Gold mole

from Wikipedia, the free encyclopedia
Gold mole
Giant gold mole (Chrysospalax trevelyani)

Giant gold mole ( Chrysospalax trevelyani )

Systematics
Class : Mammals (Mammalia)
Subclass : Higher mammals (Eutheria)
Superordinate : Afrotheria
without rank: Afroinsectiphilia
Order : Tenrecus (Afrosoricida)
Family : Gold mole
Scientific name
Chrysochloridae
Gray , 1825

The golden mole (Chrysochloridae) are a family of mammals living in southern and partly in central Africa . The animals resemble moles in their physique and also lead a similar underground way of life, but are not closely related to them.

The body shows special adaptations to a digging way of life. It is designed in the shape of a spindle to a diamond shape, externally visible ears and a tail are missing, the eyes are hidden under the fur. The front limbs in particular are particularly suitable for digging. They are short and sturdy and have large grave claws on their hands. The forearm has a third bone, the number of finger and carpal bones is reduced by characteristic adhesions. A characteristic feature of the golden mole is its silky fur, which sometimes shines metallic when exposed to certain light.

Goldmulle inhabit open to desert regions and closed forests, they can be found in both low and highlands. Due to their burrowing way of life, they represent habitat specialists who sometimes only colonize narrow habitats that are mostly bound to loose subsoil. More precise information is only available in a few cases about the way of life of the animals. They are nocturnal and solitary.

The golden mole's diet consists mainly of invertebrates such as earthworms and insects . Greatly enlarged ossicles enable them to track down the prey using the vibrations they generate . What is striking is the unstable body temperature, which adapts to the environment and therefore has an energy-saving function. Reproduction has so far been little studied, one to three young animals are born per litter, which remain as nestlings in the underground burrow. The individual development is probably very slow.

The tribal history of the family is sparse. But it goes back to the Middle Eocene around 47 million years ago. With exceptions, fossil finds only include individual skull and dentition remains. Gold moles have been known since the 18th century, but they were initially equated with moles. It was not until the turn of the 19th century that the group was separated, with the family being named in 1825.

Due to a similar way of life and comparable body characteristics, the golden mole was considered a member of the insect eater order (lipotyphla) until the end of the 20th century . More recent molecular genetic studies place them, together with the tenreks, in a completely separate group of mammals, the tenre-kitties (Afrosoricida) within the Afrotheria . The family currently comprises ten genera with 21 species, of which more than two thirds are endangered to varying degrees.

description

Habitus

The golden mole are relatively small representatives of the mammals . One of the smallest forms is the desert gold mole ( Eremitalpa granti ) with a head-trunk length of 7.6 to 8.5 cm, while the giant gold mole ( Chrysospalax trevelyani ) is 20.8 to 23.5 cm long. The body weight fluctuates accordingly between 15 and 40 g or between 410 and 500 g. Most representatives, however, have overall lengths of 9 to 14 cm and a weight of 20 to 100 g. The individual species show strong similarities in body structure, but differ in some cases considerably in terms of their fur color, which, despite the surname of the golden mole, not only appears golden yellow, but also varies between black, gray or brownish tones. The body of the gold mole is adapted to a burrowing, underground way of life. Outwardly it resembles that of the moles , with which the golden mole are not closely related. The compact trunk shows a spindle-shaped to diamond-shaped structure, a tail is not externally visible. The nose is covered by a coarse, leather-like pad that is used to support the digging. The nostrils lie in a fold of skin on the underside of the tip of the nose. Like many other burrowing mammals, the golden mole has no outer auricles, and the eyes are covered with fur. Accentuating light colored spots often appear on the face and especially in the area of ​​the eyes. The limbs are short and strong and tend to lie below the trunk, in contrast to the moles, in which they are attached to the side. The forelimbs have four toes - the outer fifth ray is reduced - and are transformed into digging tools. The claw on the middle finger (ray III) is greatly enlarged and can be longer than the forearm. On the first and second finger (ray I and II) the gold mulle each have a shorter, pointed claw, the length of which varies depending on the species. Only the desert gold mole also has a small, claw-like nail on the markedly shortened fourth finger (ray IV). The hind legs end in five toes, which are equipped with small, sharp claws. The individual toes are connected to one another by small membranes, the sole of the foot is hairless. Females have two pairs of teats .

Skull and dentition features

Top (A, B) and side (C) skull view of the yellow gold mole ( Calcochloris obtusirostris ; A), Stuhlmann's gold mole ( Chrysochloris stuhlmanni ; B) and the desert gold mole ( Eremitalpa granti ; C). The arrow in B points to the "temporal bladder", the arrow in C to the enlarged head of the hammer. In A the "temporal bladder" is missing, in C the cavity in the middle ear is enlarged and contains the extended head of the hammer.

The head is conical to teardrop-shaped, the greatest width is in the area of ​​the brain skull. The rostrum is usually elongated. The ratio of the greatest width of the skull to the greatest length varies from species to species. With long and narrow skulls, the greatest skull width only reaches 60% of the greatest skull length, with short and wide skulls the corresponding value is up to 96%. The particularly narrow-headed forms include the copper gold mole ( Amblysomus ), Arends' gold mole ( Carpitalpa ), the giant gold mole ( Chrysospalax ) or the representatives of the genera Neamblysomus and Chlorotalpa . On the other hand, the cape gold mole ( Chrysochloris ), the desert gold mole or the members of the genus Cryptochloris belong to the broad-headed mole . The rostrum is accordingly designed differently in strength and can be narrow (with a palate width of around 28% of the greatest length of the skull) or wide (with a corresponding value of up to 35%). It is noticeable that there are no constrictions in the area of ​​the eyes in the gold mole. The zygomatic arches are completely closed, but the zygomatic bone has receded and the arch consists of a bone process of the upper jaw that articulates with the temporal bone . A special feature of the gold mullets is a joint connection between the lower jaw and the hyoid bone . The articulation is between the angular process of the lower jaw and the stylohyal, which is part of the hyoid bone hanging device. A similar articulation is not known from other higher mammals. It may help the tongue chew the food.

Another abnormality can be found in the area of ​​the middle ear . In some species, the ossicles are greatly enlarged. This applies above all to the hammer (malleus), the head of which can be inflated like a ball or elongated like a club. The magnifications are sometimes enormous: in the case of the desert gold mole (body weight around 40 g) the malleus can reach a weight of 70 mg, in the case of the giant gold mole (body weight around 500 g) up to 185 mg (in comparison: in humans with an average body weight of 70 kg) the malleus 28 mg). In some cases, the enlargement of the hammer is accompanied by an increase in bone density, which in the desert gold mole reaches a value of 2.44 g / cm³. This represents one of the highest values ​​in terrestrial mammals and is only exceeded by a few whales and manatees . To accommodate the inflated hammer head, another sinus is formed on the upper side of the tympanic cavity , which in some species bulges outwardly as a bone blister over the temporal pit behind the orbit . The bony bladder is known as the temporal bulla due to its location on the temporal bone . In addition, the cochlea has a complex structure and has at least four turns with a combined combined angle of 1200 °, which is more than in numerous other underground mammals. The special characteristics of both the ossicles and the cochlea are related to the auditory perception of the golden mole. The gold mole can perceive very low-frequency tones of less than a few hundred Hertz , and they are also able to receive seismic oscillations and vibrations , for example from the movements of prey. The transmission takes place by means of bone conduction , which is triggered by the strongly asymmetrical shape of the malleus due to the enlarged head and the associated shift of the center of gravity. The ability is more pronounced in animals with an enlarged hammer than in those with a normally built hammer.

Lower jaw of the Congo gold mole ( Huetia leucorhina ; A) and Arends' gold mole ( Carpitalpa arendsi ; B). The arrow in B shows the talonid on the fourth premolar in Arends' Goldmull, which is present in the entire lower row of teeth, but is absent on the premolars and molars of the Congo Goldmull.

