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Newly discovered populations of the "terrible hairy fly", Mormotomyia hirsuta Austen (Diptera: Mormotomyiidae) in Kenya, with further observations on natural history.

INTRODUCTION

Mormotomyia hirsuta, the "terrible hairy fly", was described by E.E. Austen in 1936, based on two male specimens collected in 1933, at Ukasi Hill (as Ukazzi) in the dry eastern Sahelian zone (Coe 1999: 5) of eastern Kenya. The fly is a curious-looking, brachypterous species (Fig. 1), with long, orange-yellow setulae covering the body--a feature particularly well pronounced in males. The non-functional wings are reduced to setulose, strap-like appendages and the halteres are likewise reduced to barely visible nodular processes (Kirk-Spriggs et al. 2011). The species was sufficiently distinctive to warrant the erection of the new family Mormotomyiidae to contain it (Austen 1936). Mormotomyia hirsuta (confined to Kenya), is currently the sole representative of the family and Mormotomyiidae is the only endemic, monotypic family of flies that occurs in the Afrotropical Region.

Mormotomyia was not collected subsequently until 1948, when V.G.L. and G.R.C. van Someren rediscovered the species at the same locality, associated with a vertical fissure in a large rock at the summit of Ukasi Hill (Copeland et al. 2011). On this occasion a large number of flies were collected (including the unknown females) and immature stages were sampled in bat guano that had washed out of the rock fissure. These immature stages (egg, larva and puparium) were subsequently described by van Emden (1950), who further noted that bat guano was the substrate for larval development. Despite sporadic searches, there were no further records of Mormotomyia in the wild until 2010, when the species was again discovered at the type locality (Copeland et al. 2011; Kirk-Spriggs et al. 2011).

Mormotomyia hirsuta has long been of great interest to Diptera systematists and conservation biologists (Courtney et al. 2009: 203; Kirk-Spriggs & Stuckenberg 2009: 172; Oldroyd 1964: 184), as a result of its contentious systematic position and rarity. Due to the aberrant form, reduced head and thoracic setation and wing venation, morphological taxonomists have been unable to resolve the phylogenetic relationship of Mormotomyiidae to other families of Diptera. It has variously been placed in the Calyptratae (Pont 1980: 713); or as a possible transitional family between acalyptrate and calyptrate flies, and probably closest to the Scathophagidae (as Cordyluridae) (van Emden 1950); or as related to families of acalyptrate flies, including Sphaeroceridae (as Borboridae) (Austen 1936) and Heleomyzidae (Sphaeroceroidea) (McAlpine 1989: 1484).

Detailed study of the specimens collected in 2010 allowed redescription of the third-instar larva and puparium, using stereoscan microscopy, and description of the female reproductive tract (Kirk-Spriggs et al. 2011). This study revealed that the structure of the female reproductive tract suggested that Mormotomyia should be ascribed to the acalyptrate superfamily Ephydroidea (Kirk-Spriggs et al. 2011) and later McAlpine (2011) noted that the general structure of the antenna of Mormotomyia concurred with this view. Recent advances in molecular phylogenetics (e.g. Wiegmann et al. 2011) presented an additional technique to help resolve the phylogenetic placement of Mormotomyia, and live flies, collected into 96% ethanol, for the first time were used for such an analysis (Copeland et al. 2011). Results of this study confirm placement of the Mormotomyiidae in the Ephydroidea, as sister to the remainder of the Ephydroidea, except the Ephydridae (Winkler et al, in prep.).

While clarification of the phylogenetic placement of Mormotomyiidae represents an important milestone, other issues of biological importance remain unresolved, particularly those related to reproductive and dispersal behaviours and species conservation. Questions that still need to be posed are, for example: what is the biological significance of the pronounced male-biased sexual-size dimorphism observed in the Ukasi population (Copeland et al. 2011)?; and was Mormotomyia restricted to a single relict population, with the attendant problems of managing the conservation of an endangered species (Courtney et al. 2009: 203), or were there other, as yet undiscovered, populations? Analysis of both mitochondrial and nuclear DNA of individuals from the population of flies collected in 2010 presented evidence of outbreeding, suggesting that the Ukasi flies were probably part of a metapopulation of Mormotomyia (Copeland et al. 2011). If other populations do exist, how does a fly with non-functional wings (Fig. 1) disperse? Examination of the tarsi of Mormotomyia (Kirk-Spriggs et al. 2011) revealed none of the modifications of the tarsal claws found in other bat-associated fly species known either to be phoretic on Chiroptera, i.e. Mystacinobiidae (Holloway 1976), or ectoparasitic on them, i.e. Streblidae and Nycteribiidae (Oldroyd 1964: 184). To address these questions, further investigations were conducted. In this paper the discovery of three additional populations of Mormotomyia in eastern Kenya is reported, together with additional observations on natural history.

MATERIAL AND METHODS

Definitions

For the purposes of this paper a differentiation is made between caves and fissures (rock fissures). The former refer to openings into a rock system, often relatively large, with a floor that is either continuous with ground level, or descends below it. Fissures (Fig. 2) are defined as rock fissures above ground level, usually narrow and horizontally-oriented (i.e. much wider than high), often present on vertical or near-vertical rock faces, making access difficult for mammalian and avian predators. Virtually all fissures located during this study were horizontally-oriented. As is the case with caves, fissures are commonly inhabited by bats.

Expeditions

Between 19 April 2011 and 9 January 2013 expeditions were conducted to sites in Kenya, falling approximately along a northwest-southeast transect, from coordinates 0.4501[degrees]N 36.8852[degrees]E to 4.6154[degrees]S 39.3532[degrees]E (Fig. 3). Geological formations similar to those occurring at the type locality of Mormotomyia were examined (i.e. small inselbergs/kopjes to medium-sized hills that represent remnants of ancient basement rocks). With the exception of three caves near the Kenyan coast, all were located in dry habitats. The presence of bats was indicated by the occurrence of fresh bat guano within caves or below fissures, and by aural and visual evidence. Evidence of fly presence was sought by searching bat guano on rock faces and on the ground directly below horizontal fissures and by examination of cave floors and walls, using battery operated torches (flashlights). During dry-season expeditions evidence of recent fly occurrence was sought by searching accumulated dry guano deposits for Mormotomyia corpses. On six days (2, 3, 5, 7, 8 and 9 December 2012) at Makilu and Ngauluka Hills, lizard predation of Mormotomyia was observed through binoculars between 08h30 and 09h00. Lizards were identified using Sprawls et al. (2002).

Phenology

During observations of living flies in the field, the relative population size of Mormotomyia was monitored below one fissure at the Makilu Hill site over a period of 38 days (29 November 2012 to 7 January 2013). Living flies were counted every second day, between 08h00 and 08h30, both on guano and the rock face, to a height of c. 4 m above ground level.

Laboratory observations

When Mormotomyia were active, fresh guano was collected from below a fissure on Makilu Hill and placed in two one-litre plastic containers to a depth of c. 4 cm. Containers were covered with their original plastic lids, from which a large rectangular section had been cut and replaced with fine-meshed cloth. During transport to the laboratory, cotton material was placed over the containers. Containers were placed in a 60 cm x 45 cm x 45 cm Perspex cage and their covers removed. Emerging adults were provided with cotton wool soaked in sugar water. Containers were left in the cage after the emergence of adults to serve as possible oviposition sites. Twenty adults captured at the Makilu Hill site were placed in a two-litre plastic container containing moistened paper towelling. These were also transported to the laboratory.

Identification and deposition

Newly-sampled specimens of Mormotomyia from Mbuinzau and Makilu Hills were identified by dissection and comparison of the male terminalia with specimens from the original Ukasi Hill population. These were found to be conspecific with M. hirsuta. Voucher specimens are deposited in the National Museums of Kenya, Nairobi, the International Centre of Insect Physiology and Ecology, Nairobi and the National Museum, Bloemfontein, South Africa.

RESULTS

Exploration of caves and fissures

A total of 48 sites (Fig. 3) were examined between April 2011 and January 2013. Three of these were subterranean caves; one was a large hole in a Baobab tree, Adansonia digitata L. (Malvaceae), that housed numerous fruit bats; and the remainder were rocky hills with caves and fissures. Many of these hills had multiple caves and fissures. A total of 248 caves/fissures were investigated. Some fissures and caves, particularly those in the area of the type locality (Ukasi Hill), were investigated more than once and the total number of visits to caves/fissures was 337. Ukasi is one of a small chain of hills located within 2 km of each other, the others being Ngauluka Hill and Makilu Hill (Fig. 4). These three hills were visited on 31 separate days, comprising a total of 88 investigations of caves and fissures.

Distribution of bats and flies among caves and fissures

Appendix 1 provides a list of the areas and caves/fissures examined that contained resident bat populations, including those that also housed Mormotomyia. Caves and fissures that contained neither bats nor flies when visited are listed in Appendix 2. These data were used to test the following two hypotheses: firstly, that the distribution of bats was independent of habitat type (i.e., caves and fissures), and secondly, that the distribution of Mormotomyia was also independent of habitat type. During the survey, fissures were significantly more likely than caves to harbour bats. Bats were present in 18.8% (n = 144) of unique caves and in 46.2% (n = 104) of unique fissures (Table 1). Fissures were also significantly more likely to house Mormotomyia. Flies were found in 3.8% of fissures, while they were never observed in caves (Table 2).

