Diversity
Taphozous
is comprised of 14 species split between 2 subgenera (Wilson & Reeder, 2005). However,
the necessity of the subgenus
Liponycteris
has been called into question in recent literature (Uvizl et al., 2019). Although
many of the species have stable populations,
Taphozous hildegardeae
is a notable exception, being endangered (Webala et al., 2020).
Taphozous
species are terrestrial insectivores that are found in a wide variety of habitats,
ranging from arid deserts to humid rainforests (Patterson & Webala, 2012; Wilson &
Reeder, 2005). The presence of a glandular sac on the wing is an identifying character
of this genus, for which the common name sac-winged refers to (Hester & Myers, 2001;
Patterson & Webala, 2012).
Geographic Range
Taphozous
species can be found prominently throughout the Ethiopian and Oriental regions, with
limited distribution in the Palearctic and Australian regions as well (Wilson & Reeder,
2005). There are no known introduced populations.
Some species have heavily restricted ranges compared to other members of
Taphozous
.
Taphozous hildegardeae
is endemic to a small portion of the coastline along Kenya and Tanzania, with populations
also having been found on the nearby Zanzibar Island (Webala et al., 2020).
Taphozous australis
has a similarly small range, only occupying the northeast coast of Queensland (Armstrong,
2021).
- Biogeographic Regions
- palearctic
- oriental
- ethiopian
- australian
Habitat
Taphozous
species can be found in a diverse range of habitats, including deserts, savannas,
wetlands, coastlines, tropical rainforests, dry forests, caves, and urban areas (Armstrong,
2021; Monadjem et al., 2017a, 2017b; Webala et al., 2020). Roosting sites among species
are variable, but cliff faces, rock crevices, tree trunks, and in the case of coastal
species, sea caves, are most favorable (Armstrong, 2021; Monadjem, 2017b). Many
Taphozous
species also roost in close proximity to humans, preferring various structures ranging
from tombs to tall, inhabited brick buildings with areas such as exposed beams to
escape to when disturbed (Monadjem et al., 2017a, 2017b; O'Malley et al., 2020). Preferable
foraging habitats include open water and woodland areas, with some species traveling
as far as 15 km from their roosts to hunt (Armstrong, 2021).
- Habitat Regions
- tropical
- terrestrial
- Terrestrial Biomes
- desert or dune
- savanna or grassland
- forest
- rainforest
- scrub forest
- Aquatic Biomes
- coastal
- Wetlands
- swamp
Systematic and Taxonomic History
Current research suggests Taphozous rests within Emballonuridae , under the subfamily Taphozoinae (Wilson & Reeder, 2005). Taphozous is currently split into two subgenera, Taphozous and Liponycteris (Wilson & Reeder, 2005). However, the validity of the subgenus Liponycteris is currently being debated (Uvizl et al., 2019).
Taphozous
formerly included
Saccolaimus
as a subgenus, which was then elevated to genus status in 1991 (Chimimba & Kitchener,
1991). Currently,
Saccolaimus
is recognized as the sister taxon to
Taphozous
(Uvizl et al., 2019).
Physical Description
Taphozous species are medium to large sized bats with a forearm length ranging from 56 mm to 79 mm (Colket & Wilson, 1998; Patterson & Webala, 2012). Body and head length can range from 64 mm to 112 mm, with significant variation depending on species and location (Dengis, 1996; Harrison, 1958). Color variation includes solid browns, blacks, and tans, or in the case of Taphozous mauritianus , a unique "grizzled" patterning comprised of brown, gray, and white (Dengis, 1996). Countershading is present in some species, such as T. mauritianus and Taphozous hildegardeae (Dengis, 1996; Kingdon et al., 2018). All members of the genus possess a glandular sac on the wing (Hester & Myers, 2001; Patterson & Webala, 2012).
Sexual dimorphism in
Taphozous
is limited, however males in some species possess notable gular pouches or glands
(Muñoz-Romo & Page, 2021).
Members of the subgenus
Taphozous
have notable concavity in the frontal regions of their skull, and their bodies are
fully covered in fur (Colket & Wilson, 1998). On the contrary, members of
Liponycteris
show little concavity, with the posterior region of their bodies lacking fur (Colket
& Wilson, 1998).
- Other Physical Features
- endothermic
- bilateral symmetry
- Sexual Dimorphism
- sexes alike
Reproduction
The mating systems among most
Taphozous
species are ill-defined.
Taphozous hildegardeae
is notably polygynous, with dominant males maintaining harems of females within their
territories (Colket & Wilson, 1998). Males display territorial behavior towards other
males, and will scent mark females within their harems (Colket & Wilson, 1998).
