Description and assessment of the Enigma syndrome (ES) in leopard geckos

(Eublepharis macularius) with regard to its relevance as trait of breeding

heritable defects

Frank Krönke, Munich, Germany

Abstract

The leopard gecko, Eublepharis macularius, is probably the most commonly kept lizard
worldwide. Characteristic for this species is that an enormous number of colour forms are bred,
so-called (colour) "morphs". Since breeding means a high proportion of inbreeding, it is not
surprising that hereditary diseases arise and spread, such as Enigma syndrome (ES) or
Iridophoromas in leopard geckos. The Enigma form appeared in 2006 and to date there is no
scientific assessment of it. This means that all questions concerning this disease are
unanswered.
This work has four aims: 1. to collect the known knowledge about ES. 2. Description and
analysis of film sequences of ES-affected leopard geckos. 3. Presenting a case history of a
leopard gecko suffering from ES. 4. Assessing ES related behaviour by comparing it with
normal behaviour of healthy leopard geckos, as well as with the help of scientific literature on
welfare assessment.
It seems to be a characteristic of ES that the onset of the first symptoms, the severity of the
course, the frequency of the acute seizures are mostly very individual. The chronic and acute ES
symptoms are described. The transition from chronic to acute form can be fluid in that
individual symptoms gradually appear. In terms of assessing wellbeing, the key feature of the
chronic form is the inability to eat independently. It follows that the animals are not capable of
surviving on their own, as healthy leopard geckos are, what can be assessed as severe
impairment. Concerning the acute form, it can be stated that all behaviours do not fulfil a
biological goal and are more reactive than active. It appears that the animal can only exercise
restricted control over the behavioural processes. For these reasons, the symptoms are
considered severe, too. Furthermore, it should be remembered that there is no known method
to alleviate or treat the symptoms.
In Switzerland, the Enigma form has been banned since 2015 because it has characteristics,
which are judged as "torture breeding". In all other countries, Enigma leopard geckos can still
be bred. This should be seen as particularly critical and ethically problematic, as potential
carriers of ES are produced and marketed against better knowledge.

Keywords:
animal protection, animal welfare, colour morphs, Enigma syndrome, Eublepharis macularius,
heritable defects, reptiles

Electronic copy available at: https://ssrn.com/abstract=4626177

1. Introduction

Enigma Sydrome is a hereditary disease in leopard geckos (Eublepharis macularius) as a result of

intensive (in)breeding. Its severity varies from mild to very severe, which in any case is
associated with significant to very serious restrictions in the wellbeing of the affected animals.
There is practically no literature on this defect, which probably has several reasons: domestic
reptile husbandry plays no role in scientific research, even in animal welfare science, where the
focus is on evidence-based husbandry improvements and thus on increasing animal welfare,
this area of animal husbandry is virtually overlooked, as it is in veterinary medicine.
Economically, it does not matter and the number of animals affected is not known. These
aspects form the context of the present study. It is a descriptive qualitative analysis of various
sources (video material, lay experts comments, observation of an affected animal). This study
is certainly not exhaustive, but it is intended to provide an introduction to the topic and
summarise the limited data available.

The leopard gecko is probably the most commonly kept and bred lizard in terrarium husbandry
(Tremper 2020). The reasons for this are many, but similar to other mass-produced species such
as king python, corn snake or bearded dragon: these species are usually neither very shy nor
aggressive, they are very adaptable to different housing conditions, they are often curious and
actively turned towards humans, they are easy to feed and easy to breed and finally, they are
also very variable in their natural habitat in terms of colour and pattern (Krönke 2020). The last
characteristic is also the origin of a very unique development of these species in captivity: the
breeding of so-called "colour morphs". However, the term "morph" is not usually used
correctly in this context. “Morphology" describes and analyses the structure and form of
organisms. But it is precisely the structure and form that is more or less the same in all leopard
geckos. In contrast, colours and patterns can be highly different. In this sense, it would make
more sense to speak of "colour forms".
The "realm" of colour breeding is divided into the three large sectors of leopard gecko keepers,
breeders and traders. The group of breeders represents a continuum: at one end, professional
large-scale breeders who "produce" 50,000 or even much more leopard geckos annually and
market them internationally (Tremper 2020; Geckoboa 2, 2023), and at the other end, simple
leopard gecko keepers who hatch a few clutches of their animals per year and offer them
locally (div. local selling platforms, e.g. Morphmarket 2023; Quoka 2023).
Breeding non-avian reptiles (hereinafter referred to as reptiles) is partly done differently than
breeding dogs. Two differences in particular are striking: 1. while the “stud books” are closed
for dogs (Gansloßer et al. 2021), they are open for reptiles. 2. the focus of dog breeding is on
keeping a breed pure (Gansloßer et al. 2021), whereas with reptiles new colour forms - through
mutation and line breeding - are bred in the sense of a pure breeding line (Geckoboa 3, 2023),
as well as all possible combinations of traits are tried out, whether purposefully or not (e.g. see
morph-catalogues of large-scale breeders at Geckoboa 1, 2023; Leopardgecko 2023; Geckosetc
2023; The Urban Reptile 2023; Geckoboa 2, 2023). Both approaches have specific advantages
and disadvantages. In dogs, it is known that due to strict inbreeding, more than 700 genetically
determined hereditary diseases exist (Gansloßer et al. 2021), which is why nowadays even
mixed-breed dogs are frequently affected by these defects.
Fortunately, the situation is better with reptiles. But here, too, some genetic defects are
known. In leopard geckos, for example, the ES - Enigma syndrome - in the "Enigma morph" line
(Jessica's Animals Friends 1), tumour formation (Iridiphoroma) in Lemon Frosts (Szydlowski et

Electronic copy available at: https://ssrn.com/abstract=4626177

al. 2020; Guo et al. 2021), balance disorders in some "White and Yellow" lines (Jessica's Animals
Friends 5, 2023; own observations), bone deformations in some “Super Snows” (Jessica's
Animals Friends 6, 2023) short snouts, slowed growth, smaller final sizes and lower fertility in
Black Nights (personal observation and communications with local breeders) are to be
mentioned. Red-eyed forms seem to be very sensitive to light and usually close their eyes in
(day)light, which can be understood as a sign of discomfort. The reason for this is a lack of
pigment in the iris, as the colourless iris does not sufficiently protect the retina from incident
light (personal observation; Pogona.ch 2023; Jessica's Animal Friends 6, 2023).
Most available information on the subject of ES in leopard geckos is of a non-scientific nature
and can be found on the internet. As expected, the quality of this information is
heterogeneous. Nevertheless, there are very knowledgeable private keepers who provide
valuable informations, as well as large-scale breeders who have sound experience with
heredity and have seen mostly hundreds of animals with genetical disorders.
At all, there is little information about ES, although the Enigma mutation appeared as early as
2006 (Leopardgeckowiki 2023). Primarily, it concerns specific characteristics of colour and
pattern. Only later it became apparent that a unique clinical picture is often also associated
with this mutation. The ES can develop in different degrees of severity. Even in a mild form, the
leopard geckos are usually not able to catch food insects on their own without restrictions, as
one of the typical symptoms prevents the targeted grasping of the prey. The animals snap at
the prey several centimetres away, although the prey is directly in front of their mouth
(personal observation; Jessica's Animal Friends 2, 2023). As far as I know, no scientific study has
yet been conducted to answer the question of whether ES is actually a neurological disorder
and the exact functional mechanisms are also unknown so far. And it is still unknown why
Enigmas can remain symptom-free for several generations and then suddenly have offspring
that show ES.
Working hypothetically, it can be stated that there are no ES free breeding lines (cf. Geckoboa
2, 2023). In the White & Yellow colour form, disease symptoms sometimes also occur that could
be of neurological origin and some of them are somewhat similar to ES. But in this form there
are clearly disease-free lines (Jessica's Animal Friends 5, 2023).
Talks with hobby breeders and the observation of various leopard gecko internet forums
suggest that due to the ES the popularity of this form of the leopard gecko has decreased
considerably, or the keepers mainly take care to breed only “healthy” animals (personal
communication with local breeders; Jessica's Animals Friends 5, 2023).