Exploration in the area of Ukasi Hill (the type locality)

Twenty-three hills within 20 km of the type locality were investigated (Fig. 3). Although many caves and fissures were found to have resident bat populations (Appendix 1), evidence of Mormotomyia was found on only the two hills nearest the type locality, Makilu and Ngauluka Hills (Fig. 4). Living flies were not discovered at the type locality.

Dry-season expeditions

When visited on 11 August 2012, substantial amounts of dry guano were found below Fissure 3 at Makilu Hill (Fig. 2), much of this adhering to the steep rock face near its base. Close examination revealed what appeared to be the desiccated remains of three dead flies, embedded in a matrix of dried guano. These were highly fragile and were sampled together with the guano and taken to the laboratory for cleaning and microscopical examination, where the insect remains were confirmed as that of M. hirsuta (Figs 6, 7). Similarly, during examination of fissures and caves on Ngauluka Hill on 26 September 2012, a single corpse was found, also embedded in dried guano, at the base of the rock beneath Fissure 14. During removal, however, this corpse became detached from the guano and was lost. The rock face below both Mormotomyia-positive fissures was stained purple with pinkish streaks (e.g. Fig. 2), as was the rock face below the fissure at the type locality on Ukasi Hill, where Mormotomyia was collected in 2010 (Fig. 8). This trend suggested that bat residence in fissures was associated with a characteristic discoloration of the rock face, a feature easily observed at a distance with binoculars. This visual cue was used to focus investigations more finely on fissures with similar characteristics.

Rainy season expeditions

Based on the evidence from dry-season collections of desiccated Mormotomyia corpses from Makilu and Naguluka Hills, these and the Ukasi Hill site were closely monitored following the onset of the rains, which began in November 2012. During this time populations of live flies were discovered to be active on fresh guano covering the ground beneath fissures in the rock faces on Makilu and Ngauluka Hills (Appendix 1). Guano had recently been washed out of these fissures by precipitation. Mormotomyia were not, however, found on Ukasi Hill. Shortly thereafter, searches in similar habitat, far to the south, revealed a third population on Mbuinzau Hill (Figs 3, 5, 9, 10; Appendix 1), at a distance of 187 km from the type locality. Living flies from below fissures Makilu 3, Ngauluka 16 and Mbuinzau 12 (see Appendix 1) were collected separately into 95% ethanol, providing suitable genetic material for DNA analysis and allowing investigations of whether outbreeding had occurred between the original Ukasi Hill population and the newly discovered ones.

Phenology of Mormotomyia

At the Makilu Hill site, live flies were first observed on 29 November 2012, gradually increasing in number, until peaking abruptly on 9 December 2012 when > 800 individuals were counted (Fig. 12). Live flies were last observed on 3 January 2013.

Predation on Mormotomyia

During the period that Mormotomyia were active on Makilu and Ngauluka Hills, lizards were observed attacking flies. Observations with binoculars revealed that male and female Red-headed Rock Agamas, Agama agama L. (Agamidae), and the Five-lined Skink, Trachylepis quinquetaeniata (Lichtenstein) (Scincidae), were actively feeding on Mormotomyia at the entrance to Ngauluka fissure 16 and Makilu fissure 3 (Fig. 2). Eight lizards were observed feeding at Makilu Hill on 9 December when Mormotomyia numbers were highest. Lizards were not observed to enter the fissure itself.

In addition, the expedition to Ukasi Hill on 18 December 2011 revealed a jumping spider (Araneae: Salticidae) nest c. 2 m above the ground, mostly obscured by a tiny crack in the rock face, below the large fissure that produced Mormotomyia in late 2010. The nest was removed and, while one spider escaped capture, the nest and its remaining occupants were placed in a vial containing 95% ethanol. Subsequent examination in the laboratory revealed three nymphal and one adult salticid (probably Menemerus sp.; C. Haddad pers. comm. 2014); one mostly intact corpse of Mormotomyia embedded in spider silk along with other Mormotomyia body parts, including one head and 11 legs; and the head and partial thorax of an acridid (Orthoptera) nymph.

Laboratory observations

Of the 20 adults collected in the field only one female and one male Mormotomyia survived the journey to the laboratory. The remainder were either moribund or dead. The female specimen outlived the male specimen and died after 11 days.

Four female and one male Mormotomyia emerged in the laboratory from guano collected at the base of Makilu Hill. On two separate days the single male was found to have enclosed a female within the span of his legs (Fig. 11). In neither case was the beginning of the interaction observed and it was not possible to determine whether the same, or a different female, was "guarded". Both "guarding" periods lasted at least 10 minutes. Short video footage was recorded as part of one encounter, and a single, possible attempt at copulation appears in the film between 30 to 32 seconds (pleiocarpa1 2014a). Three other short videos of the same male illustrate feeding and grooming behaviour as it sponges sugar water placed on the surface of a rock (pleiocarpa1 20146, c, d).

DISCUSSION

Exploration

Previous successful and unsuccessful expeditions to the type locality of Ukasi Hill in search of Mormotomyia suggested that the appearance of flies was unpredictable, except for an apparent relationship with heavy precipitation events, when moist guano is washed out of fissures, thus providing a suitable medium for larval development (Austen 1936; Copeland et al. 2011; Kirk-Spriggs et al. 2011; van Emden 1950). Up to now, the dark interior of these fissures has not been examined and it is possible that breeding of Mormotomyia continues uninterrupted, even during dry periods, providing bats are in residence.

During recent expeditions, conducted between 2011 and 2013, living flies were not detected at the type locality on Ukasi Hill, where they were encountered in 2010. The absence of living flies at Ukasi Hill, although discovered contemporaneously on two nearby hills, provides further evidence of the ephemeral nature of the presence of living flies. That notwithstanding, when living flies are present at a site they may be active, and readily detected, for a considerable period of time. The presence of Mormotomyia at Makilu Hill was documented for a period lasting at least 36 days (29 November 2012 to 3 January 2013), suggesting that properly timed future expeditions would allow more extensive behavioural studies of Mormotomyia to be conducted under natural field conditions.

The survey revealed three additional sites with Mormotomyia populations, making a total of four sites, one 187 km from the other three. Together, they indicate a much wider geographical distribution than was previously thought. The survey also indicated that a similar habitat profile (fissures and not caves) was common to all sites at which the fly was found, which should facilitate the location of additional Mormotomyia sites.

Fissures were significantly more likely than caves to harbour both flies and bats (Tables 1, 2). Most caves and fissures were visited infrequently, however, and the absence of bats and flies at a site examined on one or a few occasions does not necessarily imply that they are not present at other times. Additionally, access to some recesses of investigated caves was problematic, or impossible, and in general, considerably more challenging than closely examining the well-lit ground and rock faces below fissures. These factors may have affected the detection of bats and Mormotomyia in caves and, perhaps, biased the results. Nonetheless, sampling included multiple caves and fissures across an extensive geographical range and these data suggest strongly that fissures provide a preferred habitat for both Mormotomyia and the bats on which the flies depend.

Predation on Mormotomyia

Mormotomyia adults are preyed upon opportunistically by lizards and these may play a role in limiting the population size of Mormotomyia. It is likely that jumping spiders also prey opportunistically on Mormotomyia. Insects associated with the examined salticid nest included a few Mormotomyia and a nymphal acridid grasshopper. Although no predation by spiders was observed, the presence of multiple insect bodies suggests that the salticids carry insect prey back to their nest, to consume it within the safety of the crack in the rock face. Similar behaviour has been observed in another salticid species, Heliophanus termiophagus Wesolowska & Haddad, that carries its prey into the safety of tunnels within abandoned termitaria prior to consumption (Wesolowska & Haddad 2002).

Lizards were not observed entering fissures, and the association of Mormotomyia with narrow fissures on rock faces that are difficult to reach may offer considerable protection for this species, particularly against predation by small mammals. Predation of Mormotomyia by small mammals or birds was not observed, nor was there any evidence of predation on immature stages. Vertebrate spoor were not observed on, or near, the moist guano that had accumulated below the fissures.

Female-guarding

Mormotomyia exhibits pronounced male-biased sexual size dimorphism, with seven different body part measurements significantly larger (by 33-61%) in males than females (Copeland et al. 2011). Sexual size differences in insects usually favour females and often correlate with increased fecundity. Nonetheless, there are numerous instances of the reverse being true (Copeland et al. 2011, and references therein). Larger size in males may be driven by sexual selection, size being a proxy for male fitness. Larger males may be more likely to win battles for territoriality and access to females. A form of female-guarding behaviour, whereby the male stands above the female, enclosing her within the span of the legs, may also drive increases in male size (Bonduriansky 2006).