The Goldmulle have a slightly reduced set of 36 to 40 teeth. The front set of teeth is complete, only the back teeth have receded. The dental formula is thus: . The foremost incisor in the upper row of teeth and the second incisor in the lower jaw are enlarged, the other incisors have a canine-like and small shape. The anterior premolar can be shaped variably - sectorial (with sharp cutting edges), bicuspid or tricuspid (with two or three cusps) types occur. The molars themselves have strikingly high ( hypsodontic ) crowns and a tricuspid occlusal surface pattern. The three main cusps represent the paraconus, the metaconus and the protoconus (based on the upper jaw teeth). The protoconus is often, but not in all species, greatly reduced in size, the metaconus and paraconus are close together - the latter forms the main cusp of the molars. In addition, a V-shaped shear bar (ectoloph) runs over the tooth surface, the tip of which is the paraconus. Due to the overall configuration, the tooth structure can be viewed as typical zalambdodont . In some species, a talonid is formed on the lower molars, a deep protrusion of the chewing surface, in which the protoconus of the upper molars engages when the bite is closed. If the rearmost molar occurs, it is mostly very small and either has the shape of the other molars or resembles a pin. The characteristic can, however, be strongly overprinted by the chewing of the teeth.

Skeletal features

The spine is made up of 7 cervical , 16 thoracic , 3 lumbar , 5 sacrum and 9 tail vertebrae . The cervical spine is curved downwards. The spinous processes of the first twelve thoracic vertebrae are vertical, those of the posterior thoracic and lumbar vertebrae point backwards. Of the sixteen pairs of ribs, eight are free-standing. The front ones , connected to the sternum and the sternum itself, are directed inwards, creating space for the complex foreleg muscles.

Above all, the anterior musculoskeletal system shows special adaptations to the digging way of life. The entire shoulder girdle is shifted forward so that the shoulders are roughly level with the back of the head. The body's center of gravity, which is shifted forward in this way, allows the animals to carry out a powerful forward movement when digging. The shoulder blade and sternum are markedly elongated; the former runs roughly parallel to the spine and is also extremely narrow. The greatly enlarged shoulder bone serves as the starting point for the massive shoulder muscles. There is a collarbone , but it lacks the typical curved shape. Due to the upstream position on the body, it absorbs some energy when digging. The humerus stands out due to its short, compact shape and twisted shaft. A massive deltopectoral ridge runs along this, to which parts of the arm and shoulder muscles attach. The elbow joint is strongly asymmetrical; here the laterally strongly protruding inner (medial) bone protrusion of the humerus (epicondylus medialis) stands out. As a result, the width of the lower joint end of the humerus can be 65% and more of the total bone length. The degree of formation of the medial epicondyle depends on the intensity of the excavation activity of the individual species. Likewise, the olecranon , the upper articular process of the ulna , is strongly drawn out and noticeably curved. It makes up about 35% of the total bone length. The extensor muscles of the forearm attach to both joint processes, but both bony protrusions also prevent excessive lateral movements of the arm. What should be emphasized is the formation of a rod-shaped bone, which in a sense represents a third forearm bone. It probably originated from the ossification of a tendon of a forearm muscle, possibly the flexor digitorum profundus muscle , which moves the middle finger. For this reason, it is often referred to as the "flexor" bone. The “flexor” bone lies below the ulna and is about the size of the spoke . The number of carpal bones and phalanx is reduced. The carpal bones are characterized by a flattened shape, which limits the ability of the hand to rotate. Of the four rays of the hand, the first, second and fourth consist of a metacarpal bone and two phalanges , the front phalanx being a result of the fusions of the first and second phalanx. The hypertrophied third ray represents a fusion of the metacarpal bone and the first two phalanxes and is therefore sometimes also called the “triplex” bone. Another characteristic formation is present on the second finger beam. Here the corresponding metacarpal bone is fused with the large polygonal bone and the small polygonal bone to form the so-called trapezium-trapezoid-metacarpal-II complex (also called "ttm" or "totem" bone). The individual bone adhesions develop in the course of embryonic development . The last phalanx is enlarged and split at the end, which indicates the existence of the claw. An exception is the first finger (thumb), the last link of which is not notched.

Compared to the front limbs, there are only a few characteristic features on the rear limbs. The tibia and fibula are fused together above and in the lower third. In between there is a wide space due to the clear curvature of the shafts of both bones. Likewise, the first two phalanges of rays II to IV are fused together, so that all five foot rays each have only two toe phalanges.

fur

The coat consists of the top coat and a thick undercoat . It is mostly silky and soft. The guide hairs are directed backwards and are water-repellent. The individual hairs of the cover coat are between 7 and 21 mm long, the diameter is 78 to 190 μm. They are narrower at the base than in the upper third. In cross-section, they often have a flat to pressed shape in the upper third, and a rounded shape at the base, the hair scales show a wavy arrangement, the medulla is structured like a grid. In some species there is a metallic sheen that ranges from reddish to yellowish, greenish, brownish to silver. The shine arises in the flattened, upper third of the hair, where very flat scales are formed in several layers. The light thus hits a large flat surface for reflection and is refracted several times due to the overlapping layers. Iridescent colors in living beings often offer an advantage in partner recruitment during reproduction; however, this can be ruled out in the case of the Goldmulls due to their blind nature. The shimmer of color may be a side effect caused by the flattening and thicker layering of the hair surfaces. What caused these hair changes is unknown, but they may be related to underground locomotion.

Soft tissue anatomy

The digestive tract is simple and tubular. It has a weight of 4.9 g and a length of 43.9 cm in medium-sized species such as the Hottentot gold mole rat ( Amblysomus hottentotus ). In the case of the giant gold mole, it weighs 24.5 g. The stomach takes up a total of 11 to 20% of the length of the digestive tract, the latter itself making up between 9 and 12% of the total body mass. As with many other insectivorous mammals, there is no appendix , which makes it difficult to separate the large and small intestines . The area of ​​the gastric gate (pylorus) expands very far. The entire section of the intestine is covered with thread-like extensions ( microvilli ). The urogenital system ends in a single opening, the cloaca , as in the monotone . In the males, the testicles are located in the abdominal cavity, their total weight is about 23 mg. The penis is relatively short and measures only 3 to 4 mm, penis spines are missing. The acrosome of the sperm has receded, but small barbs are formed on the head. The females have a two-horned uterus ( uterus bicornis ). The cloaca and the testicles located inside the body represent similarities with the tenreks and can be interpreted as an indication of the relationship between the two taxa. The kidneys are simply built and have a relatively large medulla and large medullary cones that extend into the ureter . Both of these suggest that the kidneys can concentrate urine to a high degree. In medium-sized species, the brain reaches a volume of 700 to 736 mg and is therefore only about half the size of similarly large elephants .

The eyes of the gold mole have receded and are hidden under the skin. In adult animals, however, there is still an eyeball that is around 0.5 mm long and 0.4 mm deep. The conjunctival sac and the lacrimal glands as well as the lacrimal ducts are also developed, but the lens , the iris and the entire eye muscles are missing . Possibly the lacrimal system in the gold mole has the same function as in other mammals and keeps the conjunctival tissue free of foreign bodies. Overall, the eye is further developed backwards than comparable with the moles, but not quite as strong as with the pouch moles . In the gold molluscs, the design of the eye begins in the embryonic stage in the same way as in other mammals , but there is hardly any further development and growth. At least five muscles are involved in the structure of the nose, some of which are attached to the zygomatic arch and extend to the tip of the nose. Despite the short and wide shape of the golden mole's nose, it is very mobile and can function as a tactile organ with which the animals search for food and dig.

distribution and habitat

The golden moles are exclusively in Africa south of the Sahara is home, focusing its range in southern Africa is. More than half of the species are endemic to the Republic of South Africa ; some representatives also inhabit areas in Namibia , Lesotho and Mozambique . Outside of southern Africa, only three species have been recorded so far: the Congo gold mole ( Huetia leucorhina ) in central , Stuhlmann's gold mole ( Chrysochloris stuhlmanni ) in central and eastern Africa and the Somalia gold mole ( Huetia tytonis ) in northeastern Africa. Because of their subterranean way of life, the gold mole are habitat specialists . As a result, their habitats are highly fragmented or narrowly limited. Often the individual species can only be identified at a few, locally narrowly limited sites - very few representatives are known from a larger distribution area. Other gold mole such as the Somalia gold mole, but also Visagie's gold mole ( Chrysochloris visagiei ), De Winton's gold mole ( Cryptochloris wintoni ) or Van Zyl's gold mole ( Cryptochloris zyli ) have so far only been detected in a few specimens.