There is no universal definition of animal welfare because it is not only a scientific concept, but
also reflects ethical standpoints and different practical views (Jones et al. 2021; Mellor 2013;
Barber 2009). Animal wellbeing implies physical, behavioural and psychological aspects, as well
as their complex interactions. A good state of welfare means being healthy, well-nourished,
feeling comfortable, have a sense of security, being able to express a range of innate/normal
behaviours and the absence of suffering, fear, pain or persistent distress. Welfare is a long-term
state that comprises the sum of experiences of an animal and its derived affective state (Jones
et al. 2021; Blackett et al. 2017). For these states to be realised, coping abilities — physiological,
behavioural or cognitive — are an important factor, too, such as being allowed choice and the
ability/self-efficacy to reach desirable outcomes (Rose et al. 2017; Clark 2017; Barber 2009;
Young 2003: Baer 1998). “The importance of good health for good welfare is one of the least
controversial aspects of the debate about animal welfare” (Keeling et al. 2018).

Electronic copy available at: https://ssrn.com/abstract=4626177

This work has four aims: 1. to collect the known knowledge, especially from the internet, about
ES. 2. Description and analysis of film sequences of ES-affected leopard geckos. 3. Presenting a
case history of a leopard gecko suffering from the ES. 4. Assessing ES related behaviour by
comparing it with normal behaviour of healthy leopard geckos, as well as with the help of
scientific literature on welfare assessment.

2. Material and Methods

2.1 Source review

A search of scientific sources was conducted in the databases medline, biosis, embase and
google scholar. A general internet search was conducted using the search engine
"duckduckgo" with English and German search terms. In addition, various websites of large
breeders, internet forums specific to leopard geckos or local platforms offering leopard geckos
for sale were specifically consulted.
A descriptive, qualitative evaluation of publicly available data sources, i.e. personal reports
from experienced keepers of leopard geckos with ES and film sequences of animals affected by
ES (n = 14) - especially on YouTube or Facebook plattforms - provided a basis. Another was the
direct observation of an ES-affected leopard gecko, which is presented as a case description.

2.2. Case description

The observation data were collected on a single ES leopard gecko on 13 observation days. The
length of a single observation unit was 60 min, resulting in a total observation time of 13 hours.
Observations were made in the evening between 8 pm and 10 pm, during the gecko's activity
period. A dim room light was available, which allowed all details in the terrarium to be seen. It
did not seem that the gecko felt disturbed by this setting. The behavioural elements were
recorded with pen and paper and later transferred to Excel tables and evaluated. In addition to
this intensive data collection, another extensive survey was conducted by spotting behaviour
approximately every 30 to 60 min between 5:30 p.m. and 1:00 a.m. over a period of about 6
weeks. The purpose of this survey was not to record detailed activity, but to record activity
over the course of the activity period to get an idea of whether the animal was behaving more
actively or passively.

2.3 Housing and husbandry

The study animal was female, about one year old, 16 cm long (total length) and weighed 19
grams. It came from a private owner. Both the dam and one sibling were asymptomatic.
Nothing was known about the sire. A faecal examination revealed an infestation with oxiuria
but no cryptosoridia. Three treatments with Panacur were carried out. The general condition
was good, the animal had always eaten with great appetite from the first day it was taken over.
For a fortnight, the gecko was housed in a quarantine box measuring 35 x 20 x 14 cm (length,

Electronic copy available at: https://ssrn.com/abstract=4626177

depth, height). This was equipped with a layer of kitchen paper, a shallow water dish, a moist
box, a plastic hiding place, and a heating mat 8 Watt, 22 x 22 cm under the hiding place.
Later on the leopard gecko was housed in a terrarium tailored to its specific needs: 60 x 40 x 40
cm (length, depth, height), 5 cm clay-sand mixture as substrate, two moist boxes (13 x 9 x 6 cm,
length, depth, height) with a layer of approx. 4 cm of moist coco humus, a heating mat 8 Watt,
22 x 22 cm, under the terrarium in the left corner, a heating rock, 10 Watt, in the right corner, a
shallow water bowl, several flat stone slabs, as well as two plastic hiding places (16 x 8 x 5 cm,
length, depth, height). The gecko had a usable surface area of about 5000 square centimetres,
with about 1/3 without cover and 2/3 under a structure providing cover. No additional lighting,
as the terrarium was in a room with several large windows. At least 50% of the back and side
windows of the terrarium were covered with dark cardboard to reduce the amount of
perceptible stimuli for the gecko.
As the leopard gecko was too small for its age and too light for its size, daily feeding with wax
moth larvae, crickets and mealworms took place after each observation. The food was either
offered by tweezers or in a small bowl (mealworms), from which the gecko could sometimes
eat on its own. Every food-item was dusted with a calcium vitamin preparation (Korvimin ZVT +
Reptil).

2.4 Comparative data set

As it is difficult to assess behavioural data without context, a data set was used that is based on
identical behavioural categories and describes normal behaviour of leopard geckos in the
terrarium without the presentation of additional stimuli (see table 1, Krönke et al. 2023).

3. Results

3.1 Source review

3.1.1. The Enigma colour form

As the phenotype is dependent on the genotype, the appearance of an Enigma leopard gecko
is always dependent on the respective colour form(s) with which it has been "mixed".
Nevertheless, the following features can be characteristic: small dot-like pattern elements,
basic colour of the tail white to bluish-grey. In matings with orange forms ("tangerines") red
eyes may occur without an amelanistic form ("albino") being crossed in at the same time.
Furthermore, an increase in colour intensity often occurs in these matings (e.g. Der
Leopardgecko 2023). In addition, there are numerous other effects of colour and pattern in
conjunction with the Enigma form (SaSobek Reptiles, 2023). Therefore, Enigmas usually look
different and are sometimes difficult to recognise as Enigmas. From time to time ES similar
symptoms occur in non-Enigmas. It is unclear whether these symptoms are independent of the
ES or whether these animals are "masked", i.e. phenotypically not recognisable as Enigmas
(Jessica's Animal Friends 2, 2023).

Electronic copy available at: https://ssrn.com/abstract=4626177

3.1.2 Symptoms

A search in a leopard gecko facebook group with about 40,000 members has shown that an
important topic are the identification of ES symptoms and the handling of ES animals (All about
leopard geckos 2023).

Onset of ES symptoms can occur at hatching or only after several, completely symptom-free
years (Jessica's Animal Friends 2, 2023). Balance and coordination disorders are characteristic
of the ES which results in often large, uncoordinated and hectic movements. The formation of
the symptoms can vary considerably. Thus, on a continuum from very severe to very mild, all
intermediate forms can occur (Jessica's Animal Friends 2, 2023; Care guide for lizards 2023).
Some authors claim that ES can worsen with age and that, especially in females, reaching
sexual maturity and/or their annual ovulations often worsen ES symptomatology (Jessica's
Animal Friends 2, 2023). Symptoms may occur in isolation from each other, immediately one
after the other, or in conjunction with normal behavioural elements. In mild forms, an ES
symptom, such as head raising, is performed respectively the movement is started, but not
completed and is immediately transformed into another normal movement (“transitional
movements”) (personal observation). Bright light, flashing light, loud noises, touching can
have a triggering effect. Moreover, poor health or poor housing conditions can also be
triggering factors. Depending on the condition of the animal, this can cause mild symptoms
such as head raising or circling, or more severe symptoms such as seizures or lateral rolling
movements (Jessica's Animal Friends 2, 2023). However, seizures – in a sense of an umbrella
term for the acute occurrence of ES symptoms - can also be initiated without traceable triggers
(personal observations).

The following symptoms of abnormal behaviour are characteristic of ES:

Inability to catch feeding insects (having bad aim):

• snap past feeding insect by several centimetres, even if it is directly in front of their
mouth
• gecko can react differently to various feeding insects
• always react more intensively than a healthy leopard gecko would do

tottering / wobbly gait:

• can be very different in intensity
• can be triggered by water drops on the floor, for example, or occur during ordinary
locomotion
• exaggerated lifting of one foot for a walking movement is also possible
• in severe cases, normal gait is no longer possible
• in severe cases, the walking direction cannot be maintained

head raising / ”star gazing”:

• from a movement or also from a resting position
• head is turned upwards at an angle of approx. 45 – 100°
• similar behaviour is also shown by healthy leopard geckos, but much less frequently and
at an angle of inclination ≤ 90°
• for seconds or minutes

Electronic copy available at: https://ssrn.com/abstract=4626177

twisting the head sideways:
• often always to the same side
• as an isolated symptom or together with other symptoms, for example together with
head raising or circling

behavioural agitation / restlessness:

• movements are started but not completed
• frequent change of the direction of movement
• no or little goal orientation
• insertion of "meaningless" behavioural elements, such as head bobbing, head tilting,
circling

closing the eyes:

• during a seizure
• during every possible movement
• in situations in which a healthy leopard gecko would not or rarely close its eyes, e.g. in a
tense posture

circling:
• running in circles, sometimes similar to a dog trying to catch its tail
• different frequency
• sometimes lasting for several minutes

physical and cognitive overstimulation / overexcitability:

• it appears that there is a hypersensitivity of the skin to any kind of touch stimuli
• a shedding can trigger such an overreaction if, for example, remnants of the old skin rub
against the new skin, which in turn can trigger strong circling
• it also appears that, for example, a moving feeding insect can cause cognitive
overstimulation, resulting in a much more intense and sustained response than in a
healthy leopard gecko
• sometimes very rapid movements of the extremities, which can be similar to vibrating

head tilting:
• lateral pendulum movements of the head, which can be unilateral or bilateral
• these movements can occur during locomotion or from a resting position

head bobbing:
• vertical pendulum movements
• often in connection with head raising

stiffening of the body and extremities (sometimes referred to as seizures):

• predominantly in severe ES forms
• stiffening of the body and extremities, which is accompanied by a temporary inability to
move
• sometimes spine pushed through to the front
• eyes often closed

Electronic copy available at: https://ssrn.com/abstract=4626177

 • usually lasting only a few seconds

lateral rolling movements (“death rolling”):

• predominantly in severe ES forms
• flipping over
• falling on the back
• rigid lying down or strained attempts to turn over again
• often in connection with stiffening of the body

states of exhaustion:
• after a seizure
• for minutes or hours

seizures:
• umbrella term for the acute occurrence of ES symptoms

(Jessica's Animal Friends 1, 2, 3, 2023 The Reptile Guide; Krönke unpublished data 2023; Dendy
Charl 2023; Wildfire Geckos 2023; Enigma Syndrome & Neurological Issues in leopard geckos
2023; Faszination Leopardgecko 2023; Forsaken December 2023; Geckoland 2023; Ice Cream
2023; Happy Gecko Farm 2023).

As a general assessment of ES, it can be said that the more frequent and intense the
symptoms, the more impaired the quality of life.
The only hypothesis on the cause, function and inheritance of ES is an online article by Ben
Bargen from 2013, which, however, has not yet been verified at the genetic level (Geckoboa 3,
2023). Bargen claims that ES affects balance and cognition and refers to this syndrome as a
neuro-degenerative disease. The tricky thing is that the first symptoms often appear late,
which is why breeders are lulled into the certainty of having healthy animals for a long time and
therefore continue to spread the harmful mutation. Environmental factors and individual
genotype would cause ES to be expressed differently in each individual. Environmental factors
include, for example, cage-mates, shipping or other stressors. He also includes other similar
syndromes, such as WYS (White & Yellow Syndrome), with ES and says that all forms of colour
are susceptible to this "balance disorder syndrome". Regarding inheritance, he says that this
syndrome is inherited in an autosomal dominant (AA) manner, i.e. homozygous recessive (aa)
animals do not develop the disease. Since the pure-bred form usually leads to the death of the
embryo, the affected animals are usually heterozygous (Aa). On the genetic level, there could
be a loss of function, which, similar to Huntington's disease in humans, causes a disturbance of
a protein synthesis. According to Bargen, these non-functional proteins could lead to partial
cell death in the brain, which could explain the symptoms. However, these hypotheses have
not yet been scientifically verified.

For a less experienced pet owner or vet, several symptoms come to mind that may be
superficially similar to ES: i.e. MDB (metabolic bone disease), injuries/head traumas, age-related
physical limitations, vision/eye problems (Jessica's Animal Friends 4, 2023).

Differences between ES and WYS:

• ES often gets worse with age, whereas WYS can get better

Electronic copy available at: https://ssrn.com/abstract=4626177

 • ES is linked to the Enigma colour form, whereas there are many WYS-free lines.
• ES is clearly more severe than WYS

Similarities ES and WYS:

Staggering gait, motor overreactions, falling over, overstimulability (Jessica's Animal Friends 5,
2023).

Picture 1: Spreading the legs and strong head tilt during a seizure

Electronic copy available at: https://ssrn.com/abstract=4626177

Picture 2: Circling movement and tilted head position

Picture 3: Laterally twisted head raising

10

Electronic copy available at: https://ssrn.com/abstract=4626177

Picture 4: Extreme inclination of the entire body shortly before falling on the back

Picture 5: Circling

11

Electronic copy available at: https://ssrn.com/abstract=4626177

Picture 6: Closing the eyes during a seizure while keeping the head tilted

3.1.3 Husbandry and housing recommendations for leopard geckos with ES:

The only possible "treatment" for ES is needs-based housing and husbandry (Jessica's Animal
Friends 1, 2023). The following recommendations should be adapted to the health condition of
the respective leopard gecko. Animals with mild symptoms may be kept almost like healthy
geckos and only need assisted feeding. The most important rule is: anything that could cause
stress should be avoided. ES-leopard gecko also require closer daily monitoring than healthy
animals.