Female-guarding behaviour of this type by a Mormotomyia male was observed, as reported in some other Diptera families. Adler and Adler (1991), for example, studied three species of Tipulidae, the males of which guard females following copulation, by standing above them, normally maintaining this position until females oviposit, or until dislodged in conflicts with conspecific males. As is the case with Mormotomyia (Copeland et al. 2011), these tipulid males had longer legs than did the females, significantly so for two of the investigated species. Interestingly, in the tipulid Limonia simulans (Walker), sexual dimorphism was also observed in the shape of the last tarsal segments that in males are sinuate on their inner surface, while in females these segments are linear (Adler & Adler 1991). A similar condition occurs in Mormotomyia, although in this case the sexual dimorphism is confined to the first tarsal segment of the mid tarsus (Austen 1936; Kirk-Spriggs et al. 2011; van Emden 1950). For both species modified tarsal segments may be involved in guarding and also mating behaviour, although direct evidence for this has not been observed. Kirk-Spriggs et al. (2011) suggested that the sexually dimorphic tarsal segments of male Mormotomyia may serve such a clasping function during copulation. Post-copulatory female-guarding behaviour is also relatively common in flies of the family Neriidae (Bonduriansky 2006; Mangan 1979; Preston-Mafham 2001) and has obvious benefits in circumstances where male competition for females is high. Even if another male successfully drives off a guarding male before oviposition is completed, engaging the interloper in battle may be enough to allow the now-unprotected female to deposit all, or most of her eggs (Adler & Adler 1991).

In laboratory observations of confined Mormotomyia, events prior to the initiation of guarding were not observed. As a result, it is not possible to distinguish between post-copulatory guarding and pre-copulatory persistence of the male, in the face of one, or more, instances of rejection by the guarded female. The duration of female-guarding was considerably longer for Mormotomyia (> 10 min, n = 2) than that reported for Tipulidae, where post-copulatory guarding episodes that were not interrupted by another male lasted for 2.3 [+ or -] 0.78 min (n = 25, Dactylobis montana (Osten Sacken)) and for 0.9 [+ or -] 0.23 min (n = 44, Limonia simulans (Walker)) (Adler & Adler 1991). Mormotomyia's longer guarding period suggests that pre-copulatory guarding may have been observed. The apparently rejected attempt at mating by the male in the video footage cited above appears to support this interpretation. The recently emerged females may not have been sufficiently sexually mature to mate. Alternatively, it is conceivable that guarding behaviour may be different when other males are present. The observations of female-guarding behaviour by Mormotomyia are based on that of a single male that may have behaved differently in the presence of potential challengers. More extensive observations of non-teneral flies in the laboratory and the field are necessary to clarify the type(s) and significance of female-guarding in this species.

Conservation status of Mormotomyia

Although Mormotomyia hirsuta is stenoecious, distributional data presented here indicates that the species is more widespread than previously thought and probably not uncommon in sites resembling those in which it has already been found. Both the macrohabitat (inselbergs) and microhabitat (fissures) in which the species occurs are little affected by human activity. Similar rocky outcrops and small hills are widely distributed in the drier areas of Kenya, particularly in the eastern and northern parts of the country. It is likely that the species will also be found to occur in the expanse of Tsavo East and Tsavo West National Parks in Kenya, through to the border with Tanzania and across it into Mkomazi National Park, the southeastern-most extension of the Sahel (Coe 1999: 5). Hitherto, Mormotomyia has been considered "the rarest fly in the world" (F.C. Thompson, pers. comm. 2010). The results reported here suggest that the conservation status of Mormotomyia is robust and that no special efforts are required to ensure its continuing survival.

Future research

Adult Mormotomyia appear to be relatively fragile insects, frequently losing legs in nature and suffering high mortality while being transported from the field to the laboratory (a c. four-hour trip over mostly smooth roads). Collection of moist guano at Mormotomyia sites is easier, however, and the transport of a reasonably small amount of guano (perhaps 2-3 litres) should produce sufficient adults to study interactions between the sexes in the laboratory, including mating and guarding behaviour. Additionally (as indicated above), at certain times the number of active flies may be substantial and observations made in the field may yield useful information, although activity of adult flies below fissures appears to be limited to individuals that have recently emerged and whose behaviour is limited to ascending the rock face and entering the fissure.

The means by which Mormotomyia disperses remain unknown. For practical reasons it was not possible to examine the interior of fissures, or undertake trapping of bats to ascertain whether the flies are phoretic as adults, a possibility that appears unlikely given that Mormotomyia lacks the modified tarsal claws apparent in fly families known to be phoretic or ectoparasitic on bats (Kirk-Spriggs et al. 2011). Answering this vexing question should be the primary objective of future research.

Finally, the discovery of three additional Mormotomyia sites provides the opportunity to compare the genetic makeup of individuals among and within multiple populations of the species. Preliminary results of mitochondrial DNA-barcoding of five individuals each from Makilu, Ukasi, and Mbuinzau Hills yielded two clusters. One cluster included all five Makilu specimens and single specimens from Ukasi and Mbuinzau. The other cluster included the other four Ukasi specimens and the final four specimens from Mbuinzau Hill, nearly 200 km away (R.S.C., unpub. data). These data hint at considerable dispersal of Mormotomyia and support earlier suggestions, based on genetic analyses, that the Ukasi population is not a genetically isolated one (Copeland et al. 2011). Analysis of nuclear DNA from the three new populations is underway (W. Booth, pers. comm. 2014) and should shed light on the question of dispersal and gene exchange of this interesting fly.

Appendix I.
Presence or absence of Mormotomyia hirsuta Austen in bat-inhabited
caves and fissures in Kenya.

Prov.       Area        Site name       Cave/fissure     Cave/
                                            name        fissure

Coast      Coast      Kisimani cave       Kisimani       cave
Coast      Coast      Mdenyenye cave     Mdenyenye       cave
Coast      Coast       Pangani cave       Pangani        cave
Coast     Kasigau    Kasigau Mountain    Kasigau 2       cave
Coast     Kasigau    Kasigau Mountain    Kasigau 3       cave
Coast      Maungu      Marasi Hill        Marasi 1       cave
Eastern   Kibwesi      Kakindu Hill      Kakindu 12      cave
Eastern   Kibwesi    Kima Kimwe Hill    Kima Kimwe 3     cave
Eastern   Kibwesi    Kima Kimwe Hill    Kima Kimwe 3    fissure
Eastern   Kibwesi    Kima Kimwe Hill    Kima Kimwe 4    fissure
Eastern   Kibwesi     Kwandula Hill      Kwandula 2     fissure
Eastern   Kibwesi     Kwandula Hill      Kwandula 2     fissure
Eastern   Kibwesi     Kwandula Hill      Kwandula 5      cave
Eastern   Kibwesi     Kwandula Hill      Kwandula 6     fissure
Eastern   Kibwesi     Kwandula Hill      Kwandula 6     fissure
Eastern   Kibwesi     Kwandula Hill      Kwandula 7     fissure
Eastern   Kibwesi     Kwandula Hill      Kwandula 7     fissure
Eastern   Kibwesi     Kwandula Hill      Kwandula 8     fissure
Eastern   Kibwesi     Kwandula Hill      Kwandula 8     fissure
Eastern   Kibwesi     Kwandula Hill      Kwandula 9     fissure
Eastern   Kibwesi     Kwandula Hill      Kwandula 9     fissure
Eastern   Kibwesi      Kwayula Hill      Kwayula 1       cave
Eastern   Kibwesi      Kwayula Hill      Kwayula 2       cave
Eastern   Kibwesi       Mbale Hill        Mbale 1        cave
Eastern   Kibwesi       Mbale Hill        Mbale 4        cave
Eastern   Kibwesi     Mbuinzau Hill      Mbuinzau 5     fissure
Eastern   Kibwesi     Mbuinzau Hill      Mbuinzau 5     fissure
Eastern   Kibwesi     Mbuinzau Hill      Mbuinzau 5     fissure
Eastern   Kibwesi     Mbuinzau Hill      Mbuinzau 5     fissure
Eastern   Kibwesi     Mbuinzau Hill      Mbuinzau 5     fissure
Eastern   Kibwesi     Mbuinzau Hill      Mbuinzau 8     fissure
Eastern   Kibwesi     Mbuinzau Hill     Mbuinzau 12     fissure
Eastern   Kibwesi     Mbuinzau Hill     Mbuinzau 12     fissure
Eastern   Kibwesi     Mbuinzau Hill     Mbuinzau 12     fissure
Eastern   Kibwesi     Mbuinzau Hill     Mbuinzau 12     fissure
Eastern   Kibwesi     Mbuinzau Hill     Mbuinzau 12     fissure
Eastern   Kibwesi     Mbuinzau Hill     Mbuinzau 13     fissure
Eastern   Kibwesi     Mbuinzau Hill     Mbuinzau 24     fissure
Eastern   Kibwesi     Vendelani Hill    Vendelani 1     fissure
Eastern   Kibwesi     Vendelani Hill    Vendelani 5      cave
Eastern   Kibwesi     Vendelani Hill    Vendelani 6      cave
Eastern   Kibwesi     Vendelani Hill    Vendelani 6     fissure
Eastern   Kibwesi     Vendelani Hill    Vendelani 6     fissure
Eastern   Kibwesi     Vendelani Hill    Vendelani 7      cave
Eastern   Kibwesi     Vendelani Hill    Vendelani 7     fissure
Eastern   Kibwesi      Yamalu Hill        Yamalu 2      fissure
Eastern   Kibwesi      Yamalu Hill        Yamalu 2      fissure
Eastern   Kibwesi      Yamalu Hill        Yamalu 6      fissure
Eastern   Kibwesi      Yamalu Hill        Yamalu 6      fissure
Eastern   Kibwesi      Yamalu Hill        Yamalu 8       cave
Eastern    Kyulu        Kyulu Hill        Kyulu 1        cave
Eastern    Mwingi     Endau Mountain       Sovi 1        cave
Eastern    Mwingi     Kalanga Hill A     Kalanga A2     fissure
Eastern    Mwingi      Kangui Hill        Kangui 2      fissure
Eastern    Mwingi      Kangui Hill        Kangui 3      fissure
Eastern    Mwingi      Kangui Hill        Kangui 4       cave
Eastern    Mwingi      Kangui Hill        Kangui 6      fissure
Eastern    Mwingi     Kavuruti Hill      Kavuruti 3     fissure
Eastern    Mwingi     Kavuruti Hill      Kavuruti 3     fissure
Eastern    Mwingi     Kavuruti Hill     Kavuruti 4b     fissure
Eastern    Mwingi    Kwanduto Hill C    Kwanduto C 5     cave
Eastern    Mwingi      Makilu Hill        Makilu 2      fissure
Eastern    Mwingi      Makilu Hill        Makilu 2      fissure
Eastern    Mwingi      Makilu Hill        Makilu 2      fissure
Eastern    Mwingi      Makilu Hill        Makilu 2      fissure
Eastern    Mwingi      Makilu Hill        Makilu 2      fissure
Eastern    Mwingi      Makilu Hill        Makilu 2      fissure
Eastern    Mwingi      Makilu Hill        Makilu 2      fissure
Eastern    Mwingi      Makilu Hill        Makilu 3      fissure
Eastern    Mwingi      Makilu Hill        Makilu 3      fissure
Eastern    Mwingi      Makilu Hill        Makilu 3      fissure
Eastern    Mwingi      Makilu Hill        Makilu 3      fissure
Eastern    Mwingi      Makilu Hill        Makilu 3      fissure
Eastern    Mwingi      Makilu Hill        Makilu 3      fissure
Eastern    Mwingi      Makilu Hill        Makilu 3      fissure
Eastern    Mwingi      Makilu Hill        Makilu 3      fissure
Eastern    Mwingi      Makilu Hill        Makilu 3      fissure
Eastern    Mwingi      Makilu Hill        Makilu 3      fissure
Eastern    Mwingi      Makilu Hill        Makilu 3      fissure
Eastern    Mwingi      Makilu Hill        Makilu 3      fissure
Eastern    Mwingi      Makilu Hill        Makilu 3      fissure
Eastern    Mwingi      Makilu Hill        Makilu 3      fissure
Eastern    Mwingi      Makilu Hill        Makilu 3      fissure
Eastern    Mwingi      Makilu Hill        Makilu 3      fissure
Eastern    Mwingi      Makilu Hill        Makilu 3      fissure
Eastern    Mwingi      Makilu Hill        Makilu 3      fissure
Eastern    Mwingi      Makilu Hill        Makilu 3      fissure
Eastern    Mwingi      Makilu Hill        Makilu 9       cave
Eastern    Mwingi      Makyui Hill       Makyui B3      fissure
Eastern    Mwingi      Makyui Hill       Makyui B3      fissure
Eastern    Mwingi      Makyui Hill       Makyui B5      fissure
Eastern    Mwingi         Miuni         Miuni baobab    baobab
                                                       tree hole

Eastern    Mwingi         Miuni         Miuni baobab    baobab
                                                       tree hole

Eastern    Mwingi       Miuni Hill        Miuni 1       fissure
Eastern    Mwingi       Miuni Hill        Miuni 1       fissure
Eastern    Mwingi       Miuni Hill        Miuni 6       fissure
Eastern    Mwingi       Miuni Hill        Miuni 6       fissure
Eastern    Mwingi       Miuni Hill        Miuni 7        cave
Eastern    Mwingi       Miuni Hill        Miuni 7        cave
Eastern    Mwingi      Mulinde Hill      Mulinde 7      fissure
Eastern    Mwingi     Ngauluka Hill      Ngauluka 1     fissure
Eastern    Mwingi     Ngauluka Hill      Ngauluka 1     fissure
Eastern    Mwingi     Ngauluka Hill      Ngauluka 2     fissure
Eastern    Mwingi     Ngauluka Hill      Ngauluka 2     fissure
Eastern    Mwingi     Ngauluka Hill      Ngauluka 2     fissure
Eastern    Mwingi     Ngauluka Hill     Ngauluka 13     fissure
Eastern    Mwingi     Ngauluka Hill     Ngauluka 14     fissure
Eastern    Mwingi     Ngauluka Hill     Ngauluka 14     fissure
Eastern    Mwingi     Ngauluka Hill     Ngauluka 14     fissure
Eastern    Mwingi     Ngauluka Hill     Ngauluka 15     fissure
Eastern    Mwingi     Ngauluka Hill     Ngauluka 15     fissure
Eastern    Mwingi     Ngauluka Hill     Ngauluka 15     fissure
Eastern    Mwingi     Ngauluka Hill     Ngauluka 15     fissure
Eastern    Mwingi     Ngauluka Hill     Ngauluka 15     fissure
Eastern    Mwingi     Ngauluka Hill     Ngauluka 15     fissure
Eastern    Mwingi     Ngauluka Hill     Ngauluka 16     fissure
Eastern    Mwingi     Ngauluka Hill     Ngauluka 16     fissure
Eastern    Mwingi     Ngauluka Hill     Ngauluka 16     fissure
Eastern    Mwingi     Ngauluka Hill     Ngauluka 16     fissure
Eastern    Mwingi     Ngauluka Hill     Ngauluka 16     fissure
Eastern    Mwingi     Ngauluka Hill     Ngauluka 16     fissure
Eastern    Mwingi     Ngauluka Hill     Ngauluka 16     fissure
Eastern    Mwingi     Ngauluka Hill     Ngauluka 17     fissure
Eastern    Mwingi     Ngauluka Hill     Ngauluka 17     fissure
Eastern    Mwingi     Ngauluka Hill     Ngauluka 17     fissure
Eastern    Mwingi     Ngauluka Hill     Ngauluka 19     fissure
Eastern    Mwingi     Nzewani Hill B     Nzewani B3     fissure
Eastern    Mwingi     Nzewani Hill B     Nzewani B3     fissure
Eastern    Mwingi     Sosoma Hill A      Sosoma A1      fissure
Eastern    Mwingi     Sosoma Hill B      Sosoma B1      fissure
Eastern    Mwingi     Sosoma Hill C      Sosoma C1       cave
Eastern    Mwingi     Sosoma Hill C      Sosoma C1       cave
Eastern    Mwingi     Sosoma Hill C      Sosoma C2       cave
Eastern    Mwingi     Sosoma Hill C      Sosoma C2       cave
Eastern    Mwingi     Sosoma Hill C      Sosoma C3      fissure
Eastern    Mwingi     Sosoma Hill C      Sosoma C3      fissure
Eastern    Mwingi     Sosoma Hill C      Sosoma C4      fissure
Eastern    Mwingi     Sosoma Hill C      Sosoma C5      fissure
Eastern    Mwingi     Sosoma Hill C      Sosoma C5      fissure
Eastern    Mwingi     Sosoma Hill C      Sosoma C6      fissure
Eastern    Mwingi     Sosoma Hill C      Sosoma C7      fissure
Eastern    Mwingi       Tivai Hill        Tivai 4        cave
Eastern    Mwingi       Tivai Hill        Tivai 4        cave
Eastern    Mwingi       Ukasi Hill        Ukasi 1       fissure
Eastern    Mwingi       Ukasi Hill        Ukasi 1       fissure
Eastern    Mwingi       Ukasi Hill        Ukasi 1       fissure
Eastern    Mwingi       Ukasi Hill        Ukasi 1       fissure
Eastern    Mwingi       Ukasi Hill        Ukasi 1       fissure
Eastern    Mwingi       Ukasi Hill        Ukasi 2        cave
Eastern    Mwingi       Ukasi Hill        Ukasi 2       fissure
Eastern    Mwingi       Ukasi Hill        Ukasi 2       fissure
Eastern    Mwingi       Ukasi Hill        Ukasi 2       fissure
Eastern    Mwingi       Ukasi Hill        Ukasi 3        cave
Eastern    Mwingi       Ukasi Hill        Ukasi 3        cave
Eastern    Mwingi       Ukasi Hill        Ukasi 4       fissure
Eastern    Mwingi       Ukasi Hill        Ukasi 4       fissure
Eastern    Mwingi       Ukasi Hill        Ukasi 4       fissure
Eastern    Mwingi       Ukasi Hill        Ukasi 6        cave
Rift      Laikipia     Ewaso Nyiro      Babu's cave      cave
Valley                    River

Prov.        Date         Coordinates      Elev.      Live
                       (decimal degrees)    (m)    Mormotomyia
                                                    present?