The different types of gold mole can be divided into two different ecological groups:

The animals are widespread both in coastal lowlands and in high mountains at altitudes of up to 4000 m. Basic prerequisites for the presence of gold mole rats are, in addition to an adequate supply of food, also loose, penetrable soils. Rocky landscapes and rivers form the limits of their expansion. The ability to adapt to landscapes changed by humans is different for the individual species. If some forms occur in a common region, such as the Fynbos gold mole ( Amblysomus corriae ) and Duthie's gold mole ( Chlorotalpa duthieae ) as well as the highveld gold mole ( Amblysomus septentrionalis ) and Sclaters gold mole ( Chlorotalpa sclateri ) exist usually different biotope demands .

way of life

Congo gold mole ( Huetia leucorhina )
Yellow gold mole ( Calcochloris obtusirostris )
Wire-haired gold mole rat ( Chrysospalax villosus )

In general, relatively little is known about the way of life of the gold mole. This is due on the one hand to the shy and subterranean way of life and on the other hand to the rarity of many species. Individual aspects of the way of life have only been researched in more detail in a few species such as the desert gold mole , the Hottentot gold mole , the giant gold mole or Stuhlmann's gold mole and the Cape gold mole ; further information is often based on random observations.

Activity times, digging and social behavior

The gold mole live largely underground. Since digging underground is around 26 times more expensive than moving above ground, the gold mole has developed some energy-saving properties. The majority of the species are nocturnal, a few can also be seen during the day. The activity time often depends on the ambient temperature. The reason for this is the animals' poor ability to maintain a stable body temperature. The average body temperature is 29 to 33 ° C and is therefore very low. Since the gold mole does not thermoregulate when it is at rest and the thermal conductivity is high, the body temperature can fluctuate greatly depending on the activity. For example, it varies from 27 to 38 ° C for the Hottentot gold mole, 26 to 36 ° C for the Cape gold mole and 18 to 32 ° C for the desert gold mole. There are optimal conditions for greater activity at outside temperatures in a range of the middle 20 degrees to middle 30 degrees, which roughly corresponds to the thermo-neutral zone. Outside of this temperature range, the body temperature fluctuates considerably and is sometimes only a few degrees above that of the surrounding soil; in the case of the desert gold mole, this difference is only 0.7 ° C. At very high and very low outside temperatures, some species fall into a torpor (rigid state), such as the Hottentot, Cape and desert gold mole. In addition, the golden mole has a comparatively low metabolic rate , which is significantly lower than that of comparably large insectivorous mammals. The metabolism increases with falling outside and body temperature, but drops again rapidly when entering the torpor. The low metabolic rate in connection with the thermolability (fluctuating body temperature) leads to an adaptive hypothermia , with which the animals are able to adapt their body temperature to the ambient temperature in order to save energy. In this sense, the frequent nocturnal lifestyle of the golden mole does not primarily serve to avoid encounters with predators , but at least for the inhabitants of dry habitats it represents an adaptation to an energy-saving way of life.

In contrast to the moles, which do their digging by rotating the upper arm to the side, golden moles are paw diggers and dig themselves forward with forward arm movements. An excavation cycle consists of a pressing phase with the head and the entire body, during which the soil substrate is compacted. This is followed by several shovel-like digging movements. The enlarged claws act like a pickaxe, the loose soil material is pushed under the belly and then scraped away with the hind legs. This process of digging is repeated cyclically; the Hottentot gold mole, for example, can dig 4 to 12 m a day, depending on the nature of the subsoil. Many species create passages on two levels: the animals look for food in tunnels just below the surface of the earth, these often exist only temporarily and are superficially recognizable as narrow ripples. Deeper corridors have individual chambers, which are often connected by tunnels and sometimes lined with plant material such as grasses like nests. They serve as resting places and to raise the young. The desert gold mole, whose habitat consists of sand dunes, on the other hand - as far as is known - does not build permanent corridors, which is due to the loose dune sand. The unstable burial tunnels usually collapse behind the animals, giving the impression that they are “swimming” through the sand. Some species are known to sometimes share burrows with beach graves , gray mulls or lamellar tooth rats . This could be a symbiotic behavior, since the golden mole does not compete with these animals for food and so the burial effort of the individual animal is reduced.

Most gold mole live solitary and are territorial, each animal has its own territory. Encounters between two animals often result in violent fights, which are fought by wrestling with the strong front paws or with bites. They emit high-pitched squeaking noises. The own building is vehemently defended against conspecifics of the same and sometimes of the opposite sex, but the territories can overlap at the edges. Only in the case of the Giant Gold Mole, based on unconfirmed reports of observations of several animals that overwintered in a burrow, is a more social behavior assumed.

nourishment

The food of the golden mole consists mainly of invertebrates such as earthworms as well as insects and their larvae. There are also hundred- and millipedes , woodlice , molluscs and spiders . Some species also ingest small vertebrates such as skinks . Foraging takes place both above and below ground. The development of the hearing and thus the hammer in the middle ear probably plays a major role here. Species with a distended malleus come to the surface more often than those with a normally built one. The former can perceive more low-frequency sounds and better locate vibrations, which helps them to find food, but also to avoid encounters with predators. Many species leave their burrows after heavy rainfall and look for something edible on the surface of the earth.

Due to the low metabolic rate and efficient kidney function , which means that urine can be concentrated in water, most species do not need to drink any fluids. This ability is particularly developed in species that live in dry habitats such as the golden mole rat. Further factors for reducing water loss are the nocturnal and subterranean way of life.

Reproduction

Little is known about the reproduction of the golden mole. For species that live in regions with strongly seasonal climatic conditions, there is a fixed mating season, for others it can take place all year round. The females of at least some species can have several litters a year, but most of the births occur in the rainy season. During courtship , the males of the Hottentot gold mole chirp, raise and lower their heads and stamp their feet on the ground. Under laboratory conditions, it was possible to observe how the males pursue the females in order to force mating, which can also lead to deaths in the females. Since the animals are blind, the search for a willing mating partner is probably carried out via calls and secretion markings . The female may choose her partner based on the length of the penis . The lack of penile spines in males means that females ovulate spontaneously .

The length of the gestation period is unknown. Usually between one and three (usually two) young animals are born. You will initially remain in a chamber that is laid out with grass. The newborns are nestled and initially hairless and have soft claws. There is hardly any data available on individual development. Corresponding observations with the giant gold mole suggest that this occurs very slowly. It is assumed from the Cape Gold Mole that the young remain in the mother's den for two to three months. The very late eruption of the permanent teeth, which only occurs shortly before reaching adulthood, is also noteworthy. Information on the life expectancy of wild animals is not available.

Natural enemies

One of the most important natural enemies of the golden mole is the barn owl , in whose crusts remains of the animals are regularly found. Other predators make gripping and crows and snakes . Among the mammals that are African striped weasel or backed jackal to call. House cats and dogs often kill individual animals, but sometimes without eating them. The animals are always alerted, especially on the surface of the earth, and flee to the nearest burrow if there is little threat. Members of some species play dead if they are touched, such as the wire-haired gold mole rat .

Systematics

External system

Internal systematics of Afrotheria according to Kuntner et al. 2011
 Afrotheria  
  Paenungulata  

 Sirenia (manatees)


   

 Hyracoidea (hyrax)


   

 Proboscidea ( Proboscidea )




  Afroinsectiphilia  

 Tubulidentata (aardvark)


   

 Macroscelidea (elephant)


  Afrosoricida  

 Chrysochloridae (golden mole)


   

 Potamogalidae (Otter shrews)


   

 Tenrecidae (Tenreks)







Template: Klade / Maintenance / Style

The golden moles are a family within the order of Tenrekartigen (Afrosoricida), its closest relatives are the tenrecs (Tenrecidae) and potamogalinae (Potamogalidae). The Tenrekartigen in turn form part of the superiority of the Afrotheria , one of the four main lines within the higher mammals . The Afrotheria are a kinship group originating predominantly from Africa or native there, whose togetherness is based primarily on molecular genetic studies and less on anatomical similarities. Two larger lines can be distinguished within the Afrotheria, the Paenungulata and the Afroinsectiphilia . The former include today's elephants , hyrax and manatees , the group has long been considered a common unit of descent. In addition to the tenre karts, the elephants and sometimes the aardvark are also placed on the second line . Some genetic studies support a sister group relationship between the Afrosoricida and the elephants, whereby both are then united under the taxon of the Afroinsectivora. Others, in turn, see the tenre cardigans in a sister group position to all other Afrotheria, while the elephants form a closer relationship with the aardvark. According to molecular genetic studies, the Afrotheria originated in the Upper Cretaceous between 90.4 and 80.9 million years ago. The split between the two main groups took place around 15 million years later. The Afrosoricida then appeared shortly before the Cretaceous-Tertiary boundary about 68 million years ago. In contrast, the gold mole only diversified in the course of the Oligocene around 28 million years ago. However, the oldest fossil record of gold molluscs to date dates back to the Eocene and is estimated to be between 48 and 41 million years old.