- Mating System
- polygynous
Taphozous species reproduction is seemingly dependent on seasonality, such as wet versus dry seasons, and geography, with breeding occurring year-round or with defined breeding seasons depending on the species and location (Colket & Wilson, 1998; Dengis, 1996; Smith et al., 2013). Many species are polyestrous in some parts of their geographic range, and then monoestrous in others (Dengis, 1996; Harrison, 1958).
Depending on the above factors, birthing can take place between February-June and October-December (Dengis, 1996; Harrison, 1958; Kingdon et al., 2013). Sperm storage and delayed fertilization is exhibited by some species, such as Taphozous nudiventris , in which sperm from September-October is stored over hibernation and then used for fertilization in March (Kingdon et al., 2013). The gestation period in this species is around 9 weeks (Kingdon et al., 2013). Taphozous melanopogon demonstrates a longer gestation period, ranging around 120-125 days (Smith et al., 2013). There is limited information on gestation periods for other Taphozous species.
Across all species, typically a single pup will be born, but twins are also possible
(Smith et al., 2013). Some polyestrous species may have two pups in a year, in separate
litters (Lim et al., 2018; Skinner & Chimimba, 2005).
- Key Reproductive Features
- iteroparous
- seasonal breeding
- year-round breeding
- gonochoric/gonochoristic/dioecious (sexes separate)
- sexual
- viviparous
- sperm-storing
- delayed fertilization
Taphozous
species are born altricial, with pups relying solely on their mothers for survival
(Kingdon et al., 2013).
Taphozous nudiventris
pups will cling onto their mothers until they are ready to fly independently, with
estimates around 8 weeks old (Dengis, 1996; Kingdon et al., 2013). There are variable
reports as to whether the pup continues to accompany the mother for those entire 8
weeks, or if at some point it begins to stay back at the roost while she forages (Kingdon
et al., 2013). Weaning begins around week 6, with the pup continuing to stay close
to its mother when foraging far from the roost (Kingdon et al., 2013). There is limited
information for the extent of parental investment in other
Taphozous
species.
- Parental Investment
- altricial
- female parental care
-
pre-weaning/fledging
-
provisioning
- female
-
protecting
- female
-
provisioning
-
pre-independence
-
provisioning
- female
-
protecting
- female
-
provisioning
Lifespan/Longevity
The longevity of Taphozous species is currently unknown.
Behavior
There is notable variation in social behaviors among Taphozous species. Taphozous perforatus , Taphozous melanopogon , and Taphozous nudiventris are gregarious, and are known to roost with hundreds and up into thousands of other individuals (Kingdon et al., 2013; Smith et al., 2013). Conversely, Taphozous mauritianus is more solitary, with as little as 5 individuals roosting together on average, but colonies of up to 30 have been observed (Skinner & Chimimba, 2005).
Most Taphozous species leave their caves soon after sunset to hunt, although T. mauritianus has been observed taking off in the daytime (Dengis, 1996; Skinner & Chimimba, 2005; Wei et al., 2008). T. melanopogon has been found to depart the roost at the same exact time regardless of the season, resulting in departure times ranging from sunset to after dark (Nowak, 1994). Many species stain the walls of their roosts with urine and gular sac secretions (Dengis, 1996; Smith et al., 2013).
Hibernation behaviors among Taphozous species varies depending on the species and location. Taphozous longimanus and T. melanopogon are suspected to hibernate based on fat accumulation during different parts of the year, but further investigation is needed (Nowak, 1994; Singh, Krishna, & Bhatnagar, 2002; Wang 2004). Taphozous nudiventris individuals in Iraq hibernate during the winter, while those in Pakistan spend the season in torpor (Kingdon et al., 2013). There is limited information on the hibernation habits of other Taphozous species.
Migratory patterns for most
Taphozous
species are ill-defined, but
T. nudiventris
has been observed to be a seasonal migrator (Kingdon et al., 2013).
- Key Behaviors
- flies
- diurnal
- nocturnal
- motile
- migratory
- hibernation
- daily torpor
- territorial
- social
- colonial
Communication and Perception
Taphozous species are known to have good eyesight, which plays an important role in navigation (Dengis, 1996; Skinner & Chimimba, 2005). Taphozous mauritianus in particular relies on vision while roosting in the daytime, and seems to avoid dark areas (Dengis, 1996; Skinner & Chimimba, 2005).
Many Taphozous species exhibit shrill sounds when injured, or may call out to other individuals while flying (Nowak, 1994). Some species are also know to emit social calls while roosting (Dengis, 1996).
Echolocation has been observed in many Taphozous species. Taphozous melanopogon exhibits low intensity, long duration calls for detecting long distance prey (Wei et al., 2008). Conversely, T. mauritianus is able to emit high intensity calls, although its vocalizations varies depending on its habitat (Dengis, 1996). Taphozous nudiventris is known to echolocate as well, but details on its vocalizations are limited (Kelbel, 2006). Taphozous species are currently understood to be oral echolocators (Dengis, 1996).