• they should be housed individually to prevent competition during feeding or "assaults"

due to uncoordinated body movements
• restricting touching and handling to the necessary minimum; slow and sensitive
movements of the hands
• no loud or sudden noises or vibrations
• cover or tape the glass walls of the terrarium with cardboard to reduce the number of
visual stimuli
• tweezer feeding, in mild cases feeding mealworms from a small bowl (cannot run away)
• assistance with shedding, e.g. by staying on damp kitchen paper in a plastic box
• no or only very low lighting of the terrarium during the day
• balanced diet with the addition of calcium vitamin supplements
• no objects in the terrarium that can fall over, are pointed or have sharp edges
• cork bark is very practical as it provides structure when, for example, a leopard gecko

12

Electronic copy available at: https://ssrn.com/abstract=4626177

 lying on its back tries to turn around again and it is soft so that the animals cannot injure
themselves
• several hiding places
• no high structures from which the gecko could fall down
• no structures on which it could get caught
• only simple-furnished terrariums
• no changes, stable conditions
• smaller terrarium size, e.g. 60 x 40 x 40 cm (length, width, height)
• some animals are oversensitive to water droplets, so do not spray
• some animals are oversensitive to a natural substrate, so use kitchen paper, cardboard,
carpet, cork wallpaper or similar
• no breeding, because it is very stressful and because the genetic defect should not be
spread further

(Care guides for pet lizards 2023; Jessica's Animal Friends 1, 2, 3, 2023; own experiences;
Reptilefiles 2023)

ES-leopard geckos probably have no pain and not only bad moments, i.e. these animals have a
reduced quality of life, but not no quality of life (Jessica's Animal Friends 2, 2023; Frank Krönke
personal observations 2023).

3.1.4 Some open questions

There are sources saying, at least in the early days when Enigma mutation was new, there were
many purebred Enigmas (e.g. Der Leopardgecko 2023). Other sources, however, claim that
purebred Enigmas are not viable and die already in the egg (Jessica's Animal Friends 2, 2023;
Care Guides for Pet Lizards 2023). And yet other sources say that homozygous Enigmas are not
necessarily incapable of life, but usually develop severe symptomatics (Care guides for pet
lizards 2023). One reason for this ambiguity could be the not yet understood mechanism that at
least phenotypically healthy animals suddenly produce sick offspring after several generations
I.e. the insufficient clarity regarding the inheritance of ES.
In this sense, there are voices claiming that every Enigma carries at least the potential to
become ill (Jessica's Animal Friends 2, 2023) and large breeders have excluded the Enigma
from their breeding for this reason. Despite intensive efforts, the latter have not succeeded in
separating the desired phenotypic traits from the genetic defect (Geckoboa 2, 2023). On the
other hand, there are large breeders, as well as hobby breeders, who regularly cross Enigmas
with other colour forms and ES symptomatology occurs only rarely or not at all (personal
communication with local hobby breeders; Geckosetc 2023; Imperial Reptiles 2023). It is
claimed that the Enigma trait is inherited in an autosomal dominant manner (Bargen 2013).
Nevertheless, there are observations that deviations from this pattern might also possible
under certain circumstances (Der Leopardgecko 2023; All about leopard geckos 2023; Leopard
Gecko Life 2023).
So, despite the profound experience of large-scale commercial breeders, the exact inheritance
of ES may not yet be fully understood. Even the “genetics" of the different colours and
patterns in the leopard gecko are still far from being understood (Geckoboa 3, 2023; Bargen
2013). The location of this mutation in the genome or a detailed description of this mutation at

13

Electronic copy available at: https://ssrn.com/abstract=4626177

the genetic level also does not yet exist. Especially when the syndrome "overstimulation" is
considered, the suspicion of insufficient stimulus processing is obvious. This is another aspect
that has not yet been clarified. It is not even known what exactly ES is from a medical
perspective, what causes the symptoms, whether it is actually a neurodegenerative disease.
There are no data available that allow a reliable statement on the development of ES
symptoms over time, i.e. whether ES worsens with increasing age or whether the course is
different for an animal with very early-onset symptoms and one with late-onset symptoms.
Furthermore, the question of why ES starts very early in some animals and only after years in
others remains unanswered.
Differences, similarities and correlations between ES and WYS are also unclear as well as the
clarification of whether ES and WYS are the same disease in different manifestations or
different causes with similar symptoms. (e.g. All about leopard geckos 2023; Leopard Gecko
Life 2023).

3.2. Case report

3.2.1 Context description

The symptoms described below are a mild form of Enigma syndrome.

I got the leopard gecko from its previous owner on 2 December 2022. It was female and is
supposed to have hatched around February 2022. It weighed 19 g and had a total length of
about 16 cm, which was a bit too small and light for its age. The previous owner acquired the
animal when it was about 4-5 months old. From the beginning, it could not eat insects on its
own, because it usually snatched at them several centimetres away.
The leopard gecko seemed to be a "motley mix" of different colour forms. Phenotypically, the
following features were visible: pure black eyes (possibly Eclipse), a small, dot-like pattern on
head and tail (Enigma), white flanks and sometimes a strong yellow back (White & Yellow),
sometimes a predominantly white-pink colouration of the entire body (perhaps Blizzard or
similar).
At first, the only behavioural abnormalities of this otherwise healthy, normal-looking animal
were a weakly pronounced movement coordination disorder ("wobbly gait"), which occurred
from time to time, as well as the inability to snap precisely at a prey. More often than normal,
the head was stretched upwards, while at the same time twisting to the left. Nevertheless, this
symptom was clearly different from the classic "star gazing", in which the head is stretched
upwards at an angle of ≥ 90°. From the beginning, during forceps feeding with small crickets,
the animal showed marked discomfort when touched at the mouth with the cricket's legs or
antennae. This behaviour would fit a hypersensitivity to touch stimuli. The gecko snaps 2-3,
sometimes 5 cm next to the insect food with great regularity. Interestingly, this animal always
snapped past the prey to the left. Furthermore, the gecko showed a few times a tilting of the
head to the left side.
What was striking about this animal was its enormous activity inside the quarantine box. It was
very active every evening for about two hours and explored intensively. Overall, however, this
behaviour appeared normal and healthy. The proportion of conspicuous behavioural elements
was low. The previous owner kept the animal in a small opaque box with very little stimuli. The
move to the quarantine box might have acted as a reinforcer of the explorative behaviour, too.
Because this leopard gecko was smaller and lighter than average, it was offered food daily and

14

Electronic copy available at: https://ssrn.com/abstract=4626177

gained significantly in size and weight: At the beginning of January 2023, it weighed 37.9 g and
had a total length of about 19.5 cm.
It was attentive and also actively perceived the "world" outside the terrarium: e.g. every
second time somebody stood in front of the terrarium, it came over and actively looked. In the
terrarium, it usually stayed in one of the numerous hiding places during the day and on or next
to the heat rock in the evening. The activity was quite different: some evenings it explored the
terrarium and used almost the entire available surface and other evenings it spent sleeping or
dozing on or next to the heat rock.
At the end of December, there were two behaviours that were ambivalent in their
interpretation: a body turn with a forward movement and a head raising followed by a body
turn. Both could be normal, healthy behaviours, but it would also be possible to see this as an
kind of "transitional movement". The latter would mean that the animal started to perform an
ES symptom - circling or head raising - but counteracted it and transformed it into another
healthy movement. These supposed transitional movements have been observed several
times. Apart from this, this leopard gecko was active at the same time, showed normal
behaviour and appeared inconspicuous or healthy.
The colouring of this leopard gecko also changed at the end of December. Whereas it had
previously been exclusively whitish-pink in colour, with a few light yellow lines on the back,
from then on the entire back showed itself in a featureless strong yellow. Only the flanks
remained white, as was typical for the "White & Yellow" colour form.
On 5 January, as well as in the two following days, the animal repeatedly showed a "squinting
of the eyes", which is untypical for normal behaviour of this kind. Furthermore, a "long
stretching or stretching" of the neck, which I have never seen in healthy leopard geckos and
could therefore be connected with ES. The transitional movements described above were also
observed several times. Furthermore, the gait was less coordinated: a "wobbly gait" was
shown several times. On 7 January, the animal was very inactive in the evening, lying on the
heat rock and giving the impression that it was doing well.

3.2.2 Acute ES Symptomatology

On 8 January, the animal showed its first severe attack/seizure of ES. A check with the previous
owner revealed that this leopard gecko had not had a seizure before. First, there was a very
strong left head torsion and shortly afterwards circling movements. This head torsion was a
characteristic of this animal and was maintained almost without interruption in the following
days. Furthermore, the following behaviours were frequently exhibited: Eye squinting, a very
wobbly gait and a behavioural restlessness, i.e. it was very active, but showed the behavioural
elements in too rapid sequence, furthermore many movements were not completed, but
instead new movements were started, which in turn were also not completed. In contrast to
the classical "star gazing" where the head is raised at an angle of ≥ 90°, this Gecko only showed
an angle of inclination of about 45° to max. 80°. The circling movements consisted of a
complete or only a 3/4 circle turn with a wobbly gait and frequent tongue flicking, exclusively
left around. They took place both without cover and under cover. Transitional movements also
occurred repeatedly. During this seizure, only a few normal movements were performed. The
acute seizure lasted about 45 minutes. Afterwards, the animal retreated under cover. It rested
there for about 30 minutes and then started circling movements again. The movements were
somewhat slower and slightly more coordinated. A neck stretching occurred. The gecko fell on

15

Electronic copy available at: https://ssrn.com/abstract=4626177

its back once out of an uncoordinated movement, but this was clearly different from a "death
roll".
The course and symptoms of the second seizure were similar to the first, frequency and
intensity were slightly lower. The circling movements were not always fully executed. The
animal licked its eyes several times. It was bright yellow all evening. Short rest periods in
between. Observation was terminated after 4 hours, the animal still showed circling
movements.
On 9th January the animal hardly moved, still held its head in a strong left twist. On 10th
January it was more active again, ate 3 crickets from the tweezers. Not only the head was
twisted to the left, but also the upper part of the body. Frequent tongue flicking and
pronounced behavioural restlessness, closed eyes during head movements. Later circling
movements and head up ≥ 90° with simultaneous left head twisting. Fell over during body
rotation and lay on the back for a short time. Resting after about five minutes of circling with
eyes closed and body position clearly tense. Did not react to opening of the terrarium, avoided
light. Rested on heat rock for a longer period of time, appeared fatigued, as if these attacks
were very strenuous and energy-sapping.
11th January still had head strongly tilted, walked around the terrarium, looked out. Head up ≥
90° with sideways twist had increased in frequency, compared to previous days. Before that it
was rather a transitional movement. Gait was somewhat more coordinated, locomotion
seemed strenuous. Explored again actively and attentively through the entire terrarium and
came to the open terrarium pane. Rested with closed eyes, but tense posture, which did not fit
normal behaviour. So the ES symptomatology was not yet suspended on the 3rd day after the
initial seizure, but significantly reduced. The symptoms slowly faded out. From January 13, it
was very little active for two weeks, but again kept the head straight throughout. It moved
from the daytime hiding place to the heat rock, where it spend the evenings.
By December 2023, the animal only showed mild symptoms (head tilting, circling) once during a
period of 5 hours. The next day, all symptoms had disappeared again. The gecko has also
become well accustomed to eating mealworms from a bowl independently and continuously.
Since then, the leopard gecko has behaved completely unobtrusively.

3.2.3 Quantitative behavioural data before the seizure

Table 1: Activity measured in behavioural units and behavioural categories

∑ behavioural ∑ behavioural all basic all resting sense of sensual all walk
elements categories needs interest behaviour security exploration around
7158 179 11 1375 75 162 4684 628
% - - 0,2 19,2 1 2,3 64,5 8,8
50%* 3579 90 6 688 38 81 2342 314
Krönke et al. 696 306 8,8 96,7 75 55,1 317,9 114,4
2023 average /
animal
% - - 1,3 13,9 10,8 7,9 45,7 16,4

*Each observation unit corresponds to 60 min. Since in the comparison data set an observation unit lasted only 30
min, the number of results is halved for better comparability. The baseline survey serves as a comparison

16

Electronic copy available at: https://ssrn.com/abstract=4626177

Because only a single animal was involved, the data analysis was limited to a few obvious
aspects. Table 1 shows that the total number of behavioural elements was higher than in the
comparison data set by a factor of 10.3, whereas the total number of behavioural categories,
i.e. different behaviours, was lower by a factor of 1.7. It was also striking that the ratio of
behavioural elements to behavioural categories differed by a factor of 40 in the case report,
but only by a factor of 2.3 in the comparison data set. This means that the Enigma animal was
much more active than the average healthy leopard gecko of the comparative study, but
expressed this mainly in behavioural intensity and not in behavioural diversity. Nevertheless, it
cannot be inferred from this that this behavioural profile was typical of Enigma leopard geckos.
Regardless of the numbers, the Enigma animal appeared predominantly normal before the
seizure and only in a few aspects somewhat conspicuous: the "head up" position was adopted
more often than by healthy leopard geckos and, as in the latter, different tilt angles could be
distinguished. Furthermore, coordination disturbances in the gait pattern could be seen, but
these occurred, depending on the daily form, with different frequency. In summary, a
qualitative description of behaviour provided more informative data than quantitative.

4. Discussion

The purpose of this study was 1. to collect all available information on ES, 2. to describe and
analyse film sequences of ES-affected leopard geckos, 3. to intensively observe an ES leopard
gecko and to write a case report, and 4. to assess the wellbeing of ES leopard geckos based on
the collected data and comparison with behavioural data of healthy animals (Krönke et al.
2023), as well as with the help of scientific literature on welfare assessment. Furthermore, it
seems necessary to include an ethical consideration, as the leopard gecko is probably the most
frequently kept lizard species in terrariums worldwide (Tremper 2020, Henkel et al. 2013,
Grießhammer et al. 2009) and therefore the potential impact is particularly high. It is
astonishing that the ES has so far been ignored by both science and the authorities, even
though the Enigma colour form appeared as early as 2006 (Leopardgeckowiki 2023).

As mentioned at the beginning, there is currently no uniform concept of what exactly is meant
by animal wellbeing (Jones et al. 2021; Dawkins 2021; Mellor 2013; Barber 2009). Nevertheless,
the following aspects always play a role: physical components such as nutrition, environmental
factors, health or behaviour and mental components such as subjective, affective states and
the role of individual experiences. In most approaches, the latter is considered to be of major
importance for wellbeing (Mellor 2013). When assessing the wellbeing of Enigma leopard
geckos, it is assumed that these animals are kept under conditions of good animal husbandry,
and therefore the aspects of nutrition, environment or housing and husbandry do not play a
role. However, if this is not the case, neglecting these factors makes Enigma leopard geckos
more vulnerable than healthy animals. The aspects of health, behaviour and subjective
affective state are of central importance.
A more biological definition of animal welfare is: “The essence of good welfare is […] being
currently healthy and also having good prospects for future health” (Dawkins 2021). Certainly,
this partial definition does not encompass everything that is understood by welfare today, but
it nevertheless emphasises a central aspect that is relevant above all others. For the Enigma

17

Electronic copy available at: https://ssrn.com/abstract=4626177

form of the leopard gecko is a mutation that is selected by humans, i.e. positively evaluated
and deliberately spread through breeding, EVEN though it jeopardises this most basic
requirement of being healthy now and in the future because of personal preferences and
interests. The second part of Dawkins' definition refers to bringing “what the animal wants”
into focus. And here, too, what has just been said applies, because no animal will want to suffer
from ES symptoms.

4.1 Assessment of wellbeing of ES leopard geckos

First, it appeared useful to differentiate two time axes that play a role in the context of ES:
chronic disease and acute disease. Furthermore, it seems to be a characteristic of ES that the
onset of the first symptoms, the severity of the course, the frequency of the acute seizures are
mostly very individual.

4.1.1 Chronic disease

Chronic aspects of ES are the following:

• Inability to catch feeding insects
• tottering/wobbly gait
• head raising
• twisting head sideways
• physical and cognitive over excitability
• neck stretching
• squinting of the eyes

Due to the low number of cases used as a basis, deviations are likely. It is important to know
that some of these behaviours are part of normal behaviour of leopard geckos and are
especially noticeable because of their greater frequency, such as head raising. Other
behaviours, such as squinting of the eyes, are very similar to normal behaviour but differ in that
they are performed in a tense posture, for example.
In terms of assessing wellbeing, the key feature here is the inability to eat independently. While
the animals may be able to eat mealworms from a shallow dish, this method is not reliably
successful. It follows that the animals are not capable of surviving on their own, as healthy
leopard geckos are, what can be assessed as severe impairment. In terms of the theory of
wellbeing, this means that there is both a health-related functional impairment and a
behavioural impairment. In the short term, the inability to eat on one's own is an experience of
frustration (cf. Lambert et al. 2019), and in the long term, malnutrition, debilitation, loss of
function and death are associated with it, as the most severe form of lack of wellbeing.
The other symptoms are to be seen on the one hand as an expression of indisposition (e.g.
head raising, squinting eyes, neck stretching) and on the other hand of a physical and/or
neurological restriction (e.g. wobbly gait, twisting head sideways, over excitability). As these
only occur occasionally in terms of their frequency and intensity and are comparatively mild,
and normal behaviour predominates, the symptoms can be judged to be little impairing.
Certainly, there are also cases in which chronic symptoms are more numerous and more
pronounced. This means that the assessment of wellbeing cannot be made in general, but in
relation to the individual animal.
A conspicuous component of the chronic symptoms in particular, and sometimes also of the

18

Electronic copy available at: https://ssrn.com/abstract=4626177

acute symptoms, is frequent tongue flicking and transitional movements. The former could be
understood as an increased need for orientation and the latter as an attempt to correct
unpleasant behaviour with another behaviour. Both would therefore be an attempt to actively
influence the situation that was presumably perceived as unpleasant.

4.1.2 Acute disease

The transition from the chronic to the acute form can be fluid in that individual symptoms
gradually appear in addition and are initially only pronounced with little intensity or are
transitional movements which increasingly change their form from an ambivalent to a clear ES
symptom.

Acute aspects of ES are the following:

• behavioural agitation/restlessness
• circling
• head tilting
• head bobbing
• stiffening of the body and extremities
• lateral rolling movements
• after a acute seizure: strong exhaustion

In contrast to chronic symptoms, all acute symptoms are clearly different from normal
behaviour, which could be seen as an indication of a link to reduced wellbeing. Furthermore, all
behaviours do not fulfil a biological goal and are more reactive than active in the sense that the
animal does not decide to do something, but is "drawn" into undesirable behaviour by the
symptoms. It appears that the animal can only exercise restricted control over the behavioural
processes. For these reasons, the symptoms are considered severe. This assessment is further
strengthened by the fact that the animals appeared to be exhausted in the aftermath of an
acute attack. Since good animal welfare is considered to result both from an absence of
negative experiences and from the presence of positive experiences or sensations (Mellor
2013), it can be unequivocally stated that due to the ES symptoms, wellbeing is considerably
reduced. As a general assessment of ES, it can be said that the more frequent and intense
these symptoms, the more impaired the quality of life. Furthermore, it should be remembered
that there is no known method to alleviate or treat the symptoms. Even in the case of a mild
seizure, as described in the case description, which affects the animal for four days.
One factor that may have been the trigger for the leopard gecko's seizure is the possibility of
sexual maturity having occurred. This assumption would fit with its age and growth spurt (cf.
Jessica's Animal Friends 2, 2023).

4.2 Assessing the quality of life of leopard geckos with ES

As more observation material on ES was available from the case description, but the ES
examples from internet sources were mostly more severe in their symptomatology, an
assessment of wellbeing is carried out separately for both data sets. A look at the recent
literature on animal welfare science shows that good welfare is much more than just the
absence of negative experiences. The presence of positive experiences as pleasure is a central

19

Electronic copy available at: https://ssrn.com/abstract=4626177

component of good welfare. Receiving food and the process of foraging or comfort behaviour
are seen as expressions of pleasure, i.e. positive affects, in animals (Boissy et al. 2007). Coping
and controllability are mechanisms of dealing with the environment that have a strong
connection to the individual's wellbeing. The possibility of influencing one's environment and
successfully overcoming challenges also cause positive affect (Boissy et al. 2007). Conversely,
this means that a reduced possibility of exerting influence and positive coping either does not
allow the positive affects to arise and/or instead negative ones do. This approach seems to me
to be justified in relation to ES, since the leopard gecko of the case description actively tried to
exert a corrective influence on ES symptoms with the help of transitional movements and
frequent tongue flicking. This means that the animal suffers not only from the acute
symptoms, but also from the frustration of not being able to intervene in the (disease) process
in a regulating way. This is probably also true for other ES leopard geckos, but based on the
short video sequences on the internet, this cannot be confirmed for all affected animals. From
this it can be deduced that due to the non-treatability of ES, all necessary precautions should
be taken to exclude symptom-triggering stress, injury possibilities or malnutrition. This can
contribute to an increase in wellbeing.

