Ashley H. Kirk-Spriggs

Rediscovery of the “terrible hairy fly”, Mormotomyia hirsuta Austen (Diptera: Mormotomyiidae), in Eastern Kenya, with Notes on Biology, Natural History, and Genetic Variation of the Ukasi Hill Population

ABSTRACT 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, exposed 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 protected site.

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.

A second New World hoverfly, Toxomerus floralis (Fabricius) (Diptera: Syrphidae), recorded from the Old World, with description of larval pollen-feeding ecology

Recently (2013-2014), several hoverfly specimens from two localities in Benin and Cameroon (West and Central Africa) were caught from a species that we could not identify using existing identification keys for Afrotropical Syrphidae. Specific identification as Toxomerus floralis (Fabricius) was accomplished using morphology and various Neotropical identification keys. Corroboration of this identification was made by sequencing of the standard COI barcode region and a subsequent BLAST-IDS in BOLD that revealed a 100% sequence similarity with Toxomerus floralis from Suriname (South America). Species identification was further supported by sequencing parts of the nuclear 18S and 28S rRNA genes. The species is widespread in Togo, Benin, Nigeria and Cameroon, and eggs, larvae and adults are abundant at several localities. Yet, the full extent of its geographic distribution within tropical Africa remains to be determined. This is only the second known established introduction of a non-African hoverfly species in the Afrotropics. Interestingly, the larvae of the species have been reported as predators of Aphididae and Delphacidae but we found them to be pollenivorous, which is a rare feeding mode within the subfamily Syrphinae. Moreover, it is the only known Syrphinae species of which the larvae feed on pollen from two plant species from different families (Cyperaceae and Orobranchaceae). This example illustrates how DNA barcoding may allow a fast and accurate identification of introduced species.

A New Species of Laboulbenia (Ascomycota) Parasitic on an African Fly (Diptera: Curtonotidae), with a Brief Review of Diptera-Associated Species of the Genus

ABSTRACT A new parasitic fungus, Laboulbenia curtonoti sp. n., associated with the endemic Madagascan fly Curtonotum balachowskyi Tsacas (Diptera: Curtonotidae), is described and figured. The new species is one of the very few Laboulbeniales which penetrate the insects cuticle by means of more-or-less developed rhizoids. A brief review of the 23 species of the genus Laboulbenia associated with Diptera is presented in a tabulated form.

New Muscidae (Diptera) Hosts of Stylogaster Macquart (Diptera: Conopidae) from the Afrotropical Region

ABSTRACT An extensive collection of Muscidae, amassed through recent expeditions to Burundi, the Democratic Republic of the Congo, Kenya, Namibia and South Africa, is now deposited at the National Museum, Bloemfontein, South Africa. Twenty-five specimens of 14 species impaled with the eggs of Stylogaster Macquart (Diptera: Conopidae) were discovered among this material. New records of Stylogaster eggs are presented for four muscid genera: Afromydaea Malloch, 1930, Coenosia Meigen, 1826, Hebecnema Schnabl, 1889 and Pseudohelina Vockeroth, 1972, and for eight species: Afromydaea debilis (Stein, 1913), Coenosia ruwenzorica (Emden, 1940), Hebecnema semiflava Stein, 1913, Limnophora obsignata (Rondani, 1866), Pseudohelina nigritarsis (Jaennicke, 1867), Pseudohelina phaeoxantha (Emden, 1951), Pyrellina abdominalis Zielke, 1971 and Pyrellina versatilis (Villeneuve, 1916). A brief discussion of these associations is presented.

A Revision of Afrotropical Quasimodo Flies (Diptera: Schizophora; Curtonotidae). Part III — the Malagasy Species of Curtonotum Macquart, with Descriptions of Six New Species

ABSTRACT The Madagascan fauna of the genus Curtonotum Macquart, 1844 is reviewed. Type material of the seven species described by Tsacas in 1974 (C. balachowskyi, C. boeny, C. keiseri, C. pauliani, C. sakalava, C. sternithrix and C. stuckenbergi) was studied and additional material of five of these is noted, substantially increasing their known distributions. Six of the seven species described by Tsacas are endemic to Madagascar; unpublished records indicate, however, that C. pauliani, occurs in Namibia and South Africa on the African continental mainland. Six additional endemic species are described as new: C. coronaeformis sp. n., C. gladiiformis sp. n., C. griveaudi sp. n., C. irwini sp. n., C. parkeri sp. n., and C. rinhatinana sp. n. The head and thorax, frons, wing, sixth sternite and hypandrium of the male of the 13 species are illustrated for the first time, as well as the highly diagnostic male phallus, both laterally and dorsally at the junction of the basiphallus and distiphallus. A key to species based on male characters is provided, and species distributions are mapped and interpreted. The biogeographical significance of the Madagascan species is discussed. An annotated checklist of Madagascan Curtonotidae is presented, and co-ordinates used to plot maps and a list of vegetation types in which species occur are provided.