Some
Taphozous
species communicate through scent marking as well, such as
Taphozous hildegardeae
,
T. nudiventris
, and
T. melanopogon
, although the purpose in some of these species is unclear (Colket & Wilson, 1998;
Kelbel, 2006; Wang, 2004).
- Other Communication Modes
- scent marks
- Perception Channels
- visual
- tactile
- acoustic
- echolocation
- chemical
Food Habits
Taphozous species are insectivorous, feeding primarily on moths (Dengis, 1996; Kingdon et al., 2013; Rydell & Yalden, 1996). Most species feed on termites, such as Macrotermes falciger , beetles, crickets, ants, and other flying insects (Dengis, 1996; Kingdon et al., 2013; Rydell & Yalden, 1996). Taphozous mauritianus in particular has been found to also feed on butterflies in the daytime (Skinner & Chimimba, 2005).
Taphozous
species use various methods to hunt, ranging from using their eyesight in the daytime,
to echolocating at night (Dengis, 1996; Skinner & Chimimba, 2005).
- Primary Diet
- carnivore
Predation
Taphozous species are primarily preyed upon by other flying animals such as owls and hawks, although predation by snakes has been reported (Dengis, 1996; Kingdon et al., 2013). Barn owls in particular have been found to prey on Taphozous nudiventris (Qumsiyeh, 1985).
Taphozous
species avoid predation primarily by roosting in well protected areas, flying quickly
and at high altitudes, and by doing most of their hunting at night (Dengis, 1996;
Wang, 2004). Their good eyesight also aids them in identifying predators in the daytime
(Dengis, 1996; Skinner & Chimimba, 2005).
- Anti-predator Adaptations
- cryptic
Ecosystem Roles
Taphozous species are effective at controlling pest populations (Fenton, Bell, & Thomas, 2011; Wang, 2004).
Many
Taphozous
species serve as the hosts for a variety of parasites, including species from the
Anchitrema
genus,
Trypanosoma longiflagellum
,
Lecithodendrium duboisi
,
Prosthodendrium parvouterus
,
Plagiorchis koreanus
, and
Pycnoporus heteroporus
. (Dengis, 1996; Kelbel, 2006).
Economic Importance for Humans: Positive
Taphozous
species are prolific in controlling pests that prove harmful to humans, namely termites
and species that negatively impact crops (Dengis, 1996; Kelbel, 2006; Kingdon et al.,
2013; Wang, 2004).
- Positive Impacts
- controls pest population
Economic Importance for Humans: Negative
Taphozous
species are capable of transmitting lyssaviruses to humans, such as rabies (Nguyen
et al., 2014). There are no other known adverse effects of
Taphozous
species on humans.
- Negative Impacts
- injures humans
Conservation Status
Most Taphozous species aren't threatened, but there are a few exceptions. Taphozous hildegardeae is endangered, with populations declining due to habitat loss (Webala et al., 2020). Although there are some populations that may reside in protected areas in Tanzania, larger colonies in Kenya mostly inhabit caves on unprotected land, which makes their roosts vulnerable to disturbance (Webala et al., 2020).
Taphozous australis
is considered near threatened as a result of development destroying and disturbing
their roosts, as well as mining activity (Armstrong, 2021). Populations have been
slowly declining in the southern portion of their range, but northern populations
are currently considered stable (Armstrong, 2021).
Additional Links
Contributors
Theo Leger (author), Colorado State University, Tanya Dewey (editor), University of Michigan-Ann Arbor.
- Palearctic
-
living in the northern part of the Old World. In otherwords, Europe and Asia and northern Africa.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- oriental
-
found in the oriental region of the world. In other words, India and southeast Asia.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- Ethiopian
-
living in sub-Saharan Africa (south of 30 degrees north) and Madagascar.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- Australian
-
Living in Australia, New Zealand, Tasmania, New Guinea and associated islands.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- tropical
-
the region of the earth that surrounds the equator, from 23.5 degrees north to 23.5 degrees south.
- terrestrial
-
Living on the ground.
- desert or dunes
-
in deserts low (less than 30 cm per year) and unpredictable rainfall results in landscapes dominated by plants and animals adapted to aridity. Vegetation is typically sparse, though spectacular blooms may occur following rain. Deserts can be cold or warm and daily temperates typically fluctuate. In dune areas vegetation is also sparse and conditions are dry. This is because sand does not hold water well so little is available to plants. In dunes near seas and oceans this is compounded by the influence of salt in the air and soil. Salt limits the ability of plants to take up water through their roots.
- tropical savanna and grassland
-
A terrestrial biome. Savannas are grasslands with scattered individual trees that do not form a closed canopy. Extensive savannas are found in parts of subtropical and tropical Africa and South America, and in Australia.