Internal system

Internal systematics of the gold mole according to Asher et al. 2010
 Chrysochloridae  




 Eremitalpa


   

 Huetia


   

 Cryptochloris


   

 Chrysochloris





   

 Chrysospalax 



   

 Calcochloris



   

 Chlorotalpa


   


 Carpitalpa


   

 Neamblysomus



   

 Amblysomus



Template: Klade / Maintenance / 3

Template: Klade / Maintenance / Style

The internal systematic structure of the gold mole was evaluated differently over the decades, which was based on a different weighting of characteristics. After several genera had already been described in the 19th century, Robert Broom listed them all within Chrysochloris at the beginning of the 20th century , but subdivided the genus based on the presence or absence of an externally visible bone vesicle on the temporal fossa of the skull and the number of teeth ( 36 or 40) in many ways. In the 1920s, Austin Roberts subdivided the gold mulle at the generic level, introduced several new genera and established some of the previously described new ones, which became established in the period that followed. However, in later decades there was frequent amalgamation and separation of individual genres, which in turn resulted from the different emphasis on individual characteristics. In most cases these attempts at classification were not based on statistical surveys. This only changed in the mid-1990s with the investigations of Gary N. Bronner , who carried out intensive morphometric and cytogenetic analyzes. The current systematic classification of the gold mole on species and genus level is largely based on Bronner.

The exact relationships between the individual genera of the golden mole remained unclear for a long time and were only examined in individual cases. In 1957 Alberto M. Simonetta then divided the family into three subfamilies. The training of the hammer on the middle ear served as the basis for this . Although numerous scientists in the 19th and early 20th centuries had pointed to the sometimes noticeably enlarged ossicles, it was Clive Forster Cooper who described them in more detail in 1928, but at that time he did not recognize their possible taxonomic value. Simonetta divides the Goldmulle as follows:

Morphology of the hammer and the anvil at Gold Mullen: Hottentot Goldmull ( Amblysomus hottentotus ; A), Sclater Goldmull ( chlorotalpa sclateri ; B), Arends' Goldmull ( Carpitalpa arendsi ; C), Cape Goldmull ( Chrysochloris asiatica ; D), Grant's golden mole ( Eremitalpa granti ; E), wire-haired golden mole rat ( Chrysospalax villosus ; F)
  • Amblysominae Simonetta , 1957: with a normally built malleus ( Amblysomus , Neamblysomus , Calcochloris , Huetia )
  • Chrysochlorinae Gray , 1825: with a greatly elongated head of the malleus ( Carpitalpa , Chlorotalpa , Chrysochloris , Cryptochloris )
  • Eremitalpinae Simonetta , 1957: with a spherically inflated head of the malleus ( Chrysospalax , Eremitalpa )

Simonetta assumed a gradual development of the head of the hammer from a relatively small one in the Amblysominae to a relatively large one in the Eremitalpinae. Other authors, however, later saw the spherically inflated or club-like elongated malleus head only as a variation of a feature that cannot be clearly separated even within the individual species and genera. They therefore summarized the Eremitalpinae with the Chrysochlorinae in a subfamily, the Chrysochlorinae.

At the beginning of the 21st century, the method of molecular genetics produced a picture that deviated somewhat from this traditional structure of the golden mole. As a result, two major main lines can be identified within the family, but they only partially coincide with the subfamilies. One group consists of the genera Amblysomus and Neamblysomus as well as Carpitalpa , the first two have a small hammer head, the latter a slightly enlarged one. They are informally referred to as the "amblysomine group". Opposite this is a "chrysochlorine group" with genera and species with an enlarged head of the malleus. The "chrysochlorine group" also includes, with Calcochloris and Huetia , shapes with a normal sized hammer. Chlorotalpa , whose hammer head is also enlarged , occupies a rather middle position between these two clades . It can therefore be assumed that the special characteristics in the area of ​​the middle ear do not go back to a single event within the tribal history of the golden mole, but rather represent a more homoplastic characteristic that arose several times within the family. In 2018, a new preliminary attempt at structuring the gold mole was presented. This includes the three previously established subfamilies. The Amblysominae would therefore be composed of the genera Amblysomus , Neamblysomus , Carpitalpa , Chlorotalpa , Chrysospalax and Calcochloris , the Chrysochlorinae in turn from the genus Chrysochloris (including Kilimatalpa and Cryptochloris ) and the Eremitalpinae from the genus Eremital . In addition, he provides with the Huetinae consisting of the genus Huetia a fourth independent subfamily.

Overview of the recent and fossil genera of the gold mole

The gold mole family is divided into ten genera living today, to which a total of 21 species currently belong:

  • Family: Chrysochloridae Gray , 1825
  • Amblysomus Pomel , 1848 (copper gold mole; five recent species)
  • Calcochloris Mivart , 1867 (yellow gold mole; a recent species)
  • Carpitalpa Lundholm , 1955 (Arends' Goldmull; a recent species)
  • Chlorotalpa Roberts , 1924 (two recent species)
  • Chrysochloris Lacépède , 1799 (Cape gold mulle; three recent species)
  • Chrysospalax Gill , 1883 (giant gold mole; two recent species)
  • Cryptochloris Shortridge & Carter , 1938 (two recent species)
  • Eremitalpa Roberts , 1924 (desert gold mole; a recent species)
  • Huetia Forcart , 1942 (two recent types)
  • Neamblysomus Roberts , 1924 (two recent species)

Studies from 2015 suggest that greater biodiversity is to be expected within the gold mole. This results from the strong adaptation to certain habitats with the formation of subpopulations, some of which exist in isolation from one another. The view is confirmed by further genetic analyzes from 2018. Their results support the recognition of further species, especially those of the genera Eremitalpa and Amblysomus . The total number of species of the gold mole could increase to at least around two dozen. In addition, the studies advocate an independent genus status for Kilimatalpa , which was previously managed within Chrysochloris , while Cryptochloris should only be understood as a subgenus of Chrysochloris .

The following types of fossil are recognized:

Another genus, Eochrysochloris Seiffert, Simons, Ryan, Bown & Attia , 2007, probably does not belong directly to the gold mole, but to another family field, the Chrysochloroidea .

Research history

Taxonomy

John Edward Gray
Carl von Linné

The first scientific naming of a representative of the gold mole was made by Linnaeus (1707–1778) in his Systema Naturae from 1758. He named today's Cape gold mole as Talpa asiatica , which he placed on the one hand to the Eurasian moles , on the other hand its distribution in Asia suspected. The information that Linnaeus had about the Cape Goldmull was taken from Albert Seba's work Thesaurus , published in 1734 . Bernard Germain Lacépède (1756-1825) introduced the generic name Chrysochloris for the Cape gold mole in 1799 and thus named a representative of the genus of the gold mole for the first time. The name Chrysochloris is borrowed from the Greek and consists of the words χρύσεος ( chryseos "golden") and χλωρός ( chlōrós "light green" or "fresh"). However, Chloris also refers to the Greek goddess of flowers, who finds her Roman counterpart in Flora . The generic name is not to be understood as a reference to the coat color, which can be very different, but refers to the metallic sheen of the coat under certain lighting conditions. From Chrysochloris also a surname Chrysochloridae derives. In general, this is today ascribed to John Edward Gray (1800–1875) and reference is made to a publication by him from 1825 in which Gray attempted to classify mammals into families and tribes . This used Gray although calling Chrysochlorina with which he the golden moles of the moles (talpina), the shrews (soricina), the hedgehogs (Erinacina), the tenrecs (Tenrecina) and tree shrews deposed (Tupaina) and all the family Talpidae united. The correct and correct spelling of the Chrysochloridae family goes back to St. George Jackson Mivart , who first used it in 1868. For a long time he was also considered the first name of the family.

On the position of the gold mole within the higher mammals

In 1758, Linnaeus brought the gold mole into connection with the Eurasian moles. He arranged them in the group of "Bestiae", in which he saw, among other things, the shrews and hedgehogs, but also the armadillos , the possums and pigs . Even after Lacépède discontinued the golden mole from the moles forty years later by establishing the genus Chrysochloris , the close grouping to other insect-eating animals remained. Johann Karl Wilhelm Illiger saw the golden mole as well as the moles, shrews and hedgehogs as well as the tenreks together in a group he called "Subterranea", which is the first time that he summarized all those taxa that were later to be called "insect eaters". Informally, Georges Cuvier used the term Les insectivores in 1817 with a similar composition to Illiger, the official name Insectivora comes from Thomas Edward Bowdich from 1821. Bowdich distinguished two groups within the Insectivora, one with long internal incisors like the gold mole and hedgehogs and a second with large canine teeth, such as the moles and tenreks.