Coast     26/7/2012    -4.61542 39.35284    20         no
Coast     26/7/2012    -4.61415 39.35420    20         no
Coast     26/7/2012    -4.61535 39.35316    20         no
Coast      8/4/2012    -3.84727 38.67464    581        no
Coast      8/4/2012    -3.84713 38.67485    597        no
Coast      8/4/2012    -3.57323 38.74625    646        no
Eastern   12/10/2012   -2.08336 38.33989    715        no
Eastern   12/10/2012   -2.06258 38.32144    726        no
Eastern   21/12/2012   -2.06258 38.32144    726        no
Eastern   12/10/2012   -2.06229 38.32131    733        no
Eastern   14/10/2012   -2.02291 38.33135    711        no
Eastern   20/12/2012   -2.02291 38.33135    711        no
Eastern   14/10/2012   -2.02141 38.33038    676        no
Eastern   14/10/2012   -2.01911 38.32964    679        no
Eastern   20/12/2012   -2.01911 38.32964    679        no
Eastern   14/10/2012   -2.01883 38.32967    681        no
Eastern   20/12/2012   -2.01883 38.32967    681        no
Eastern   14/10/2012   -2.01887 38.32968    673        no
Eastern   20/12/2012   -2.01887 38.32968    673        no
Eastern   14/10/2012   -2.01864 38.32964    661        no
Eastern   20/12/2012   -2.01864 38.32964    661        no
Eastern   17/10/2012   -2.00177 38.32431    702        no
Eastern   17/10/2012   -2.00128 38.32418    712        no
Eastern   16/10/2012   -2.00465 38.33027    672        no
Eastern   16/10/2012   -2.00367 38.33051    674        no
Eastern   30/10/2012   -2.37266 37.91180   1216        no
Eastern   19/12/2012   -2.37266 37.91180   1216        no
Eastern   20/12/2012   -2.37266 37.91180   1216        no
Eastern   21/12/2012   -2.37266 37.91180   1216        no
Eastern   22/12/2012   -2.37266 37.91180   1216        no
Eastern   30/10/2012   -2.36823 37.91150   1227        no
Eastern   1/11/2012    -2.37011 37.90958   1143        no
Eastern   19/12/2012   -2.37011 37.90958   1143        no
Eastern   20/12/2012   -2.37011 37.90958   1143        yes
Eastern   21/12/2012   -2.37011 37.90958   1143        yes
Eastern   22/12/2012   -2.37011 37.90958   1143        no
Eastern   01/11/2012   -2.36721 37.90947   1207        no
Eastern   20/12/2012   -2.37024 37.90682   1029        no
Eastern   15/10/2012   -2.01073 38.33111    651        no
Eastern   15/10/2012   -2.00970 38.33023    662        no
Eastern   15/10/2012   -2.00968 38.33059    666        no
Eastern   19/12/2012   -2.00968 38.33059    666        no
Eastern   22/12/2012   -2.00968 38.33059    666        no
Eastern   15/10/2012   -2.00975 38.33069    665        no
Eastern   19/12/2012   -2.00975 38.33069    665        no
Eastern   13/10/2012   -2.04735 38.33130    912        no
Eastern   22/12/2012   -2.04735 38.33130    912        no
Eastern   13/10/2012   -2.04797 38.33171    904        no
Eastern   22/12/2012   -2.04797 38.33171    904        no
Eastern   13/10/2012   -2.04885 38.33116    837        no
Eastern    9/5/2012    -2.92915 38.40625    593        no
Eastern   19/12/2011   -1.22799 38.56463    650        no
Eastern   15/8/2012    -0.81018 38.39499    873        no
Eastern   29/9/2012    -0.79458 38.53382    729        no
Eastern   29/9/2012    -0.79484 38.53313    737        no
Eastern   29/9/2012    -0.79477 38.53315    735        no
Eastern   29/9/2012    -0.79445 38.53371    741        no
Eastern   13/8/2012    -0.85850 38.51793    622        no
Eastern   11/12/2012   -0.85850 38.51793    622        no
Eastern   13/8/2012    -0.85851 38.51819    630        no
Eastern   16/8/2012    -0.79727 38.37453    707        no
Eastern   11/8/2012    -0.83673 38.55806    677        no
Eastern   30/11/2012   -0.83673 38.55806    677        yes
Eastern   2/12/2012    -0.83673 38.55806    677        yes
Eastern   4/12/2012    -0.83673 38.55806    677        no
Eastern   9/12/2012    -0.83673 38.55806    677        no
Eastern   13/12/2012   -0.83673 38.55806    677        no
Eastern   14/12/2012   -0.83673 38.55806    677        yes
Eastern   11/8/2012    -0.83623 38.55784    686        no1
Eastern   29/11/2012   -0.83623 38.55784    686        yes
Eastern   30/11/2012   -0.83623 38.55784    686        yes
Eastern   2/12/2012    -0.83623 38.55784    686        yes
Eastern   4/12/2012    -0.83623 38.55784    686        yes
Eastern   5/12/2012    -0.83623 38.55784    686        yes
Eastern   7/12/2012    -0.83623 38.55784    686        yes
Eastern   9/12/2012    -0.83623 38.55784    686        yes
Eastern   13/12/2012   -0.83623 38.55784    686        yes
Eastern   14/12/2012   -0.83623 38.55784    686        yes
Eastern   16/12/2012   -0.83623 38.55784    686        yes
Eastern   19/12/2012   -0.83623 38.55784    686        yes
Eastern   22/12/2012   -0.83623 38.55784    686        yes
Eastern   25/12/2012   -0.83623 38.55784    686        yes
Eastern   28/12/2012   -0.83623 38.55784    686        yes
Eastern   31/12/2012   -0.83623 38.55784    686        yes
Eastern    3/1/2013    -0.83623 38.55784    686        yes
Eastern    6/1/2013    -0.83623 38.55784    686        no
Eastern    9/1/2013    -0.83623 38.55784    686        no
Eastern   8/11/2011    -0.83642 38.55582    999        no
Eastern   30/9/2012    -0.87016 38.52322    619        no
Eastern   12/12/2012   -0.87016 38.52322    619        no
Eastern   30/9/2012    -0.87043 38.52298    639        no
Eastern   12/8/2012    -0.88113 38.55712    564        no

Eastern   6/12/2012    -0.88113 38.55712    564        no

Eastern   12/8/2012    -0.86047 38.55015    587        no
Eastern   6/12/2012    -0.86047 38.55015    587        no
Eastern   12/8/2012    -0.86056 38.54947    577        no
Eastern   6/12/2012    -0.86056 38.54947    577        no
Eastern   12/8/2012    -0.86031 38.54934    596        no
Eastern   6/12/2012    -0.86031 38.54934    596        no
Eastern   14/8/2012    -0.84208 38.43270    729        no
Eastern   10/8/2012    -0.82251 38.54564    634        no
Eastern   29/11/2012   -0.82251 38.54564    634        no
Eastern   10/8/2012    -0.82241 38.54544    619        no
Eastern   29/11/2012   -0.82241 38.54544    619        no
Eastern   10/12/2012   -0.82241 38.54544    619        no
Eastern   26/9/2012    -0.82596 38.55215    749        no
Eastern   26/9/2012    -0.82576 38.55230    752        no1
Eastern   1/12/2012    -0.82576 38.55230    752        no
Eastern   8/12/2012    -0.82576 38.55230    752        no
Eastern   26/9/2012    -0.82716 38.55269    737        no
Eastern   1/12/2012    -0.82716 38.55269    737        no
Eastern   7/12/2012    -0.82716 38.55269    737        no
Eastern   8/12/2012    -0.82716 38.55269    737        yes
Eastern   10/12/2012   -0.82716 38.55269    737        no
Eastern   13/12/2012   -0.82716 38.55269    737        no
Eastern   26/9/2012    -0.82702 38.55299    728        no
Eastern   1/12/2012    -0.82702 38.55299    728        no
Eastern   7/12/2012    -0.82702 38.55299    728        yes
Eastern   8/12/2012    -0.82702 38.55299    728        yes
Eastern   10/12/2012   -0.82702 38.55299    728        yes
Eastern   13/12/2012   -0.82702 38.55299    728        no
Eastern   14/12/2012   -0.82702 38.55299    728        yes
Eastern   8/12/2012    -0.82681 38.55320    711        no
Eastern   10/12/2012   -0.82681 38.55320    711        no
Eastern   13/12/2012   -0.82681 38.55320    711        no
Eastern   10/12/2012   -0.82693 38.55164    713        no
Eastern   28/9/2012    -0.88033 38.55646    555        no
Eastern   12/12/2012   -0.88033 38.55646    555        no
Eastern   27/9/2012    -0.88110 38.67256    503        no
Eastern   27/9/2012    -0.88130 38.67316    501        no
Eastern   27/9/2012    -0.88357 38.67567    509        no
Eastern   14/12/2012   -0.88357 38.67567    509        no
Eastern   27/9/2012    -0.88342 38.67582    535        no
Eastern   14/12/2012   -0.88342 38.67582    535        no
Eastern   27/9/2012    -0.88344 38.67584    514        no
Eastern   14/12/2012   -0.88344 38.67584    514        no
Eastern   27/9/2012    -0.88325 38.67636    525        no
Eastern   27/9/2012    -0.88321 38.67635    541        no
Eastern   14/12/2012   -0.88321 38.67635    541        no
Eastern   27/9/2012    -0.88309 38.67631    531        no
Eastern   14/12/2012   -0.88274 38.67634    504        no
Eastern   13/8/2012    -0.84618 38.52211    672        no
Eastern   11/12/2012   -0.84618 38.52211    672        no
Eastern   18/12/2011   -0.81713 38.54225    720     no (2-3)
Eastern   28/11/2012   -0.81713 38.54225    720        no
Eastern   29/11/2012   -0.81713 38.54225    720        no
Eastern   3/12/2012    -0.81713 38.54225    720        no
Eastern   9/12/2012    -0.81713 38.54225    720        no
Eastern   18/12/2011   -0.81429 38.54543    667        no
Eastern   1/10/2012    -0.81895 38.54274    664        no
Eastern   3/12/2012    -0.81895 38.54274    664        no
Eastern   9/12/2012    -0.81895 38.54274    664        no
Eastern   18/12/2011   -0.81587 38.54164    670        no
Eastern   1/10/2012    -0.81858 38.54328    707        no
Eastern   1/10/2012    -0.81731 38.54401    754        no
Eastern   3/12/2012    -0.81731 38.54401    754        no
Eastern   09/12/2012   -0.81731 38.54401    754        no
Eastern   1/10/2012    -0.81692 38.54370    774        no
Rift      19/4/2011    0.45013 36.88520    1658        no
Valley

(1) Dried corpses found, embedded in guano matrix.