4.2.1 Assessing the quality of life of a leopard gecko with ES (case description)

As mentioned above, not much can be deduced from the observation data of a single animal.
The pronounced activity before the seizure and the fact that this behaviour was predominantly
normal behaviour, i.e. fulfils a biological purpose, such as exploration or display of wellbeing
(resting heat rock), indicates that the animal felt well. Neither escape behaviour (e.g. pane
standing or pane scratching) nor shut down behaviour (e.g. staying in hiding place) was shown,
which would have been a clear indication of discomfort. Another indication of well-being was
the fact that the animal was only active during its natural activity time. The rest of the time was
spent sleeping or resting in a hiding place, which is also normal behaviour (Mohanty et al. 2021;
Libourel et al. 2015). It can thus be stated that a leopard gecko suffering from ES can certainly
have a good quality of life outside of its seizures. Here, too, the aforementioned applies that an
assessment cannot take place in principle, but only bound to the individual. Krönke et al. (2023)
serves as a basis for comparison and description of "normal behaviour" in leopard geckos: the
characteristics of wellbeing were described and analysed with about 380 hours of observation
time.
Recent research has shown that the quality of the animal-human relationship can be
considered a key factor in how an animal interacts with its environment. In this sense, the
increase in a positive animal-human relationship can be understood as an increase in the
welfare state (Jones et al. 2021). Consequently, it can be concluded that the active interest of
the leopard gecko in people who are in front of its terrarium or open the windows of the
terrarium, as described above, can be understood as an expression of welfare. Another
example: if the hand was held in the terrarium, the leopard gecko regularly smelled it,
sometimes climbed onto the hand and rested in the warm hands for about a quarter of an hour
before becoming restless.

20

Electronic copy available at: https://ssrn.com/abstract=4626177

4.2.2 Assessing the quality of life of leopard geckos with ES (internet sources)

There are numerous videos on the internet that show ES in many different facets (All about
leopard geckos 2023). Many of the examples show severe, acute forms of ES. As the film
sequences are short and the clips focus on the main symptoms, there is little contextual
information. It seems unquestionable that such an animal is unable to control its behaviour
during a seizure. None of these behaviours serve a biological purpose. In the analogy from the
case description, it is assumed that these states are perceived as very stressful and unpleasant.
There is little information about the frequency and duration of these seizures and also details
about the animals (e.g. age, sex, onset of symptoms, development of symptomatology, etc.).
The only conclusion that can be drawn from this is that the more frequently these seizures
occur, the more the well-being of the respective animal is impaired. However, this also means
that the geckos can have phases of relatively good wellbeing between the acute phases.

4.3 Ethical evaluation of the breeding and distribution of leopard geckos of the Enigma form

“As human caretakers, with complete control over captive animals' care and environment, we
have an ethical obligation to safeguard, and indeed promote the welfare of those animals
under our care” (Benn et al. 2019). In a moral philosophical consideration, it can be stated that
human beings should treat each subject with respect and not as a mere means to its ends. That
is, we should not deal with them in a way that disregards their autonomy, that is, their ability to
decide how they want to act or want to be. What matters, is that individuals be recognised as
having inherent worth, valuable in themselves, regardless of their usefulness to others. And
because they have inherent value, they are entitled to a certain kind of treatment, namely,
respectful treatment, which acknowledges their independent worth and which forbids their
being used merely for others' purposes. Harms, intentionally done to anyone subject cannot be
justified by aggregating benefits derived by others (Fischer 2021). If we keep an animal not only
as a means to an end, for example as an aesthetic object, then we must be concerned about
the animal's welfare and needs for the animal's sake, i.e. for its own sake (Korsgaard 2018). This
means that we are responsible for ensuring that the animal is well, which excludes accepting
suffering.
What distinguishes us humans is our capacity for empathy, which on the one hand means
putting ourselves in the shoes of other creatures and feeling them out, and on the other hand
enables us to realise that other creatures mean something to themselves in the same way that
we humans mean something to ourselves. What distinguishes us humans is reason, which leads
us to the conclusion that every animal must be regarded as an "end in itself" and whose fate is
significant. In other words, an animal is entitled to be treated according to human moral
standards because they are sentient beings who perceive the world in categories of good or
bad and act accordingly (Korsgaard 2018).
Although there are different moral philosophical points of view, the above has great relevance
in relation to the Enigma form of the leopard gecko. To ignore the individual suffering of
animals with ES because not every Enigma gecko also develops ES, and Enigma causes "such
beautiful colour effects", is to reduce animals to a mere means to an end. In the sense of
Dawkins (2021), no animal wants to be chronically ill and in the sense of Kant's moral
philosophy, it is a disrespectful and therefore morally condemnable act to ignore the inherent
value and autonomy of an individual. Not everything that is possible is good or morally

21

Electronic copy available at: https://ssrn.com/abstract=4626177

acceptable.

4.4 Legal aspects

In Switzerland, the Enigma form has been banned since 2015 because, according to the
Ordinance on Animal Welfare in Breeding, it has the following characteristics, which are judged
as "torture breeding": "Damage that leads to functional failures or behavioural deviations, that
severely impairs the general condition, that leads to severe disturbances of the body functions.
Sufferings which make normal behaviour impossible. Profound interventions in the capabilities
of an animal that lead to severe restrictions in the ability to react to environmental stimuli"
(BLV 2014; original in German).
In Germany, too, there is a report by the Federal Ministry of Food and Agriculture which defines
what is meant by "torture breeding" and prohibits it, citing the Animal Welfare Act.
Furthermore, it should serve as information for the public, "if breeders are not aware of the
necessary correlations and consequences of their actions, do not observe them and exceed the
necessary limits of their creative possibilities (e.g. breeding with defective genes or
overtyping), there is a danger that they will impair the welfare of the animals with their
breeding objectives" (BMEL 2005; original in German). According to the German Animal
Welfare Act, it is forbidden to breed vertebrates that show pain, suffering or damage as a
result of the breeding. It is also not permitted to keep or exhibit them. “Torture breeding” is
furthermore defined as traits which are deliberately promoted or tolerated and which manifest
themselves, among other things, in behavioural disorders or limited self-preservation. In this
context, it is sufficient if pain, suffering or damage is already present to a minor extent (BMEL
2005). So, formally, there are theoretically possibilities to prevent breeding and distribution of
ES leopard geckos. The problem here, however, is that the above-mentioned expert opinion
only refers to the breeding of birds and mammals, and the breeding of reptiles is not regulated
in any detail. And this despite the fact that presumably thousands of these animals are kept in
Germany alone. The fact that the private husbandry of reptiles epitomises a blind spot of legal
and scientific attention is concerning. The 1990s saw an increasing public interest in reptiles -
probably linked to the increasing “options” of the expanding internet. This development led to
a boom from the year 2000 onwards (BNA 2020; Benn et al. 2019; Schuppli et al. 2014; Warwick
2014; Arbuckle 2013; Herrel et al. 2013; Whitham et al. 2013). Nowadays there are millions of
privately kept reptiles in Europe, the USA and parts of Asia (Warwick 2014, 1990).
Fischer (2021) points out an interesting aspect: if the law does not protect what it should
protect, such as freedom from pain, suffering or fear, then this constitutes a morally
problematic lapse.

4.5 Conclusion

As already described, the use of Enigma leopard geckos varies greatly. There are large breeders
who have completely removed this form from their breeding portfolio and explicitly justify this
with the disease potential of this form: “Efforts to remove the disorder [ES] have proven
unsuccessful over and over. Be very careful of breeders claims to no "Enigma syndrome". I
personally won't work with them any more due to the issues” (Geckoboa 2, 2023). Many
others, whether large-scale or hobby breeders, do not do this, perhaps pointing out the risk.

