
DITTMAR, K., PORTER, M.L., MURRAY, S. & WHITING, M.F. 2006. Molecular phylogenetic analysis of nycteribiid
phylogeographic origins. Molecular Phylogenetics and Evolution 38: 155-170.
& OLDROYD a. 88. Family Streblidae
- A M Hutson
HUTSON, A.M. & OLDROYD
a. 88. Family Streblidae. In: Crosskey, R.W., ed., Catalogue of the Diptera
of the Afrotropical Region
------1980b. 89. Family Nycteribiidae. In: Crosskey, R.W., ed., Catalogue of the Diptera of the Afrotropical
Region
KIRK-SPRIGGS
Curtonotum (Diptera: