Malaysian Black Scorpion – Detailed Guide: Care, Diet, and Breeding

Malaysian Black Scorpion (Heterometrus spinifer) – profile Care, Diet, and Breeding
Heterometrus spinifer (the Malaysian Black scorpion), is one of the renowned scorpion species in the hobby, owing to their impressive size, hardiness, and adaptability to a variety of conditions.

This is fantastic hardy beginner-friendly species for anyone new to keeping scorpions.

In this guide, I gathered everything we currently know about Heterometrus spinifer including ideal tank setups, habits, healthy diets, breeding, compatibility, etc.

Important: Unfortunately, Heterometrus spinifer is not widely available in the exotic pet trade anymore. Because of overharvesting, the species has been protected from commercial exploitation since 2005 by the Malaysian government.

All scorpions of this species are only available as captive-born.

Another problem is that Heterometrus species are often sold under the wrong names. Some of these species can be very hard to identify until the adult stage. For example, most Heterometrus spinifer on the market, in reality, are Heterometrus silenus (ex. Heterometrus petersii).

Quick Notes about Malaysian Black Scorpion

Name Malaysian Black Scorpion
Other Names
The Thai black scorpion, the Giant forest scorpion, or the Giant black scorpion
Scientific Name Heterometrus spinifer 
Type
Terrestrial
Tank size (minimum) 10 gallons (~40 liters)
Keeping Easy
Breeding Easy  
Average size 4 – 5.3 inches (10 – 14 cm)
Optimal Temperature 75 – 82°F (24 – 28 °C)
Water type Freshwater bowl
Moisture 70 – 90 %
Substrate 3 – 6 inches (7 – 15 cm) deep
Diet Carnivore  
Temperament Semi-aggressive 
Life span up to 10 years
Color Form Brown to black with reddish-brown telson 

Interesting fact: Scorpions are considered to be living fossils as they have existed on the earth since the Silurian period, around 430 million years ago. During their long-term evolution, scorpions kept their morphology almost unchanged.

Etymology of Malaysian Black Scorpion

The word ‘Heterometrus’ came from the Greek word ‘Héteros’ meaning ‘Different or other’ and the Greek word ‘Metrus’ = ‘Measurement’. 

  • Heterometrus spinifer. Spinifer is from ‘Spīna’, meaning ‘Thorn, spine, prickle’ + ‘-fer’ (‘Carrying or bearing’).

Distribution of Malaysian Black Scorpion

The distribution of Heterometrus spinifer is still a matter of discussion and debate even in scholarly literature.

It was believed that this species was distributed in southeast Asia, including Malaysia, Thailand, Cambodia, and Vietnam. The type locality ‘India’ was regarded as erroneous even then.

Nowadays, scientists presume that Heterometrus spinifer can be found only throughout peninsular Malaysia (recorded from the Federal Territory of Kuala Lumpur and the states of Johor, Kedah, Kelantan, Melaka, Negeri Sembilan, Pahang, Perak, Perlis, Selangor, and Terengganu) and also recorded from Tioman Island.

Natural Habitat of Malaysian Black Scorpion

Heterometrus spinifer is a terrestrial species. They live in the tropical rainforests in the dark undergrowth under logs, trees, rocks, leaf litter, or other debris.

They are exceptionally biodiverse animals and can easily utilize and adapt to various microhabitats to survive.

Description of Malaysian Black Scorpion

Malaysian Black Scorpion (Heterometrus spinifer) – Detailed Guide Care, Diet, and Breeding
photo by Petr Hamerník, Zoo Praha

Heterometrus spinifer is a relatively large scorpion. On average, fully grown adults can reach 4 – 5.3 inches (or 10 –14 cm) in length. 

These scorpions have a bulky cephalothorax, a broad-segmented abdomen, and a long thin delicate tail-like structure called a telson.

Like most scorpion species, Heterometrus spinifer has very bad eyesight.

To find food and navigate through the environment, they rely on their vibration-sensing organs – little tiny hairs on their body, pincers, pectines, and legs.

The coloration is uniformly black, only the manus and telson may be reddish brown.

Related article:

Difference Between Heterometrus spinifer and Heterometrus silenus

It is very easy to confuse these species because they are quite similar in color and appearance, especially, when they are young (up to instar 5).

Nonetheless, it is extremely important not to confuse them!