(2) Mormotomyia was collected below this fissure from November to
December 2010.

(3) Mormotomyia corpses were found in the nest of a salticid spider,
located within a small crack in the rock face below this fissure

APPENDIX 2
Caves and fissures with neither Chiroptera nor Mormotomyia.

Prov.      Area        Site name          Cave/        Fissure/
                                         fissure         cave
                                           name

Coast     Kasigau       base of         Kasigau 1        cave
                        Kasigau
                       Mountain

Coast     Kasigau       base of         Kasigau 4        cave
                        Kasigau
                       Mountain

Coast     Kasigau       base of         Kasigau 5        cave
                        Kasigau
                       Mountain

Coast     Kasigau       base of         Kasigau 6        cave
                        Kasigau
                       Mountain

Eastern   Kibwesi    Kakindu Hill       Kakindu 1      fissure
Eastern   Kibwesi    Kakindu Hill       Kakindu 2      fissure
Eastern   Kibwesi    Kakindu Hill       Kakindu 3        cave
Eastern   Kibwesi    Kakindu Hill       Kakindu 4        cave
Eastern   Kibwesi    Kakindu Hill       Kakindu 5        cave
Eastern   Kibwesi    Kakindu Hill       Kakindu 6        cave
Eastern   Kibwesi    Kakindu Hill       Kakindu 7        cave
Eastern   Kibwesi    Kakindu Hill       Kakindu 8        cave
Eastern   Kibwesi    Kakindu Hill       Kakindu 9        cave
Eastern   Kibwesi    Kakindu Hill       Kakindu 10       cave
Eastern   Kibwesi    Kakindu Hill       Kakindu 11       cave
Eastern   Kibwesi    Kanziko Hill       Kanziko 1        cave
Eastern   Kibwesi    Kanziko Hill       Kanziko 2        cave
Eastern   Kibwesi    Kanziko Hill       Kanziko 3        cave
Eastern   Kibwesi    Kanziko Hill       Kanziko 4        cave
Eastern   Kibwesi    Kanziko Hill       Kanziko 5        cave
Eastern   Kibwesi    Kanziko Hill       Kanziko 6      fissure
Eastern   Kibwesi    Kanziko Hill       Kanziko 7        cave
Eastern   Kibwesi   Kima Kimwe Hill    Kima Kimwe 1      cave
Eastern   Kibwesi   Kima Kimwe Hill    Kima Kimwe 2    fissure
Eastern   Kibwesi   Kima Kimwe Hill    Kima Kimwe 5    fissure
Eastern   Kibwesi    Kwandula Hill      Kwandula 1     fissure
Eastern   Kibwesi    Kwandula Hill      Kwandula 3     fissure
Eastern   Kibwesi    Kwandula Hill      Kwandula 4       cave
Eastern   Kibwesi    Kwandula Hill     Kwandula 10     fissure
Eastern   Kibwesi    Kwayula Hill       Kwayula 3        cave
Eastern   Kibwesi    Kwayula Hill       Kwayula 4        cave
Eastern   Kibwesi    Kwayula Hill       Kwayula 5      fissure
Eastern   Kibwesi     Mbale Hill         Mbale 2         cave
Eastern   Kibwesi     Mbale Hill         Mbale 3         cave
Eastern   Kibwesi     Mbale Hill         Mbale 5         cave
Eastern   Kibwesi     Mbale Hill         Mbale 6         cave
Eastern   Kibwesi     Mbale Hill         Mbale 7         cave
Eastern   Kibwesi     Mbale Hill         Mbale 8         cave
Eastern   Kibwesi     Mbale Hill         Mbale 9       fissure
Eastern   Kibwesi     Mbale Hill         Mbale 10      fissure
Eastern   Kibwesi    Mbuinzau Hill      Mbuinzau 1       cave
Eastern   Kibwesi    Mbuinzau Hill      Mbuinzau 2     fissure
Eastern   Kibwesi    Mbuinzau Hill      Mbuinzau 3       cave
Eastern   Kibwesi    Mbuinzau Hill      Mbuinzau 4       cave
Eastern   Kibwesi    Mbuinzau Hill      Mbuinzau 6       cave
Eastern   Kibwesi    Mbuinzau Hill      Mbuinzau 7       cave
Eastern   Kibwesi    Mbuinzau Hill      Mbuinzau 9       cave
Eastern   Kibwesi    Mbuinzau Hill     Mbuinzau 10       cave
Eastern   Kibwesi    Mbuinzau Hill     Mbuinzau 11       cave
Eastern   Kibwesi    Mbuinzau Hill     Mbuinzau 14     fissure
Eastern   Kibwesi    Mbuinzau Hill     Mbuinzau 15     fissure
Eastern   Kibwesi    Mbuinzau Hill     Mbuinzau 16     fissure
Eastern   Kibwesi    Mbuinzau Hill     Mbuinzau 17       cave
Eastern   Kibwesi    Mbuinzau Hill     Mbuinzau 18     fissure
Eastern   Kibwesi    Mbuinzau Hill     Mbuinzau 19       cave
Eastern   Kibwesi    Mbuinzau Hill     Mbuinzau 20     fissure
Eastern   Kibwesi    Mbuinzau Hill     Mbuinzau 21     fissure
Eastern   Kibwesi    Mbuinzau Hill     Mbuinzau 22       cave
Eastern   Kibwesi    Mbuinzau Hill     Mbuinzau 23     fissure
Eastern   Kibwesi    Miusyani Hill      Miusyani 1     fissure
Eastern   Kibwesi    Miusyani Hill      Miusyani 2     fissure
Eastern   Kibwesi    Miusyani Hill      Miusyani 3     fissure
Eastern   Kibwesi    Miusyani Hill      Miusyani 4       cave
Eastern   Kibwesi    Miusyani Hill      Miusyani 5     fissure
Eastern   Kibwesi    Miusyani Hill      Miusyani 6     fissure
Eastern   Kibwesi    Miusyani Hill      Miusyani 7       cave
Eastern   Kibwesi   Syokivulu Hill     Syokivulu 1     fissure
Eastern   Kibwesi   Syokivulu Hill     Syokivulu 2     fissure
Eastern   Kibwesi   Syokivulu Hill     Syokivulu 3     fissure
Eastern   Kibwesi   Syokivulu Hill     Syokivulu 4     fissure
Eastern   Kibwesi   Syokivulu Hill     Syokivulu 5       cave
Eastern   Kibwesi   Syokivulu Hill     Syokivulu 6     fissure
Eastern   Kibwesi   Syokivulu Hill     Syokivulu 7       cave
Eastern   Kibwesi   Syokivulu Hill     Syokivulu 8       cave
Eastern   Kibwesi   Syokivulu Hill     Syokivulu 9     fissure
Eastern   Kibwesi   Syokivulu Hill     Syokivulu 10    fissure
Eastern   Kibwesi   Vendelani Hill     Vendelani 2       cave
Eastern   Kibwesi   Vendelani Hill     Vendelani 3     fissure
Eastern   Kibwesi   Vendelani Hill     Vendelani 4       cave
Eastern   Kibwesi   Vendelani Hill     Vendelani 8     fissure
Eastern   Kibwesi   Vendelani Hill     Vendelani 9     fissure