The affiliation of the golden mole to the insectivores was only rarely questioned in the following century and a half; the only debates were largely the more precise relationships within the organizational unit. Wilhelm Peters worked out two nameless groups in 1863, one with an appendix , to which he referred the elephants and shrews , and one without, which included all other insectivora including the golden mole. Three years later , Ernst Haeckel gave these two groups the formal names Menotyphla (with appendix) and Lipotyphla (without appendix) in the second volume of his work General Morphology of Organisms . Theodore Gill pursued a different approach in the mid-1880s by distinguishing between the Zalambdodonta and the Dilambdodonta within the Insectivora. The division was based on the tooth design, with the zalambdodonta having a V-shaped enamel ridge on the molars (from Greek ζα ( za- ) for an excess and the Greek letter Λ (lambda)), the dilambdodonta a W-shaped (from Greek δι ( ie "two") and the Greek letter Λ (lambda)). The golden mole, together with the tenreks and the slit weevils, formed the zalambdodont insectivores, while the hedgehogs, moles and shrews (and also the elephants and shrews) belonged to the dilambdodonts. Gill recognized a closer relationship between the Goldmulle and the Tenreks. The views of Haeckel and Gill were partially adopted by William K. Gregory in his 1910 work The orders of Mammals . He led the actual Insectivora under the term Lipotyphla, but he positioned the Menotyphla outside of the insect eater. He kept the golden mole, tenreks and the slit weevils in the Zalambdodonta, but divided the remaining Dilambdodonta into the two large groups Erinaceomorpha (hedgehogs) and Soricomorpha (shrews and moles). In contrast to other authors, Robert Broom completely excluded the gold mulle from insectivores in 1915 and 1916 due to the design of the skull and also denied that it was closely related to the tenreks; he created a classification group for the gold mulle under the name Chrysochloridea. However, George Gaylord Simpson contradicted this in his general taxonomy of mammals published in 1945 and saw this step as too radical. He subdivided the then Insectivora into five recent superfamilies, one of which represented the gold mole under the name Chrysochloroidea, while he saw the tenreks as another superfamily (Tenrecoidea) and placed them in relative proximity to the gold mole. A closer relationship between the two groups within the insectivore was discussed again and again later, for example by Percy M. Butler in 1956, who took over the division of Gregory into the two large groups of the Erinaceomorpha and the Soricomorpha and referred the golden mole and tenreks to the Soricomorpha. For him, the golden mulle represented derived forms within the Soricomorpha due to the heavily embossed skull due to the underground way of life. Malcolm C. McKenna came to a similar result, but with a different conclusion, in 1975. He also assigned the golden mulle to the Soricomorpha, the but differed from these by a complete zygomatic arch. But since the cheekbone is missing, McKenna saw the closed arch in the gold mole as a new, secondary formation. In his Classification of Mammals, published in 1997 together with Susan K. Bell , McKenna excluded the golden mole from the Soricomorpha and established it in its own classification group within the insectivore.

It was not until the late 1990s that molecular genetic studies rearranged the relationships between the original insectivores (Insectivora, later Lipotyphla). Accordingly, the golden mole do not form a monophyletic unit with the moles or the shrews, but together with the tenreks form a group of insect-eating animals that is endemic in Africa , which was then referred to the separate order of the tenre-kitties (Afrosoricida; actually translated "African shrews"). In addition, a closer relationship between the Tenre-kitties and the elephants, the aardvark, the elephants, the snakes and the manatees and thus to mostly African native animals could be recognized, altogether an externally strongly heterogeneous group, which received the name Afrotheria . Subsequent analyzes underpinned this result, among other things through the isolation of a specific retroposon , the so-called Afro SINE , which all representatives of Afrotheria have in common. For this reason, the similarities of the golden mole with moles and other insectivores are based solely on convergence .

Tribal history

The fossil history of the gold mole is sparse. All extinct representatives are already largely similar in their physique to today's species. They have a teardrop-shaped skull with ten teeth on each half of the jaw and, as far as the body skeleton has been handed down, show adaptations to a digging way of life. Differences can often only be seen in different tooth features and in the increasing reduction of the last molar.

Eocene

The earliest known finds date back to the Eocene and could be discovered in southern Africa. Some isolated molars, a lower jaw and possibly also individual upper jaw fragments come from the Black Crow find locality in the diamond restricted area of Namibia and are assigned to the genus Diamantochloris . With an age of about 48 to 41 million years, the finds belong to the Middle Eocene. The lower teeth already show the fusion of two of the three main cusps on the chewing surface (paraconid and metaconid), which is typical for Goldmulle. Since a particularly large talonid is also formed, other authors do not see Diamantochloris in the immediate predecessor line of today's gold mulle. The same site also contained a remains of the upper jaw and isolated lower teeth. They belong to an animal with comparable tooth characteristics, but which was significantly smaller than Diamantochloris . In 2019 the finds were described as belonging to the genus Damarachloris . Namachloris is a little younger , the evidence of which is based on more than 100 finds. The genus was discovered on the Eocliff , also in the restricted area. The formation of the formation is about 41 to 38 million years in the past, which corresponds to the Upper Eocene. According to the mammal fauna found, a somewhat younger age would also be possible. In addition to complete skulls and mandibles, the fossil material is composed of almost all parts of the body skeleton. For the first time, the “flexor” bones could be detected for a fossil gold mole, as well as the “triplex” and the “totem” bones are documented. The limbs show adaptations to a sand-digging way of life. The finds probably go back to prey remains of birds of prey or owls that hunted here on the bank of a former lake.

Oligocene

From the subsequent Oligocene , remains of gold mullets or their closest relatives have so far only come down to us from the Gebel-Qatrani formation of the Fayyum basin in northern Egypt . Two mandibular fragments belonging to the genus Eochrysochloris were found. The estimated age of 34 to 30 million years corresponds to the lower section of the geological series. The three main cusps of the molars are a noticeable feature, all of which are almost the same size and independent. It is possible that Eochrysochloris is not directly related to the modern Goldmullen, but belongs to a side branch.

Miocene

After that, gold mole can only be found again with Prochrysochloris in the course of the Miocene . The most extensive material to date came to light in western Kenya in the area of ​​Lake Victoria and consists of almost half a dozen skulls and some remains of the lower jaw. The Legetet Formation and the Kapurtay Agglomerates , both of which belong to the Lower Miocene and are between 20 and 17.5 million years old, are important deposits here . The latter includes, among other things, the important site of Songhor . As with some older fossil finds of the golden mole, the rear molar of Prochrysochloris was still largely unreduced in size, there was no prominent bony bulge at the temporal pit as in the Cape golden mole . The same genus could also be detected with a lower jaw fragment in Arrisdrift in southern Namibia. The find is a little younger than that from East Africa, the fossil-bearing strata date to the beginning of the Middle Miocene. Further mandibular fragments and individual teeth of Prochrysochloris are available from the same region from the more or less equally old sites of Elisabethfeld and Langental .

Pliocene and Pleistocene

In the Pliocene , forerunners of today's species appear for the first time. Two representatives of Chrysochloris have been documented from Langebaanweg in southwestern South Africa , which is at the beginning of this phase with an age of 5 million years . On the basis of the extensive fossil material, which consists of skull and lower jaw remains as well as parts of the body skeleton, conclusions can be drawn about sand-digging animals comparable to the desert gold mole . A few finds of Amblysomus , Neamblysomus , Chlorotalpa and Chrysospalax date from the transition to the Pleistocene around 2.5 million years ago . They were discovered in the important South African cave sites of Makapansgat and Sterkfontein . From near Sterkfontein, from Bolt's farm, comes a 29 mm long and 20 mm wide skull, which is reminiscent of that of the copper gold mulle due to its narrow construction. In contrast to these, however, it has a bulging, bony bladder on the temporal fossa, which does not occur in today's representatives of the genus. The bulge also shows lateral enlargements, which in turn is not documented with such structures in any other form of the gold mole. The skull was therefore referred to Proamblysomus .