- savanna
-
A grassland with scattered trees or scattered clumps of trees, a type of community intermediate between grassland and forest. See also Tropical savanna and grassland biome.
- temperate grassland
-
A terrestrial biome found in temperate latitudes (>23.5° N or S latitude). Vegetation is made up mostly of grasses, the height and species diversity of which depend largely on the amount of moisture available. Fire and grazing are important in the long-term maintenance of grasslands.
- forest
-
forest biomes are dominated by trees, otherwise forest biomes can vary widely in amount of precipitation and seasonality.
- rainforest
-
rainforests, both temperate and tropical, are dominated by trees often forming a closed canopy with little light reaching the ground. Epiphytes and climbing plants are also abundant. Precipitation is typically not limiting, but may be somewhat seasonal.
- scrub forest
-
scrub forests develop in areas that experience dry seasons.
- coastal
-
the nearshore aquatic habitats near a coast, or shoreline.
- swamp
-
a wetland area that may be permanently or intermittently covered in water, often dominated by woody vegetation.
- urban
-
living in cities and large towns, landscapes dominated by human structures and activity.
- suburban
-
living in residential areas on the outskirts of large cities or towns.
- polygynous
-
having more than one female as a mate at one time
- iteroparous
-
offspring are produced in more than one group (litters, clutches, etc.) and across multiple seasons (or other periods hospitable to reproduction). Iteroparous animals must, by definition, survive over multiple seasons (or periodic condition changes).
- seasonal breeding
-
breeding is confined to a particular season
- year-round breeding
-
breeding takes place throughout the year
- sexual
-
reproduction that includes combining the genetic contribution of two individuals, a male and a female
- viviparous
-
reproduction in which fertilization and development take place within the female body and the developing embryo derives nourishment from the female.
- sperm-storing
-
mature spermatozoa are stored by females following copulation. Male sperm storage also occurs, as sperm are retained in the male epididymes (in mammals) for a period that can, in some cases, extend over several weeks or more, but here we use the term to refer only to sperm storage by females.
- delayed fertilization
-
a substantial delay (longer than the minimum time required for sperm to travel to the egg) takes place between copulation and fertilization, used to describe female sperm storage.
- altricial
-
young are born in a relatively underdeveloped state; they are unable to feed or care for themselves or locomote independently for a period of time after birth/hatching. In birds, naked and helpless after hatching.
- female parental care
-
parental care is carried out by females
- diurnal
-
- active during the day, 2. lasting for one day.
- nocturnal
-
active during the night
- motile
-
having the capacity to move from one place to another.
- migratory
-
makes seasonal movements between breeding and wintering grounds
- hibernation
-
the state that some animals enter during winter in which normal physiological processes are significantly reduced, thus lowering the animal's energy requirements. The act or condition of passing winter in a torpid or resting state, typically involving the abandonment of homoiothermy in mammals.
- territorial
-
defends an area within the home range, occupied by a single animals or group of animals of the same species and held through overt defense, display, or advertisement
- social
-
associates with others of its species; forms social groups.
- colonial
-
used loosely to describe any group of organisms living together or in close proximity to each other - for example nesting shorebirds that live in large colonies. More specifically refers to a group of organisms in which members act as specialized subunits (a continuous, modular society) - as in clonal organisms.
- visual
-
uses sight to communicate
- acoustic
-
uses sound to communicate
- chemical
-
uses smells or other chemicals to communicate
- scent marks
-
communicates by producing scents from special gland(s) and placing them on a surface whether others can smell or taste them
- visual
-
uses sight to communicate
- tactile
-
uses touch to communicate
- acoustic
-
uses sound to communicate
- echolocation
-
The process by which an animal locates itself with respect to other animals and objects by emitting sound waves and sensing the pattern of the reflected sound waves.
- chemical
-
uses smells or other chemicals to communicate
- cryptic
-
having markings, coloration, shapes, or other features that cause an animal to be camouflaged in its natural environment; being difficult to see or otherwise detect.
- causes disease in humans
-
an animal which directly causes disease in humans. For example, diseases caused by infection of filarial nematodes (elephantiasis and river blindness).
- carnivore
-
an animal that mainly eats meat
- insectivore
-
An animal that eats mainly insects or spiders.
- endothermic
-
animals that use metabolically generated heat to regulate body temperature independently of ambient temperature. Endothermy is a synapomorphy of the Mammalia, although it may have arisen in a (now extinct) synapsid ancestor; the fossil record does not distinguish these possibilities. Convergent in birds.
- bilateral symmetry
-
having body symmetry such that the animal can be divided in one plane into two mirror-image halves. Animals with bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends. Synapomorphy of the Bilateria.
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