First of all, these species should not be mixed for safety reason. If you cannot figure out what species you have, your scorpions do not have this problem.

They can tolerate the company of their species members. However, they do not like intruders of other scorpion species and can fight with them. Basically, this is a good example of a species competition in nature.

Second, it is the price. Heterometrus spinifer is priced higher. Most people want them.

Similarities:

  • totally black/dark brown carapace,
  • the granulation pattern on the carapace (under the median eyes),
  • little granulation on the pedipalps.

Differences:

  • Chelae. Heterometrus spinifer has slightly narrower chelae (length to width ratio 2.4–2.6 ) compared to Heterometrus silenus (length to width ratio 2–2.4).
  • Telson. Heterometrus spinifer has reddish-brown telson as adults (pale yellow/orange as juveniles). Heterometrus Silenus has usually dark brown to completely black telson.
Some sources say that Heterometrus spinifer may also have a dark brown or dark telson. Many sellers (intentionally or not) may tell you that as well.

The problem though is that even if it is true, such individuals are so rare (I would say – unique!) that should not be taken into account. Keep that in mind, next time to see something like that.

Related article:

Lifespan of Malaysian Black Scorpion

Currently, there is no available data on the longevity of wild scorpions.

In captivity, Heterometrus spinifer can live up to 10 years under optimal conditions and good care.

Note: Lifespan is likely shorter in the wild because of diseases, predation, and poor living conditions.

Typical Behavior of Malaysian Black Scorpion

Malaysian Black Scorpion (Heterometrus spinifer) – profile Care, Diet, and Breeding
photo by Виктор Гриц

Heterometrus spinifer is nocturnal specie. The pick of their activity starts at dusk and stops during the night, before sunrise.

In addition, they are very skittish and would prefer to go and hide under logs, stones, leaves, or whatever’s inside the enclosure.

They are not active scorpions. So, do not expect these scorpions to stay in the open and/or crawl around their habitat for a long time. In some cases, they can hide for weeks. Do not worry too much. This is normal, especially, when they are just introduced into a new enclosure. They also become a bit more lethargic in the pre-molt state.

Heterometrus spinifer is a burrowing species. They enjoy doing it.

Sometimes they may even block the entrance for hibernation when conditions are not good enough. This is called diapause – a period of suspended development.

Fully grown Malaysian black scorpions are one of the few species that can relatively safely cohabit with minimal aggression or cannibalism of their species members. While cannibalism does occur, it is mostly between young scorpions.

Although it is possible to keep this species communally it is still not considered a good idea. Even a small risk of aggression should be eliminated if you want to keep them healthy and happy.

Heterometrus spinifer is slightly more aggressive compared to other species in the genus Heterometrus. Nonetheless, they mostly react defensively. Thus, if not provoked, they can be relatively safely handled without any problem.

Females with their young (nymphs) become territorial and more aggressive, and instead of running away they grasp or may even try to sting.

Note: Scorpions respond by pulling the claws towards the mouth, erecting the tail, and scuttling backward (defensive withdrawal).

Features:

  • Communal: Yes (with caution, still not recommended)
  • Active: No
  • Peaceful: Yes (generally)
  • Burrowers: Yes
  • Venomous: Yes

Venom of Malaysian Black Scorpion

The venoms of the black scorpion Heterometrus spp. including the Malaysian black scorpion (Heterometrus spinifer) have been reported to be of a lower order of toxicity.

Heterometrus spinifer does not spray venom, these scorpion sting.

Although their venom peptides are powerful weapons to capture small prey and defend themselves against enemies, it generally should not cause too many problem (if any) for healthy adults. Something like a bee sting or even weaker.

Nonetheless, DO NOT allow children to touch Heterometrus spinifer.

Some people (especially children) may be allergic to their toxins and their venom can cause intense localized pain: redness, swelling, and itching can occur as well. In this case, if you get stung, you need to get to the hospital ASAP.

Interesting fact: Scorpions can sting many times, but their venom becomes depleted with each sting.

Diet of Malaysian Black Scorpion

In the wild, Heterometrus spinifer is considered to be generalist predator on a variety of prey, such as insects and other small animals they can capture and overpower, such as:

  • bugs,
  • crickets,
  • mealworms,
  • moths
  • snails,
  • spiders,
  • small vertebrates (like lizards, snakes, frogs, rodents, etc.).