Threat and protection

Several species are listed as endangered or threatened by the IUCN . The main causes of the threat are the loss of habitat through the expansion of human settlements in general, through conversion into fields or pasture areas, through mining or through infrastructure measures as a result of the tourist development of certain regions. Locally, the pursuit of domestic dogs and cats also have an impact on the populations . In addition, many species only inhabit a small area and are therefore particularly susceptible to disturbance. Twelve of the 21 species are "threatened with extinction" according to IUCN ( critically endangered ), "high risk" ( endangered , "at risk") ( vulnerable ) or "near threatened" ( near threatened ), for three more species lacking accurate data this however, they are also likely to be threatened. A larger part of the species is present in nature reserves, but in many cases there is no more detailed information on the biology of the animals.

literature

  • Gary N. Bronner: Family Chrysochloridae Golden-moles. In: Jonathan Kingdon, David Happold, Michael Hoffmann, Thomas Butynski, Meredith Happold and Jan Kalina (eds.): Mammals of Africa Volume I. Introductory Chapters and Afrotheria. Bloomsbury, London, 2013, pp. 223-257
  • Gary N. Bronner and Nigel C. Bennett: Order Afrosoricida. In: John D. Skinner and Christian T. Chimimba (Eds.): The Mammals of the Southern African Subregion. Cambridge University Press, 2005, pp. 1-21
  • Ronald M. Nowak: Walker's Mammals of the World . The Johns Hopkins University Press, Baltimore 1999, ISBN 0-8018-5789-9
  • Gerhard Storch: Lipotyphla, insect eater. In: Wilfried Westheide, Reinhard Rieger (Ed.): Special Zoology. Part 2: vertebrate or skull animals . Spectrum Academic Publishing House, Heidelberg - Berlin 2004, 712 pages, ISBN 3-8274-0307-3
  • William A. Taylor, Samantha Mynhardt and Sarita Maree: Chrysochloridae (Golden moles). In: Don E. Wilson and Russell A. Mittermeier (Eds.): Handbook of the Mammals of the World. Volume 8: Insectivores, Sloths and Colugos. Lynx Edicions, Barcelona 2018, pp. 180–203 ISBN 978-84-16728-08-4
  • Don E. Wilson and DeeAnn M. Reeder (Eds.): Mammal Species of the World . 3rd edition. The Johns Hopkins University Press, Baltimore 2005, ISBN 0-8018-8221-4