Eastern   Kibwesi     Yamalu Hill        Yamalu 1        cave
Eastern   Kibwesi     Yamalu Hill        Yamalu 3        cave
Eastern   Kibwesi     Yamalu Hill        Yamalu 4        cave
Eastern   Kibwesi     Yamalu Hill        Yamalu 5        cave
Eastern   Kibwesi     Yamalu Hill        Yamalu 7        cave
Eastern   Mwingi     Ethombe Hill     Ethombe Hill 1     cave
Eastern   Mwingi     Ethombe Hill     Ethombe Hill 2     cave
Eastern   Mwingi    Kalanga Hill A      Kalanga A1     fissure
Eastern   Mwingi    Kalanga Hill A      Kalanga A3     fissure
Eastern   Mwingi    Kalanga Hill A      Kalanga A4       cave
Eastern   Mwingi    Kalanga Hill A      Kalanga A5       cave
Eastern   Mwingi    Kalanga Hill A      Kalanga A6       cave
Eastern   Mwingi    Kalanga Hill A      Kalanga A7       cave
Eastern   Mwingi    Kalanga Hill B      Kalanga B1       cave
Eastern   Mwingi    Kalanga Hill B      Kalanga B2       cave
Eastern   Mwingi      Kangui Hill        Kangui 1        cave
Eastern   Mwingi      Kangui Hill        Kangui 5        cave
Eastern   Mwingi      Kangui Hill        Kangui 7      fissure
Eastern   Mwingi      Kangui Hill        Kangui 8        cave
Eastern   Mwingi      Kangui Hill        Kangui 9        cave
Eastern   Mwingi     Kavuruti Hill      Kavuruti 1     fissure
Eastern   Mwingi     Kavuruti Hill      Kavuruti 2       cave
Eastern   Mwingi     Kavuruti Hill     Kavuruti 4A       cave
Eastern   Mwingi    Kwanduto Hill A    Kwanduto A1     fissure
Eastern   Mwingi    Kwanduto Hill A    Kwanduto A2       cave
Eastern   Mwingi    Kwanduto Hill B    Kwanduto B1       cave
Eastern   Mwingi    Kwanduto Hill C    Kwanduto C1       cave
Eastern   Mwingi    Kwanduto Hill C    Kwanduto C2       cave
Eastern   Mwingi    Kwanduto Hill C    Kwanduto C3       cave
Eastern   Mwingi    Kwanduto Hill C    Kwanduto C4       cave
Eastern   Mwingi    Kwanduto Hill D    Kwanduto D1       cave
Eastern   Mwingi    Kwanduto Hill D    Kwanduto D2       cave
Eastern   Mwingi    Kwanduto Hill D    Kwanduto D3       cave
Eastern   Mwingi      Makilu Hill        Makilu 1      fissure
Eastern   Mwingi      Makilu Hill        Makilu 4        cave
Eastern   Mwingi      Makilu Hill        Makilu 5        cave
Eastern   Mwingi      Makilu Hill        Makilu 6        cave
Eastern   Mwingi      Makilu Hill        Makilu 7        cave
Eastern   Mwingi      Makilu Hill        Makilu 8        cave
Eastern   Mwingi     Makyui Hill A      Makyui A1        cave
Eastern   Mwingi     Makyui Hill A      Makyui A2        cave
Eastern   Mwingi     Makyui Hill A      Makyui A3        cave
Eastern   Mwingi     Makyui Hill B      Makyui B1        cave
Eastern   Mwingi     Makyui Hill B      Makyui B2        cave
Eastern   Mwingi     Makyui Hill B      Makyui B4        cave
Eastern   Mwingi     Mathini Hill       Mathini 1        cave
Eastern   Mwingi     Mathini Hill       Mathini 2      fissure
Eastern   Mwingi      Miuni Hill         Miuni 2         cave
Eastern   Mwingi      Miuni Hill         Miuni 3       fissure
Eastern   Mwingi      Miuni Hill         Miuni 4         cave
Eastern   Mwingi      Miuni Hill         Miuni 5       fissure
Eastern   Mwingi     Mulinde Hill       Mulinde 1        cave
Eastern   Mwingi     Mulinde Hill       Mulinde 2        cave
Eastern   Mwingi     Mulinde Hill       Mulinde 3      fissure
Eastern   Mwingi     Mulinde Hill       Mulinde 4        cave
Eastern   Mwingi     Mulinde Hill       Mulinde 5        cave
Eastern   Mwingi     Mulinde Hill       Mulinde 6      fissure
Eastern   Mwingi     Mulinde Hill       Mulinde 8        cave
Eastern   Mwingi     Ngauluka Hill      Ngauluka 3     fissure
Eastern   Mwingi     Ngauluka Hill      Ngauluka 4     fissure
Eastern   Mwingi     Ngauluka Hill      Ngauluka 5       cave
Eastern   Mwingi     Ngauluka Hill      Ngauluka 6       cave
Eastern   Mwingi     Ngauluka Hill      Ngauluka 7     fissure
Eastern   Mwingi     Ngauluka Hill      Ngauluka 8     fissure
Eastern   Mwingi     Ngauluka Hill      Ngauluka 9     fissure
Eastern   Mwingi     Ngauluka Hill     Ngauluka 10       cave
Eastern   Mwingi     Ngauluka Hill     Ngauluka 11       cave
Eastern   Mwingi     Ngauluka Hill     Ngauluka 12       cave
Eastern   Mwingi     Ngauluka Hill     Ngauluka 18       cave
Eastern   Mwingi    Nzewani Hill B      Nzewani B1       cave
Eastern   Mwingi    Nzewani Hill B      Nzewani B2     fissure
Eastern   Mwingi    Nzewani Hill C      Nzewani C1     fissure
Eastern   Mwingi    Nzewani Hill D      Nzewani D1     fissure
Eastern   Mwingi    Nzewani Hill E      Nzewani E1       cave
Eastern   Mwingi    Nzewani Hill E      Nzewani E2       cave
Eastern   Mwingi     Sosoma Hill A      Sosoma A2        cave
Eastern   Mwingi     Tiumboni Hill      Tiumboni 1       cave
Eastern   Mwingi     Tiumboni Hill      Tiumboni 2     fissure
Eastern   Mwingi      Tivai Hill         Tivai 1       fissure
Eastern   Mwingi      Tivai Hill         Tivai 2         cave
Eastern   Mwingi      Tivai Hill         Tivai 3         cave
Eastern   Mwingi      Tivai Hill         Tivai 5         cave
Eastern   Mwingi      Tivai Hill         Tivai 6         cave
Eastern   Mwingi      Tivai Hill         Tivai 7         cave
Eastern   Mwingi      Tivai Hill         Tivai 8         cave
Eastern   Mwingi      Tivai Hill         Tivai 9         cave
Eastern   Mwingi      Tivai Hill         Tivai 10      fissure
Eastern   Mwingi      Ukasi Hill         Ukasi 1         cave
Eastern   Mwingi      Ukasi Hill         Ukasi 5         cave
Eastern   Mwingi      Ukasi Hill         Ukasi 7         cave
Eastern   Mwingi     unnamed Hill                        cave
Eastern   Mwingi     unnamed Kopje                       cave
                     west of Ukasi