They use sensory systems (long and very thin sensory hairs located all over the body) other than vision to locate prey.

Interesting facts:

  • Once these scorpions attain a certain size and become strong enough, they only use their claws to dispatch any normal prey item. Whereas young may try to sting prey as well.
    Note: This is, probably, because venom production may be costly from an energetics standpoint and large prey items are often rejected in favor of smaller items that do not require stinging.
  • Adult scorpions can easily survive without food for over several months.
Important: Large prey may harm the scorpion. In captivity, you should never give them prey that can fight back aggressively.

In the terrarium, the acceptable food items include:

  • crickets,
  • mealworms,
  • cockroaches,
  • red runner roaches,
  • worms.

How Often to Feed Malaysian Black Scorpion?

Scorpions have a very slow metabolism and do not move a lot. Therefore, feeding adults once a week is more than enough.

Females will require more food after getting their young.

Some Feeding Tips

  • Time. Heterometrus spinifer is nocturnal. Therefore, it is highly recommended to feed them at night (at least in the evening). By doing so, you will replicate the conditions and environment under which they eat naturally.
  • Prey size. It is absolutely important that prey be smaller than the scorpions to ensure that they can grab it easily with their claws. For example, adult scorpions can eat full-grown cricket (1 inch or 2.5 cm long).

         Ideally, their food should not be larger than their stomach.

  • Prey activity. These scorpions are not active hunters. They will not pursue their prey. Like most scorpion species, they prefer to wait for their moment. So, do not give them prey that will burrow, jump or fly around their enclosure. For example, remove the wings, chop the forelimbs, break the legs, or squash the head so that they wiggle and are easily accessible to the scorpions.
  • Refuses to eat. Do not leave the live food in the enclosure. If your scorpion does not want to eat you need to remove it the next day.
  • Scorplings. Young scorpions can be too weak to overpower their prey. However, they will consume the crickets that have just died.
  • Check the hiding spots. Keep in mind that scorpions often drag and store food in their hiding spots for later consumption. Check them from time to time to prevent any mold, mites, or bacterial contaminations. If it is not eaten in 2 days, remove it.

Feeding Problems with Malaysian Black Scorpion

Some hobbyists complain that their scorpions stopped eating for weeks. This often happens right after purchase. Some potential reasons include:

  • Shipment stress. Give them time to adapt to their new home.
  • Preparation for a molt. Changes in their eating habits change drastically before molting. You will see a sharp decrease in appetite. They may start eating less and eventually stop eating altogether.
  • Not hungry. Scorpions have a very low metabolism and can live without food for months.
  • Low temperature. When the temperature drops below the optimal range, Heterometrus spinifer becomes very lethargic.
  • Diet control. In some cases, we can have the opposite problem – a scorpion becomes so fat that it looks like its body squeezing out of its exoskeleton. Check its diet. You are giving them way too much.
  • Gravid female. Make sure that your scorpion is not a gravid female. In this case, you will also see a little bit of white space between each tergite segments and the pleural membrane will be exposed. This is normal.

Features:

  • Diet Type: Carnivore
  • Food Preference: Alive prey.
  • Feeding Frequency: 1 time a week (for adults) and 2 – 3 times a week (for babies)
Malaysian Black scorpions usually eat the prey alive. So, if you are a sensitive person, you should not keep them as pets.

Related article:

Keeping and Housing Malaysian Black Scorpion

The quality of the habitat is the main reason why scorpions have a reduced lifespan or die during molting.

Although these scorpions are very hardy, there are still some important factors that must be considered when caring for them.

It is crucial to understand that to keep them healthy and happy, we have to mimic their natural habitat.

Enclosure Type:

First of all, you will need to choose an enclosure that provides the appropriate levels of heat and humidity for Heterometrus spinifer. There are several options here.

  1. Aquarium
PROS CONS
It is made of ecological material (glass) There can be problems with the ventilation
Good for humidity  Heavy
Easy to buy  Transportation problems
  1. Terrarium
PROS CONS
It is made of ecological materials. Some models have problems with ventilation
Good for humidity. High cost and weight
Some models have good ventilation  
  1. Plastic container
PROS CONS
Cheap Plastic is not the most ecological material, especially in hot temperatures
Good for humidity Nor aesthetically pleasing
Ventilation can be done very easy*  
Easy to move  

*Note: Even if there are no holes in the plastic container, we can easily do those ourselves. Drill a few small holes on two opposite sides of the walls.

Cross ventilation between opposite walls is the ideal condition – closer to the base on one side and closer to the upper edge on the other side. It will improve natural airflow.

Important: Although Heterometrus spinifer prefers digging to climbing, any enclosure should be escape-proof anyway.

Related article:

Tank Size (Enclosure):

Even though Heterometrus spinifer can grow pretty big, on the internet you may find that some hobbyists keep their scorpions in tiny enclosures while others vote for large tanks.

In my opinion, the tank size depends on 2 main factors:

  • Size of the scorpion.
  • Density.

Personally, I would not recommend keeping young scorpions (up to 2 – 3 years old) together because of potential cannibalism, especially, during molting.

Young scorpions should have an enclosure that is at least 3-4 times larger than their size. It should be enough to create at least one hiding place for them.

Note: Do not keep them in little tiny boxes, as some people do. Sure, your scorpions will survive but will they be happy?

As for the adult Heterometrus spinifer, the minimum recommended tank size for one individual is a 10-gallon (40-liter) tank. A pair of adults will require at least 15 gallons (60 liters).

Having a larger tank is always preferable because it can be easier to make diverse areas for them to hide. Hiding is extremely important for them.

Tip: Long tanks are better than tall tanks of the same size for keeping scorpions. They need mostly a place to crawl from side to side, not up and down.

Light:

No special requirements.

Heterometrus spinifer are nocturnal creatures. Basically, using ambient light is enough.

UV lighting warning: Like other scorpions, they give off an amazing fluorescent bluish-green appearance under UV light (‘Blacklight’). Therefore, people often use this kind of lighting.

This is wrong.

UV lighting is not completely safe for scorpions. Ultraviolet is a part of the light from the sun, its rays can cause damage to the outer layers of their exoskeleton (eventually, causing molting problems). That is why they do not like to be exposed to the sun.

UV light can be used only for short periods of time or as an option to make the environment safe in the house while there is a scorpion living. It can help in locating the scorpion if it escapes its cage.

Related article:

Temperature:

The temperature inside the enclosure should always be between 75 – 82°F (24 – 28°C). Heterometrus spinifer does love warm temperatures. 

Keeping them below this range (down to 68°F (20°C)) is possible but it will reduce their metabolism and make them slower.

Note: Scorpions are cold-blooded animals. It means that they do not have control of their heat balance. Thus, their body temperature varies with the temperature of the environment.

Important: Heaters should never be placed under their enclosure because they can overheat your substrate and burn or kill burrowed scorpions there.

The best (safest) option will be to put the heater to the side of the enclosure, ideally,  some part of the heater should be above the substrate line to heat the air as well.

Note: It is important to also use a thermostat to help moderate the temperature and keep it in the desired range.

Humidity:

Although Heterometrus spinifer lives on land, these terrestrial scorpions require humid air to breathe, move and grow properly.

It is recommended to have a humidity level of more than 60%. The ideal range is considered to be between 70 and 90 %.

Keep one part of the enclosure a little bit more damp by gently watering the substrate. It will give your scorpion the choice of where it would like to be at this or that moment.

Tip: One of the easiest ways to keep the humidity up is to use lots of sphagnum moss inside their enclosure.
Note: Do not spray/mist scorpions directly, it freaks them out and they really hate it.

Related article:

Water:

Like most scorpion species, Heterometrus spinifer also needs freshwater. So, it is important to provide them with a shallow dish that they can easily crawl out of.

They do not need much, because these scorpions can hold on to the water for a very long time!

Some hobbyists may disagree with me by saying that these scorpions get most of their hydration from eating prey items. In addition, they do not need any dishes because even little water droplets on the side of the container will be enough for them.

It is true.

Nonetheless, having a water dish helps to maintain humidity. Also, it is always better to provide something extra for our pets.

Ideally, we need to give them something like bottled spring water. But if you decide to use tap water, let it age for 24 hours before using it. Tap water contains chlorine and it can be toxic to them. 

  • Do water changes every 5 – 7 days.

Substrate:

As I have already mentioned, Heterometrus spinifer are obligate burrowers. So, you must provide enough deep moist substrate to simulate a natural behavior.

Scorpions burrow to hide and rest. It stresses them a lot if they cannot do that.

The general rule of thumb is that the substrate should at least equal the length of the scorpion itself. If your setup allows you should always opt for deeper!

Young scorpions do not dig much whereas adults require more depth (5 – 6 inches or 12 – 15 cm deep minimum). 

Interesting fact: Asian forest scorpions are very sensitive to substrate vibrations.

The substrate should always be kept moist enough. It means that it should hold its shape when you squeeze it. However, not so wet that it drips or pools water.

Some popular choices of substrate include:

  • coco fiber (Eco earth),
  • peat moss,
  • organic topsoil,
  • Jungle mix soil,
  • Zoo Meds Creatures Creature soil,
  • Reptisoil,
  • The Bio Dude Terra.

It is also possible to use a combination of different substrates, for example:

  • coco fiber and sand and (5:1 ratio). It is easy to maintain and it holds moisture very well.
  • a mixture of peat moss and organic topsoil (1:1 ratio).

Do not use only sand! Sand often becomes supersaturated even though the top couple of inches are completely dry.

How often should we change the substrate?

Compared to many other animals, scorpions do not produce a lot of waste and when they do, it is also easy to remove.

Nonetheless, in my opinion, it is still better to change the substrate at least every 6 months.

Hiding places:

Ideally, you need to make sure that your Malaysian Black scorpion has lots of hiding places. They do enjoy hiding and burrowing, it makes them happy.

They need to be able to completely conceal themselves.

Use driftwood, rocks, coconuts, pieces of cork bark, etc. It will make them feel like they are still in the Asian rainforests.

Important: If you are planning to house several Malaysian Black scorpions, each of them needs a hiding place to feel secure in different parts of the enclosure!

Examples of Tank Equipment
(with links to check the price on Amazon)

Handling Malaysian Black Scorpion

Heterometrus spinifer is venomous species with powerful pair of claws, so you need to handle them with utmost care.

These scorpions are not pets you can play with!

Make sure children do not touch and/or provoke the scorpion. It is against ethics and provoking the scorpion has often its consequences.

Do not take them out just because you want to. You should really handle him as little as possible. They do not benefit from being handled and get stressed very easily.

Nonetheless, if you want/need to take them, it will be better to use rubber gloves:

  • put your hand in front of the scorpion,
  • nudge it from any side with a brush (pen, etc.),
  • the scorpion will turn around to face the ‘threat’,
  • nudge and slightly push it until it will back up onto your hand.
  • Heterometrus spinifer may behave defensively but they usually do not attack once they are in your hand.
  • DO NOT do sudden moves.
  • DO NOT press them down, they hate it!

Another word of caution is that Heterometrus spinifer can pinch you and even break the skin. As a result, you can flick your hand instinctively and your pet will go flying somewhere.  

Heterometrus spinifer can get hurt easily, even with their hard shells.

What is a bioactive enclosure for Malaysian Black Scorpion?

This is a self-sustaining ecosystem that does not require any (or little) maintenance. It also includes living organisms that work together in a balance within a tank.

Generally, when hobbyists are talking about a bioactive enclosure for their scorpions, they mean that there will be isopods and springtails to clean after their scorpions. These little bugs help to keep mold and decaying matter under control as well.

Warning: Some species of isopods are protein hungry. It can be dangerous for molting scorpions because during this process they are completely defenseless until their skeleton fully hardens.

Therefore, if you are not sure, it is better to play safe and add only springtails. These insects are detritivores that recycle dead organic material including scorpion’s poop.

Breeding Malaysian Black Scorpion

Maturity:

Heterometrus spinifer becomes adult and reaches sexual maturity in the 2nd to 3rd year. Generally, it takes them from 6 to 7 molts to reach maturity.

Sexing:

According to the studies, Heterometrus spinifer is not sexually dimorphic in its claws or tails. You have to check their pectines.

  • Look at their undercarriage behind the walking legs to find two little feathers (pectines). In males, pectines (the sensitive organs) are longer and more feathery. Females have comparatively smaller and less feathery pectines as compared to males.

It can be really hard to see for the untrained eye.

Mating and fertilization:

According to the study, the main structure of the courtship in different species of Asian forest scorpions is similar:

  • the male uses its claws to guide the female over a spermatophore during mating;
  • the female usually follows the leading male in a classical promenade (mating dance) until a suitable spot is found for spermatophore deposition;
  • then, the male assists the female in positioning her genital aperture over the spermatophore;
  • after that, the pair separates.