Individual evidence

  1. ^ Robert Broom: Some new and some rare golden moles. Annals of the Transvaal Museum 20, 1946, pp. 329-335
  2. ↑ a b c d e f g h i j k l m n o p Gary N. Bronner: Family Chrysochloridae Golden-moles. In: Jonathan Kingdon, David Happold, Michael Hoffmann, Thomas Butynski, Meredith Happold and Jan Kalina (eds.): Mammals of Africa Volume I. Introductory Chapters and Afrotheria. Bloomsbury, London, 2013, pp. 223-257
  3. ↑ a b c d e f g h i j k l m n o p q Gary N. Bronner and Nigel C. Bennett: Order Afrosoricida. In: John D. Skinner and Christian T. Chimimba (Eds.): The Mammals of the Southern African Subregion. Cambridge University Press, 2005, pp. 1-21
  4. ↑ a b c d e f g h i j k William A. Taylor, Samantha Mynhardt and Sarita Maree: Chrysochloridae (Golden moles). In: Don E. Wilson and Russell A. Mittermeier (Eds.): Handbook of the Mammals of the World. Volume 8: Insectivores, Sloths and Colugos. Lynx Edicions, Barcelona 2018, pp. 180–203 ISBN 978-84-16728-08-4
  5. ^ A b Robert Broom: On the structure of the skull in Chrysochloris. Proceedings of the Zoological Society of London 1916, pp. 449-459
  6. ^ Gary N. Bronner, Elizabeth Jones and DJ Coetzer: Hyoid-dentary articulations in golden moles (Mammalia: Insectivore; Chrysochloridae). Zeitschrift für Mammaliankunde 55, 1990, pp. 11-15
  7. ^ Matthew J. Mason: Morphology of the middle ear of golden moles (Chrysochloridae). Journal of Zoology 260, 2003, pp. 391-403
  8. ^ Matthew J. Mason: Functional Morphology of the Middle Ear in Chlorotalpa Golden Moles (Mammalia, Chrysochloridae): Predictions From Three Models. Journal of Morphology 261, 2004, pp. 162-174
  9. ^ A b Matthew J. Mason, Sarah J. Lucas, Erica R. Wise, Robin S. Stein and Melinda J. Duer: Ossicular density in golden moles (Chrysochloridae). Journal of Comparative Physiology A 192, 2006, pp. 1349-1357
  10. ↑ Joseph Hyrtl: Comparative anatomical studies on the internal auditory organ. Prague, 1845, pp. 1–139 (pp. 10–14 and 59–60) ( [1] )
  11. ^ A b c Robert Broom: A contribution to the knowledge of the cape golden moles. Transactions of the South African Philosophical Society 18, 1907, pp. 283-311 ( [2] )
  12. ↑ Nick Crumpton, Nikolay Kardjilov and Robert J. Asher: Convergence vs. Specialization in the Ear Region of Moles (Mammalia). Journal of Morphology 276, 2015, pp. 900-914
  13. ↑ Matthew J. Mason and Peter M. Narins: Seismic sensitivity in the Desert golden mole (Eremitalpa granti): A review. Journal of Comparative Psychology 116 (2), 2002, pp. 258-263
  14. ↑ Matthew J. Mason: Bone conduction and seismic sensitivity in golden moles (Chrysochloridae). Journal of Zoology 260, 2003, pp. 405-413
  15. ↑ UB Willi, GN Bronner and PM Narins: Ossicular differentiation of airborne and seismic stimuli in the Cape golden mole (Chrysochloris asiatica). Journal of Comparative Physiology A 192, 2006, pp. 267-277
  16. ^ Matthew J. Mason: Of mice, moles and guinea pigs: Functional morphology of the middle ear in living mammals. Hearing Research 301, 2013, pp. 4-18.
  17. ^ Robert Broom: Some observations on the dentition of Chrysochloris and on the Tritubercular Theory. Annals of the Natal Museum 2, 1916, pp. 129-139 ( [3] )
  18. ^ Robert J. Asher and Marcelo R. Sánchez-Villagra: Locking Yourself Out: Diversity Among Dentally Zalambdodont Therian Mammals. Journal of Mammalian Evolution. 12 (1/2), 2005, pp. 265-282
  19. ↑ a b c d J. P. Gasc, FK Jouffroy, S. Renous and F. von Blottnitz: Morphofunctional study of the digging system of the Namib Desert Golden mole (Eremitalpa granti namibensis): cinefluorographical and anatomical analysis. Journal of Zoology 208, 1986, pp. 9-35
  20. ^ A b c Robert J. Asher and D. Margaret Avery: New Golden Moles (Afrotheria, Chrysochloridae) from the Early Pliocene of South Africa. Palaeontologia Electronica 13 (1), 2010, p. 3A ( [4] )
  21. ↑ a b Märtha Kindahl: The embryonic development of the hand and foot of Eremitalpa (Chrysochloris) granti (Broom). Acta Zoologica 30, 1949, pp. 133-152
  22. ^ A b Gillian M Puttick and Jennifer UM Jarvis: The Functional Anatomy of the Neck and Forelimbs of the Cape Golden Mole, Chrysochloris Asiatica (Lipotyphla: Chrysochloridae). Zoologica Africana 12, 1977, pp. 2, 445-458
  23. ↑ a b c d Martin Pickford: Late Eocene Chrysochloridae (Mammalia) from the Sperrgebiet, Namibia. Communications of the Geological Survey of Namibia 16, 2015, pp. 153–193
  24. ↑ Michael F. Schneider and Victorino a Buramuge: Atlas of the microscopic hair structure of Southern African shrews, hedgehogs, goldem moles and elephant-shrews (Mammalia). Bonn zoological contributions 54 (3), 2006, pp. 103–172
  25. ↑ Holly K. Snyder, Rafael Maia, Liliana D'Alba, Allison J. Shultz, Karen MC Rowe, Kevin C. Rowe and Matthew D. Shawkey: Iridescent color production in hairs of blind golden moles (Chrysochloridae). Biology Letters 8, 2012, pp. 393-396
  26. ↑ Julia Boonzaier, Elizabeth L. Van der Merwe, Nigel C. Bennett and Sanet H. Kotzé: Comparative Gastrointestinal Morphology of Three Small Mammalian Insectivores: Acomys spinosissimus (Rodentia), Crocidura cyanea (Eulipotyphla), and Amblysomus hottentotus) (Afrosoricidaida). Journal of Morphology 274 (6), 2013, pp. 615-626
  27. ↑ Cassandra Pinheiro, Michaela L. Venter, Lauren Sahd, Gary N. Bronner and Sanet H. Kotzé: Comparative gastrointestinal morphology of seven golden mole species (Mammalia: Chrysochloridae) from South Africa. Journal of Morphology 279, 2018, pp. 1776-1786
  28. ^ Matthew RE Symonds: Phylogeny and life histories of the 'Insectivora': controversies and consequences. Biological Reviews 80, 2005, pp. 93-128
  29. ↑ a b Tarryn A. Retief, Nigel C. Bennett, Anouska A. Kinahan and Philip W. Bateman: Sexual selection and genital allometry in the Hottentot golden mole (Amblysomus hottentotus). Mammalian Biology 78, 2013, pp. 356-360
  30. ↑ J. Michael Bedford, Orin B. Mock and Steven M. Goodman: Novelties of conception in insectivorous mammals (Lipotyphla), particularly shrews. Biological Reviews 79, 2004, pp. 891-909
  31. ↑ a b Laura J. Fielden, Michael R. Perrin and GC Hickman: Water metabolism in the Namib desert golden mole, Eremitalpa granti namibiensis (Chrysochloridae. Comparative Biochemistry and Physiology 96A (1), 1990, pp. 227-234
  32. ↑ Jason A. Kaufman, Gregory H. Turner, Patricia A. Holroyd, Francesco Rovero and Ari Grossman: Brain Volume of the Newly-Discovered Species Rhynchocyon udzungwensis (Mammalia: Afrotheria: Macroscelidea): Implications for Encephalization in Sengis. PlosOne 8 (3), 2013, p. E58667, doi: 10.1371 / journal.pone.0058667
  33. ^ Georgina Sweet: The Eyes of Chrysochloris hottentota and C. asiatica. Journal of Cell Science 53 (2), 1909, pp. 327-338
  34. ↑ V. Gubbay: A comparison of the development of the rudimentary eye of Eremitalpa granti (Broom) with did of the normal eye of Elephantulus myurus jamesoni (Chubb). South African Journal of Science 52 (8), 1956, pp. 182-186 and 193-198
  35. ↑ Howard P. Whidden: Extrinsic Snout Musculature in Afrotheria and Lipotyphla. Journal of Mammalian Evolution 9 (1/2), 2002, pp. 161-184
  36. ↑ a b R. S. Seymour, PC Withers and WW Weathers: Energetics of burrowing, running, and free-living in the Namib Desert golden mole (Eremitalpa namibensis). Journal of Zoology 244, 1998, pp. 107-117
  37. ↑ a b c d e M. A. Kuyper: The ecology of the golden mole Amblysomus hottentotus. Mammal Review 15 (1), 1985, pp. 3-11
  38. ↑ Laura J. Fielden, JP Waggoner, Michael R. Perrin and GC Hickman: Thermoregulation in the Namib Desert golden mole, Eremitalpa granti namibensis (Chrysochloridae). Journal of Arid Environments 18, 1990, pp. 221-237
  39. ^ NC Bennett and AC Spinks: Thermoregulation and metabolism in the Cape golden mole (Insectivora: Chrysochloris asiatica). Journal of Zoology 76, 1995, pp. 957-971
  40. ↑ Laura J. Fielden, GC Hickman and Michael R. Perrin: Locomotory activity in the Namib Desert golden mole Eremitalpa granti namibensis (Chrysochloridae). Journal of Zoology 226, 1992, pp. 329-344
  41. ↑ Laura J. Fielden, Michael R. Perrin and GC Hickman: Feeding ecology and foraging behavior of the Namib Desert golden mole, Eremitalpa granti namibensis (Chrysochloridae). Journal of Zoology 220, 1990, pp. 367-389
  42. ↑ Jennifer UM Jarvis: Notes on the golden mole, Chrysochloris stuhlmanni Matschie, from the Ruwenzori Mountains, Uganda. East African Wildlife Journal 12, 1974, pp. 163-166
  43. ^ Matthew J. Mason and Peter M. Narins: Seismic Signal Use by Fossorial Mammals. American Zoologist 41 (5), 2001, pp. 1171-1184
  44. ^ Gary N. Bronner: Notes on the early post-natal development of a giant golden mole (Günther, 1875) born in captivity (Mammalia: Insectivora; Chrysochloridae). Koedoe 35 (2), 1992, pp. 57-58
  45. ^ A b Matjaž Kuntner, Laura J. May-Collado and Ingi Agnarsson: Phylogeny and conservation priorities of afrotherian mammals (Afrotheria, Mammalia). Zoologica Scripta 40 (1), 2011, pp. 1-15
  46. ↑ a b Peter J. Waddell, Hirohisa Kishino and Rissa Ota: A Phylogenetic Foundation for Comparative Mammalian Genomics. Genome Informatics 12, 2001, pp. 141-154
  47. ^ A b Robert W. Meredith, Jan E. Janečka, John Gatesy, Oliver A. Ryder, Colleen A. Fisher, Emma C. Teeling, Alisha Goodbla, Eduardo Eizirik, Taiz LL Simão, Tanja Stadler, Daniel L. Rabosky, Rodney L. Honeycutt, John J. Flynn, Colleen M. Ingram, Cynthia Steiner, Tiffani L. Williams, Terence J. Robinson, Angela Burk-Herrick, Michael Westerman, Nadia A. Ayoub, Mark S. Springer and William J. Murphy: Impacts of the Cretaceous Terrestrial Revolution and KPg Extinction on Mammal Diversification. Science 334, 2011, pp. 521-524