Prov.        Date         Coordinates      Elev.
                       (decimal degrees)    (m)

Coast      8/4/2012    -3.84724 38.67684    629

Coast      8/4/2012    -3.84646 38.68218    653

Coast      8/4/2012    -3.84629 38.68196    666

Coast      8/4/2012    -3.84621 38.68162    663

Eastern   11/10/2012   -2.07396 38.33571    796
Eastern   11/10/2012   -2.07356 38.33587    834
Eastern   11/10/2012   -2.07349 38.33589    833
Eastern   11/10/2012   -2.07335 38.33587    839
Eastern   11/10/2012   -2.07319 38.33575    848
Eastern   11/10/2012   -2.07332 38.33591    866
Eastern   11/10/2012   -2.07333 38.33571    832
Eastern   11/10/2012   -2.07488 38.33629    759
Eastern   11/10/2012   -2.07619 38.33603    715
Eastern   12/10/2012   -2.08290 38.34098    756
Eastern   12/10/2012   -2.08315 38.34038    792
Eastern   17/10/2012   -2.00080 38.32550    713
Eastern   17/10/2012   -1.99869 38.32583    737
Eastern   17/10/2012   -1.99846 38.32587    728
Eastern   17/10/2012   -1.99823 38.32585    725
Eastern   17/10/2012   -1.99802 38.32580    712
Eastern   17/10/2012   -1.99829 38.32516    680
Eastern   17/10/2012   -1.99851 38.32526    691
Eastern   12/10/2012   -2.06292 38.32174    694
Eastern   12/10/2012   -2.06263 38.32138    720
Eastern   12/10/2012   -2.06225 38.32119    744
Eastern   14/10/2012   -2.02398 38.32894    683
Eastern   14/10/2012   -2.02286 38.33139    721
Eastern   14/10/2012   -2.02272 38.33134    702
Eastern   14/10/2012   -2.01764 38.32897    643
Eastern   17/10/2012   -2.00106 38.32397    706
Eastern   17/10/2012   -2.00071 38.32389    716
Eastern   17/10/2012   -2.00025 38.32307    685
Eastern   16/10/2012   -2.00446 38.33051    664
Eastern   16/10/2012   -2.00367 38.33088    720
Eastern   16/10/2012   -2.00368 38.33016    691
Eastern   16/10/2012   -2.00325 38.32997    714
Eastern   16/10/2012   -2.00309 38.32973    716
Eastern   16/10/2012   -2.00326 38.32955    699
Eastern   16/10/2012   -2.00207 38.32915    700
Eastern   16/10/2012   -2.00260 38.33023    678
Eastern   30/10/2012   -2.37274 37.90781   1013
Eastern   30/10/2012   -2.37267 37.90971   1092
Eastern   30/10/2012   -2.37361 37.91168   1147
Eastern   30/10/2012   -2.37359 37.91176   1162
Eastern   30/10/2012   -2.37290 37.91229   1238
Eastern   30/10/2012   -2.37079 37.91155   1239
Eastern   30/10/2012   -2.36967 37.91082   1238
Eastern   30/10/2012   -2.36901 37.91003   1236
Eastern   30/10/2012   -2.37248 37.90983   1110
Eastern   1/11/2012    -2.36527 37.91017   1225
Eastern   1/11/2012    -2.36369 37.91051   1231
Eastern   1/11/2012    -2.36325 37.91050   1214
Eastern   1/11/2012    -2.36221 37.90979   1165
Eastern   3/11/2012    -2.37412 37.91677   1088
Eastern   3/11/2012    -2.37417 37.91540   1141
Eastern   3/11/2012    -2.37547 37.91558   1123
Eastern   3/11/2012    -2.37596 37.91427   1146
Eastern   3/11/2012    -2.37851 37.91420   1055
Eastern   3/11/2012    -2.37942 37.91420   1025
Eastern   10/10/2012   -2.08115 38.21447    773
Eastern   10/10/2012   -2.08137 38.21422    799
Eastern   10/10/2012   -2.08119 38.21420    802
Eastern   10/10/2012   -2.08109 38.21411    794
Eastern   10/10/2012   -2.08035 38.21337    818
Eastern   10/10/2012   -2.08006 38.21323    825
Eastern   10/10/2012   -2.07997 38.21319    827
Eastern   2/11/2012    -2.38710 37.91752   1054
Eastern   2/11/2012    -2.38727 37.91777   1059
Eastern   2/11/2012    -2.38760 37.91805   1076
Eastern   2/11/2012    -2.39007 37.91826   1097
Eastern   2/11/2012    -2.39031 37.91854   1080
Eastern   2/11/2012    -2.39212 37.91894   1064
Eastern   2/11/2012    -2.39243 37.91725   1037
Eastern   2/11/2012    -2.39003 37.91654   1056
Eastern   2/11/2012    -2.38770 37.91729   1039
Eastern   2/11/2012    -2.38707 37.91659    998
Eastern   15/10/2012   -2.01075 38.33102    652
Eastern   15/10/2012   -2.01047 38.33075    648
Eastern   15/10/2012   -2.01016 38.33043    659
Eastern   15/10/2012   -2.00978 38.33082    660
Eastern   15/10/2012   -2.01034 38.33148    661
Eastern   13/10/2012   -2.04794 38.33131    910
Eastern   13/10/2012   -2.04749 38.33166    923
Eastern   13/10/2012   -2.04779 38.33169    913
Eastern   13/10/2012   -2.04785 38.33169    919
Eastern   13/10/2012   -2.04887 38.33147    883
Eastern   20/4/2011    -0.87742 38.09740   1148
Eastern   20/4/2011    -0.88003 38.09830   1124
Eastern   15/8/2012    -0.81272 38.39620    762
Eastern   15/8/2012    -0.81133 38.39447    793
Eastern   15/8/2012    -0.81169 38.39437    789
Eastern   15/8/2012    -0.81180 38.39421    780
Eastern   15/8/2012    -0.81173 38.39402    770
Eastern   15/8/2012    -0.81207 38.39373    742
Eastern   15/8/2012    -0.80575 38.38523    736
Eastern   15/8/2012    -0.80453 38.38404    748
Eastern   29/9/2012    -0.79422 38.53416    717
Eastern   29/9/2012    -0.79476 38.53317    737
Eastern   29/9/2012    -0.79392 38.53353    749
Eastern   29/9/2012    -0.79368 38.53318    719
Eastern   29/9/2012    -0.79331 38.53275    680
Eastern   13/8/2012    -0.85716 38.52114    662
Eastern   13/8/2012    -0.85959 38.52099    643
Eastern   13/8/2012    -0.85851 38.51819    630
Eastern   16/8/2012    -0.78889 38.36361    663
Eastern   16/8/2012    -0.78884 38.36356    661
Eastern   16/8/2012    -0.79583 38.37329    691
Eastern   16/8/2012    -0.79711 38.37458    695
Eastern   16/8/2012    -0.79722 38.37455    702
Eastern   16/8/2012    -0.79724 38.37461    705
Eastern   16/8/2012    -0.79725 38.37445    713
Eastern   16/8/2012    -0.79607 38.37098    691
Eastern   16/8/2012    -0.79606 38.37092    688
Eastern   16/8/2012    -0.79617 38.37092    679
Eastern   11/8/2012    -0.83690 38.55811    680
Eastern   11/8/2012    -0.83535 38.58826    651
Eastern   11/8/2012    -0.83324 38.55795    649
Eastern   11/8/2012    -0.83473 38.55265    609
Eastern   12/8/2012    -0.83894 38.55846    670
Eastern   12/8/2012    -0.83855 38.55812    698
Eastern   30/9/2012    -0.87019 38.52430    607
Eastern   30/9/2012    -0.87027 38.52449    611
Eastern   30/9/2012    -0.86962 38.52448    611
Eastern   30/9/2012    -0.86978 38.52344    618
Eastern   30/9/2012    -0.86999 38.52328    628
Eastern   30/9/2012    -0.87057 38.52283    630
Eastern   30/9/2012    -0.90326 38.49417    576
Eastern   30/9/2012    -0.90361 38.49396    579
Eastern   12/8/2012    -0.86020 38.55010    582
Eastern   12/8/2012    -0.85887 38.54932    591
Eastern   12/8/2012    -0.86062 38.54931    582
Eastern   12/8/2012    -0.86052 38.54952    581
Eastern   14/8/2012    -0.84257 38.43364    665
Eastern   14/8/2012    -0.84242 38.43357    691
Eastern   14/8/2012    -0.84264 38.43279    700
Eastern   14/8/2012    -0.84232 38.43269    705
Eastern   14/8/2012    -0.84201 38.43250    708
Eastern   14/8/2012    -0.84201 38.43275    732
Eastern   14/8/2012    -0.84193 38.43505    689
Eastern   10/8/2012    -0.82223 38.54538    628
Eastern   10/8/2012    -0.82196 38.54612    675
Eastern   10/8/2012    -0.82291 38.54604    636
Eastern   10/8/2012    -0.82112 38.55160    649
Eastern   10/8/2012    -0.82122 38.55200    646
Eastern   10/8/2012    -0.82481 38.55333    646
Eastern   26/9/2012    -0.82312 38.54681    644
Eastern   26/9/2012    -0.82425 38.55124    754
Eastern   26/9/2012    -0.82485 38.55102    768
Eastern   26/9/2012    -0.82543 38.55152    749
Eastern   10/12/2012   -0.82688 38.55154    701
Eastern   28/9/2012    -0.88006 38.55631    552
Eastern   28/9/2012    -0.88022 38.55677    553
Eastern   28/9/2012    -0.88067 38.55747    550
Eastern   28/9/2012    -0.88202 38.55713    559
Eastern   28/9/2012    -0.87752 38.55538    551
Eastern   28/9/2012    -0.87747 38.55520    552
Eastern   27/9/2012    -0.88098 38.67161    502
Eastern   14/8/2012    -0.79809 38.49091    646
Eastern   14/8/2012    -0.79816 38.49090    648
Eastern   13/8/2012    -0.84853 38.52280    674
Eastern   13/8/2012    -0.84828 38.52303    707
Eastern   13/8/2012    -0.84625 38.52219    672
Eastern   13/8/2012    -0.84584 38.52203    662
Eastern   13/8/2012    -0.84318 38.52161    649
Eastern   13/8/2012    -0.84118 38.52415    676
Eastern   13/8/2012    -0.84126 38.52449    692
Eastern   13/8/2012    -0.84131 38.52458    705
Eastern   13/8/2012    -0.84115 38.52470    707
Eastern   1/10/2012    -0.81947 38.54282    625
Eastern   1/10/2012    -0.81717 38.54421    776
Eastern   1/10/2012    -0.81680 38.54399    767
Eastern   21/4/2011    -0.94212 38.08205    996
Eastern   18/12/2011   -0.79808 38.49096    660


ACKNOWLEDGEMENTS

This research was funded through a grant (to R.S.C), from the Mohamed Bin Zayed Species Conservation Fund. We are grateful to P.M. Kabiro of ICIPE's Earth Observation Unit who contributed the map in Fig. 3. We thank Chief B. Musoo for permission to work in the Ukasi area. C.M. Maithya, J. Muriuki, M. Musingila, K. Ngalu, M.K. Kirk-Spriggs and M. Kodheki assisted in the field.

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Robert S. Copeland (1) *, Josephat Bukhebi (2) and Ashley H. Kirk-Spriggs (3)

(1) International Centre of Insect Physiology and Ecology, P.O. Box 30772, Nairobi 00100, and Division of Invertebrate Zoology, National Museums of Kenya, P.O. Box 40658, Nairobi, (00100) Kenya; rcopeland@icipe.org

(2) international Centre of Insect Physiology and Ecology, P.O. Box 30772, Nairobi, 00100 Kenya; jbukhebi@icipe.org

(3) Department of Entomology, National Museum, P.O. Box 266, Bloemfontein 9300 and Honorary Research Fellow, School of Life Sciences, University of KwaZulu-Natal, Private Bag X01, Scottsville, 3209 South Africa; ashley.kirk-spriggs@nasmus.co.za

* Corresponding author

http://africaninvertebrates.org

urn:lsid:zoobank.org:pub:C26A287C-ADAC-49F5-A011-17DB4A5B9D0E

TABLE 1
Relationship of caves and fissures to the presence of bats
(Chi-square = 20.22; p < 0.001).

Presence/absence     Caves   Fissures

Chiroptera present    27        48
Chiroptera absent     117       56

TABLE 2
Relationship of caves and fissures to the presence of Mormotomyia
(p = 0.03, Fisher's Exact Test).

Presence/absence       Caves   Fissures

Mormotomyia present      0        4
Mormotomyia absent      144      100
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Author:Copeland, Robert S.; Bukhebi, Josephat; Kirk-Spriggs, Ashley H.
Publication:African Invertebrates
Article Type:Report
Geographic Code:6KENY
Date:Dec 1, 2014
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