A new species of Alhajarmyia Stuckenberg (Diptera: Vermileonidae), the first wormlion fly described from East Africa and its biogeographical implications

A second species of the genus Alhajarmyia Stuckenberg (A. stuckenbergi Swart, Kirk-Spriggs & Copeland, sp. n.), is described and figured, from the Eastern Arc Mountains of Kenya (Kasigau Mountain and Taita Hills), being the first vermileonid recorded from East Africa. The species is shown to differ from its congener, A. umbraticola (Stuckenberg & Fisher), described from Oman in the Arabian Peninsula, based on external characters including male and female terminalia. An identification key is provided together with distribution maps for the two species, and biogeographical aspects are discussed.

Newly Discovered Populations of the “Terrible Hairy Fly”, Mormotomyia hirsuta Austen (Diptera: Mormotomyiidae) in Kenya, with Further Observations on Natural History

ABSTRACT 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.

A revision of Afrotropical Quasimodo flies (Diptera: Schizophora; Curtonotidae). Part IV—the continental Afrotropical species of Curtonotum Macquart, with descriptions of thirteen new species and a combined phylogenetic analysis of the Curtonotidae.

A first comprehensive phylogeny of the entire family Curtonotidae is generated from molecular markers and morphology. The molecular data set comprises 33 taxa (30 in-group Curtonotidae; three out-groups: Camilla, Diastata, Drosophila) and 4 gene regions from 3 genes: 2 non-contiguous fragments from the CPSase (carbamoylphosphate synthetase) domain of the nuclear protein coding gene CAD (= CAD1 and CAD3); a fragment from the coding region of TPI (triosephosophate isomerase); and a fragment of the mitochondrial gene CO1 (cytochrome oxidase 1). We performed Bayesian like-lihood analyses in the program MrBayes 3.2; maximum likelihood analyses in the program Garli 2.0; and parsimony analysis in TNT on the concatenated genetic dataset. A data matrix of 62 discrete, morphological features of imagines was compiled from 75 taxa (70 in-group Curtonotidae taxa and five out-group exemplars: Amiota, Camilla, Diastata, Drosophila, Stegana), and these data are presented as Appendix II. For the combined morphological and molecular data a Bayes-ian likelihood analysis in the program MrBayes 3.2 and a parsimony analysis in TNT were performed, and for the morphological dataset a parsimony analysis was carried out in TNT. Results of the molecular and morphological analyses attest to the monophyly of the Curtonotidae and clearly indicate two primary clades, with Axinota + Curtonotum being sister to the remainder of the Curtonotidae. Curtonotum sensu stricto (sensu Klymko and Marshall 2011) is here adopted and ten newly-defined species-groups of the genus Curtonotum are recognised the: anus; boeny; campsiphallum; gonzo; platyphallum; rinhatinana; saheliense; striatifrons; stuckenbergi; and uncinatum species-groups. The following nomenclatorial changes are proposed: Cyrtona appendiculata Séguy, 1938 is formally reinstated as a valid species and is re-moved as a junior synonym of Cyrtona pictipennis (Thomson, 1869). The former variety name sublineata (Duda, 1939) is upgraded to a specific name, as Parapsinota sublineata (Duda, 1939). The continental Afrotropical fauna of the genus Curtonotum Macquart, 1844 is revised and a diagnosis of the genus is provided. Known biology, behaviour and published information on immature stages of the genus are briefly reviewed. Type material of 12 of the 13 named species (C. angolense Tsacas, C. campsiphallum Tsacas, C. cuthbertsoni Duda, C. herrero Tsacas, C. pauliani Tsacas, C. platyphallum Tsacas, C. quinquevittatum Curran, C. saheliense Tsacas, C. sao Tsacas, C. simile Tsacas, C. striatifrons Malloch and C. tigrinum Séguy), was studied and errors in previous interpretations and designation of type specimens are resolved. Curtonotum pauliani is the only species occurring on both the continental African mainland and Madagascar. The type spec-imen of C. maculiventris (Enderlein) is lost and a neotype is here designated. One species synonymy is proposed: C. tigrinum Séguy, 1933 = C. maculiventris (Enderlein, 1917), syn. n. Additional material of the aforementioned species is noted, substantially increasing their known distributions. Thirteen species are described as new, namely: C. bicuspis Kirk- Spriggs, sp. n., C. cimbebas Kirk-Spriggs, sp. n., C. constance Kirk-Spriggs, sp. n., C. freidberg Kirk-Spriggs, sp. n., C. gonzo Kirk-Spriggs, sp. n., C. hay Kirk-Spriggs, sp. n., C. litoralis Kirk-Spriggs, sp. n., C. marriott Kirk-Spriggs, sp. n., C. mcgregor Kirk-Spriggs, sp. n., C. moffatt Kirk-Spriggs, sp. n., C. tsacas Kirk-Spriggs, sp. n., C. uncinatum Kirk- Spriggs, sp. n. and C. unicuspis Kirk-Spriggs, sp. n. The head and thorax, frons, wing, fifth sternite and hypandrium of the male of the 25 named species are illustrated for the first time, as well as the highly diagnostic male phallus, from both the right and left sides laterally. A key to species based on male characters is provided, and species distributions are mapped and interpreted according to major vegetation types, topography and humidity zones. The biogeographical signif-icance of the continental Afrotropical species is discussed. Co-ordinates used to plot maps and a list of Major Habitat Types and Vegetation Types in which species occur are provided as Appendix III.