Generally, the mating dance lasts from 30 min up to 1.5 hours.

Gestation:

Females have gestation periods of up to 9 – 12 months and produce fairly small broods (20–40 nymphs).

Note: It is highly recommended to separate the male and the female as a mother scorpion becomes highly defensive.

Scorplings:

Malaysian Black Scorpion (Heterometrus spinifer) Breeding (scorplings)
photo by Timur Zinov

Heterometrus spinifer does not lay eggs. These scorpions give birth to young scorpions, generally called Nymphs or Scorplings.

This process is very stressful for the mother and can last 1 – 2 days. Do not touch or move anything for 2 days from the moment of birth. Keep her in a dark place and do not disturb her.

After that, start intensively feeding her. Check temperature and humidity. Make sure that there is enough airflow as well.

Scorplings are incapable of living separately from the female until their first molt. The mother carries the entire brood on her back. 

During this period, there is no need to feed baby scorpions, their mother will take care of that. Just give her more food than you normally would. She will catch the prey, tear it apart and let the babies feed on it.

As in most other scorpion species, about a week after the first molt, the second instars scorplings start leaving their mother’s back.

Tip: To improve the survival rate of scorplings, keep your temperature between 79 – 86°F (26 – 30°C) and humidity above 80%.

Separation:

Once scorplings leave its parent back, it will be better to separate them from the mother and each other as well.

At this stage, they need a lot of nutrition. Considering the fact they are often too weak to overpower their prey they can start cannibalizing fast.

Generally, females of Heterometrus spinifer do not hunt down their offspring. Nonetheless, natural predation can still happen, especially, if she is stressed. Therefore, the second instar scorplings should be separated from the mother.

They do not require a lot of space and can be kept in small food storage containers with adequate ventilation (just make small holes in the plastic). Provide them with damp substrate and hiding spots.

Baby scorpions have the same diet as adult scorpions but in smaller sizes:

  • small pinhead crickets (¼ inches or 0.5 cm long),
  • small worms,
  • fruitflies,
  • moths, etc.

Do not give them large live prey they cannot catch! It is recommended to feed them more frequently compared to adults – 2 times a week.

In addition, remove the water dish from their enclosure (tiny scorplings can easily drown there).

Malaysian Black Scorpion and Suitable Tankmates

It can be risky to house multiple Heterometrus spinifer in the same tank. Gravid females, young and sub-adults, in particular, are extremely likely to fight and even kill one another when housed together.

It is not recommended to keep Heterometrus spinifer with other species of genus Heterometrus. They will fight eventually.

Ideally, Heterometrus spinifer is better in solitary confinement.

In Conclusion

The Malaysian Black Scorpions (Heterometrus spinifer) are not hard to care for once you understand their needs and establish a routine.

Nonetheless, these scorpions are mildly venomous and are not very active. So, if you were looking for something to be constantly moving around or you can play with, then this is not the pet for you.

Scorpion is one of the most ancient animals on the earth and now we are keeping them as pets. You should only acquire them if you are willing to give them the special care and attention they need.

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References:

  1. Nirthanan, Selvanayagam, et al. “Assignment of voltage-gated potassium channel blocking activity to κ-KTx1. 3, a non-toxic homologue of κ-hefutoxin-1, from Heterometrus spinifer venom.” Biochemical pharmacology4 (2005): 669-678.
  2. Kovařík, František. “A review of the genus Heterometrus Ehrenberg, 1828, with descriptions of seven new species (Scorpiones, Scorpionidae).” Euscorpius15 (2004): 1-60.
  3. Stockmann, R., Ythier, E. in Scorpions of the World N.A.P edn 392 (2010).
  4. Kovařík, F. 2004. A Review of the Genus Heterometrus Ehrenberg, 1828, with Descriptions of Seven New Species (Scorpiones, Scorpionidae). 2004, No. 15.
  5. Systematic revision of the Asian forest scorpions (Heterometrinae Simon, 1879), revised suprageneric classification of Scorpionidae Latreille, 1802, and revalidation of Rugodentidae Bastawade et al., 2005 (Bulletin of the American Museum of Natural History, no. 442). Prendini, Lorenzo; Loria, Stephanie F.

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