  48. ↑ Erik R Seiffert: A new estimate of afrotherian phylogeny based on simultaneous analysis of genomic, morphological, and fossil evidence. BMC Evolutionary Biology 7, 2007, p. 224, doi: 10.1186 / 1471-2148-7-224
  49. ↑ C. Gilbert, PC O'Brien, G. Bronner, F. Yang, A. Hassanin, MA Ferguson-Smith and TJ Robinson: Chromosome painting and molecular dating indicate a low rate of chromosomal evolution in golden moles (Mammalia, Chrysochloridae) . Chromosome Research 14, 2006, pp. 793-803
  50. ↑ a b c d e Martin Pickford: Chrysochloridae (Mammalia) from the Lutetian (Middle Eocene) of Black Crow, Namibia. Communications of the Geological Survey of Namibia 16, 2015, pp. 105–113
  51. ↑ a b c Robert J. Asher, Sarita Maree, Gary Bronner, Nigel C Bennett, Paulette Bloomer, Paul Czechowski, Matthias Meyer and Michael Hofreiter: A phylogenetic estimate for golden moles (Mammalia, Afrotheria, Chrysochloridae). BMC Evolutionary Biology 10, 2010, p. 69, doi: 10.1186 / 1471-2148-10-69
  52. ^ Austin Roberts: Some additions to the list of South African mammals. Annals of the Transvaal Museum 10 (2), 1924, pp. 59-76
  53. ^ A b c Alberto M. Simonetta: A new golden mole from Somalia with an appendix on the taxonomy of the family Chrysochloridae (Mammalia, Insectivora). Monitore Zoologico Italiano NS Supplement 2, 1968, pp. 27-55
  54. ^ F. Petter: Remarques sur la systematique des Chrysochlorides. Mammalia 45 (1), 1981, pp. 49-53
  55. ^ Gary N. Bronner: Geographic patterns of morphometric variation in the Hottentot golden mole, Amblysomus hottentotus (Insectivora: Chrysochloridae). A multivariate analysis. Mammalia 60 (4), 1996, pp. 729-751
  56. ^ Gary N. Bronner: Non-eographic variation in morphological characteristics of the Hottentot golden mole, Amblysomus hottentotus (Insectivora: Chrysochloridae). Mammalia 60 (4), 1996, pp. 707-727
  57. ^ Gary N. Bronner: Cytogenetic Properties of Nine Species of Golden Moles (Insectivora: Chrysochloridae). Journal of Mammalogy 76 (3), 1995, pp. 957-971
  58. ^ A b Gary N. Bronner: Systematic revision of the Golden mole genera Amblysomus, Chlorotalpa and Calcochloris (Insectivora, Chrysochloromorpha, Chrysochloridae). University of Natal, Pretoria, 1995, pp. 1–346 (p. 268)
  59. ↑ Clive Forster Cooper: On the ear region of certain of the Chrysochloridae. Philosophical Transactions of the Royal Society of London. Series B 435, 1928, pp. 265-282
  60. ^ Acacia von Mayer, G. O'Brian and EE Sarmiento: Functional and systematic implications of the ear in golden moles (Chrysochloridae). Journal of Zoology 236m 1995, pp. 417-430
  61. ^ A b Gary Bronner: An imminent updated (2017) taxonomy for golden moles. Afrotherian Conservation 14, 2018, pp. 57–59
  62. ↑ Samantha Mynhardt, Sarita Maree, Illona Pelser, Nigel C. Bennett, Gary N. Bronner, John W. Wilson and Paulette Bloomer: Phylogeography of a Morphologically Cryptic Golden Mole Assemblage from South-Eastern Africa. PlosONE 10 (12), 2015, p. E0144995, doi: 10.1371 / journal.pone.0144995
  63. ^ A b Percy M. Butler and A. Tindell Hopwood: Insectivora and Chiroptera from the Miocene rocks of Kenya Colony. Fossil Mammals of Africa 13, 1957, pp. 1–35 ( [5] )
  64. ^ A b Robert J. Asher: Tenrecoidea. In: Lars Werdelin and William Joseph Sanders (eds.): Cenozoic Mammals of Africa. University of California Press, Berkeley, London, New York, 2010, pp. 99-106
  65. ^ A b Martin Pickford: New Chrysochloridae (Mammalia) from the middle Eocene of Black Crow, Namibia. Communications of the Geological Survey of Namibia 21, 2019, pp. 40–47
  66. ↑ a b Erik R. Seiffert, Elwyn L. Simons, Timothy M. Ryan, Thomas M. Bown and Yousry Attia: New remains of Eocene and Oligocene Afrosoricida (Afrotheria) from Egypt, with implications for the origin (s) of afrosoricid zalambdodonty . Journal of Vertebrate Paleontology 27 (4), 2007, pp. 963-972
  67. ^ A b Carl von Linné: Systema naturae. 10th edition, 1758, volume 1, p. 53 ( [6] )
  68. ↑ Oldfield Thomas: On the probable identity of certain specimens, formerly in the Lidth de Jeude Collection, and now in the British Museum, with those figured by Albert Seba in his 'Thesaurus' of 1734. Proceedings of the Zoological Society 1892, p. 309-318 ( [7] )
  69. ↑ John Edward Gray: An outline of an attempt at the disposition of Mammalia into tribes and families, with a list of the genera apparently appertaining to each tribe. Annals of Philosophy NS 10, 1825, pp. 337–344 ( [8] )
  70. ↑ St. George Jackson Mivart: Notes on the Osteology of the Insectivora. Journal of Anatomy and Physiology 2 (1), 1868, pp. 117–154 (p. 150) ( [9] )
  71. ^ A b George Gaylord Simpson: The principles of classification and a classification of mammals. Bulletin of the American Museum of Natural History 85, 1945, pp. 1–350 (pp. 49 and 175–179)
  72. ^ Johann Karl Wilhelm Illiger: Prodromus Systematis Mammalium et Avium. Berlin, 1811, pp. 1–301 (p. 126) ( [10] )
  73. ↑ Georges Cuvier: Le regne animal distribué d'apres son organization pour servir de base a l'histoire naturelle des animaux et d'introduction à l'anatomie comparée. Paris, 1817, pp. 1–540 (pp. 131–138) ( [11] )
  74. ^ Thomas Edward Bowdich: An analysis of the natural classifications of Mammalia for the use of students and travelers. Paris, 1821, pp. 1–115 (pp. 31–33) ( [12] )
  75. ↑ Wilhelm Peters: About the mammalian species Selenodon. Treatises of the Royal Academy of Sciences in Berlin 1863, pp. 1–22 ( [13] )
  76. ↑ Ernst Haeckel: General Morphology of Organisms. Volume 2. Berlin, 1866, pp. 1–462 (p. CLX) ( [14] )
  77. ^ Theodore Gill: Insectivora. In: John Sterling Kingsley (ed.): Standard natural history. Volume 5. Boston, 1885, pp. 1–535 (pp. 134–158) ( [15] )
  78. ^ William King Gregory: The orders of mammals. Bulletin of the American Museum of Natural History 27, 1910, pp. 1–524 (pp. 464–465)
  79. ^ Robert Broom: On the Organ of Jacobson and its relations in the "Insectivora". Part II. Talpa, Centetes and Chrysochloris. Proceedings of the Zoological Society of London 1915, pp. 347-354 ( [16] )
  80. ^ Percy M. Butler: The skull of Ictops and the classification of the Insectivora. Proceedings of the Zoological Society of London 126, 1956, pp. 453-481
  81. ↑ Malcolm C. McKenna: Toward a phylogenetic classification of the Mammalia. In W. Patrick Luckett and Frederick S. Szalay (Eds.): Phylogeny of the primates: a multidisciplinary approach. New York, London, 1975, pp. 21-46
  82. ↑ Malcolm C. McKenna and Susan K. Bell: Classification of mammals above the species level. Columbia University Press, New York, 1997, pp. 1-631 (pp. 272-295)
  83. ↑ Mark S. Springer, Gregory C. Cleven, Ole Madsen, Wilfried W. de Jong, Victor G. Waddell, Heather M. Amrine, and Michael J. Stanhope: Endemic African mammals shake the phylogenetic tree. Nature 388, 1997, pp. 61-64
  84. ↑ a b Michael J. Stanhope, Victor G. Waddell, Ole Madsen, Wilfried de Jong, S. Blair Hedges, Gregory C. Cleven, Diana Kao and Mark S. Springer: Molecular evidence for multiple origins of Insectivora and for a new order of endemic African insectivore mammals. PNAS 95, 1998, pp. 9967-9972 ( [17] )
  85. ↑ Masato Nikaido, Hidenori Nishihara, Yukio Hukumoto and Norihiro Okada: Ancient SINEs from African Endemic Mammals. Molecular Biology and Evolution 20 (4), 2003, pp. 522-527
  86. ↑ Hidenori Nishihara, Yoko Satta, Masato Nikaido, JGM Thewissen, Michael J. Stanhope and Norihiro Okada: A Retroposon Analysis of Afrotherian Phylogeny. Molecular Biology and Evolution 22 (9), 2005, pp. 1823-1833
  87. ^ Martin Pickford: Diamantochloris mandible from the Ypresian / Lutetian of Namibia. Communications of the Geological Survey of Namibia 19, 2018, pp. 51–65
  88. ^ A b Robert J. Asher: Recent additions to the fossil record of tenrecs and golden moles. Afrotherian Conservation 15, 2019, pp. 4–13
  89. ^ Percy M. Butler: Macroscelidea, Insectivora and Chiroptera from the Miocene of East Africa. Palaeovertebrata 14 (3), 1984, pp. 117-200
  90. ↑ Pierre Mein and Martin Pickford: Insectivora from Arrisdrift, a basal Middle Miocene locality in southern Namibia. Memoir of the Geological Survey of Namibia 19, 2003, pp. 143-146
  91. ↑ Pierre Mein and Martin Pickford: Early Miocene Insectivores from the Northern Sperrgebiet, Namibia. Memoir of the Geological Survey of Namibia 20, 2008, pp. 169-183
  92. ^ G. de Graaff: A new chrysochlorid from Makapansgat. Palaeontologia Africana 5, 1958, pp. 21-27
  93. ^ A b Robert Broom: On two Pleistocene Golden moles. Annals of the Transvaal Museum 20, 1941, pp. 215-216
  94. ^ Diana M. Avery: An assessment of the Lower Pleistocene micromammalian fauna from Swartkrans Members 1-3, Gauteng, South Africa. Geobios 31 (3), 1998, pp. 393-414
  95. ^ Diana M. Avery: Notes on the systematics of micromammals from Sterkfontein, Gauteng, South Africa. Palaeontologia Africana 36, ​​2000, pp. 83-90
  96. ^ Diana M. Avery: The Plio-Pleistocene vegetation and climate of Sterkfontein and Swartkrans, South Africa, based on micromammals. Journal of Human Evolution 41, 2001, pp. 113-132
  97. ^ IUCN: The IUCN Red List of Threatened Species. Version 2015. ( [18] ), accessed June 24, 2016

Web links

Commons : Goldmulle  - Collection of images, videos and audio files