22

Electronic copy available at: https://ssrn.com/abstract=4626177

This should be seen as particularly critical and ethically problematic, as potential carriers of ES
are produced and marketed against better knowledge. Due to the manifold "mixing" of the
most diverse colour forms with Enigma, and the possibly not yet fully understood inheritance,
the harmful genes are potentially distributed throughout the entire gene pool. This is a
condition that should not occur in good and therefore responsible animal husbandry.
In animal husbandry, the animal should be kept for its own sake and not as an object or a mere
means to an end. However, if a leopard gecko is bred and sold only for its external
characteristics, in the case of Enigma for example the red eyes, the intense colouring or the
deviating patterning, although the health risk is known, then this action is ethically
questionable and not desirable. This both puts the health and welfare of the individual animal
at risk and increases the likelihood that this fate will befall more and more animals as harmful
genes accumulate in the population of leopard gecko keepers (Care Guides for pet reptiles
2023).
From the facts presented, it has become clear that ES can severely reduce quality of life and
wellbeing in phases, but that in some cases quality of life can still be better in other phases. The
severity of the symptomatology combined with the numerous unanswered questions as for
example, about the origin, heredity or triggering of ES appears to be sufficient to argue against
further propagation and spread of this "colour form" of the leopard gecko. Even though many
Enigmas do not suffer from ES, it does not seem ethically appropriate to find it acceptable for
individual animals to suffer from ES against their better judgement.
Fischer (2021) points out a further important connection: Morality and law should not be
considered the same thing. Because many people refer to laws to put their actions in a morally
good light and argue that what the law allows cannot be wrong after all. In other words, even
if the Animal Welfare Act does not explicitly prohibit the breeding and marketing of the Enigma
form of the leopard gecko, this definitely does not mean that it is morally right and good.
Because breeding always means inbreeding to a large extent, it is not understandable why we
have to go through the same experiences again and again. It is known from dog breeding that
there are over 700 genetically fixed mutations (Gansloßer et al. 2021) that cause dogs to suffer.
Even if reptiles may be somewhat less susceptible in this respect, examples such as ES,
Iridophoromas in Lemon Frost leopard geckos and other examples show that (in)breeding can
also be associated with adverse consequences in reptiles. Why is it no longer sufficient today to
enjoy the nature and natural appearance of a leopard gecko? And shouldn't those for whom
this is not enough perhaps better refrain from keeping animals?!

23

Electronic copy available at: https://ssrn.com/abstract=4626177

Conflict of interest
The author declare no conflict of interest.

Correspondence: [email protected]
Dr. Frank Krönke, Birkerstr. 12, 80636 München (Munich), Germany

Financing / Funding / Affiliation

This study received no funding and is not affiliated with any institution. It was carried out
exclusively in my spare time, all costs were borne by myself.

24

Electronic copy available at: https://ssrn.com/abstract=4626177

Sources

A) PRINTED

Arbuckle, Kevin 2013:

Folklore husbandry and a philosophical model for the design of captive management regimes.
Herpetological Review 44(3): 448-452.

Baer, Janet 1998:

A veterinanry perspective of potential riskfactors in environmental enrichment. In:
Shepherdson, David J; Mellen, Jill D; Hutchins, Michael “Second Nature. Environmental
enrichment for captive animals”, p. 277-301.

Barber, Josph C.E. 2009:

Programmatic approaches to assessing and improving animal welfare in zoos and aquariums.
Zoo Biology 28(6):519-530.

Benn, Amelia L; McLelland, David L; Whittaker, Alexandra L 2019:

A review of welfare assessmenr methods in reptiles, and preliminary application of the welfare
Quality® Protocol to the pygmy blue-tongue skink, Tiliqua adelaidensis, using animal based
measures. Animals 9(1): 1-27.

Bleckett, T.A.; McKenna, C.; Kavanagk, L.; Morgan, D.R. 2017:

The welfare of wild animals in zoological institutions: are we meeting our duty of care?
International Zoo Yearbook 51: 187-202

Boissy, Alain; Manteufel, Gerhard; Jensen, Margit; Moe, Randi Oppermann; Spruijt, Berry 2007:
Assessment of positive emotions in animals to improve their welfare. Physiology and Behaviour
92(3): 375-397.

Clark, Fay 2017:

Cognitive enrichment and welfare: current approaches and future directions. Animal Behaviour
and Cognition 4(1): 52-71.

Dawkins, Marian Stamp 2021:

The science of animal welfare: understanding what animals want. Oxford University Press.
Oxford, UK.

Fischer, Bob 2021:

Animal Ethics. A contemporary introduction. Routledge Contemporary Introductions to
Philosophy. Routledge.

Gansloßer, Udo, Adler, Yvonne, Braun, Gudrun 2021:

Hunderassen. Zoologie, Zucht und Verhalten neu betrachtet. Franckh-Kosmos-Verlags-GmbH.
Stuttgart. 608 p

Grießhammer, Karsten; Köhler, Gunther 2009:

25

Electronic copy available at: https://ssrn.com/abstract=4626177

Leopardgeckos. Pflege, Zucht, Erkrankungen, Farbvarianten. Herpeton Verlag, Offenbach.

Guo, Longhua, Bloom, Joshua; Sykes, Steve; Huang, Elaine; Kashif, Zain; Pham, Elise; Ho,
Katarina; Alcaranz, Ana; Xiao, Xinshu Grace; Duarte-Vogel, Sandra; Kruglyak, Leonid 2021:
Genetics of white color and iridophoroma in “Lemon Frost” leopard geckos. PloS Genet 17(6):
e1009580; doi.org/10.1371/journal.pgen.1009580.

Henkel, Friedrich Wilhelm; Köthing, Michael; Schmidt, Wolfgang 2000:

Leopardgeckos. Natur und Tier Verlag, Münster.

Herrel, Antony; van der Meijden, Arie 2014:

An analysis of the live reptile and amphibian trade in the USA compared to the global trade in
endangered species. The Herpetological Journal 24(2):103-110.

Jones, Narelle; Sherwin, Sally L; Robbins, Rachel; McLelland, David J; Whittaker, Alexandra L
2021:
Welfare assessment tools in zoos: from theory to practice. Veterinary Science 9(4).
doi:10.20944/preprints202112.0027.v2

Keeling, Linda J; Rushen, Jeffrey; Duncan, Ian JH 2018:

Understanding animal welfare. In: Appleby, Michael C; Olsson, Anna S; Galindo, Francisco
“Animal welfare”. CABI, Oxfordshire, UK.

Korsgaard, Christine M. 2018:

Fellow creatures. Our obligations to the other animals. Oxford University Press. Oxford, UK.

Krönke, Frank 2020:

Leopardgeckos (Gattung Eublepharis) - Top angesagt oder todlangweilig? Ein Blick auf die
Terraristik - und darüber hinaus. (Cover story) REPTILIA 25(6): 12-25.

Krönke, Frank; Xu, Lisa 2023 (submitted & accepted)

Sensual stimulation as a means of sustained enhancement of well-being in leopard geckos,
Eublepharis macularius. Animals 2023.

Lambert, Helen; Carder, Gemma; D'Cruze, Neil 2019:

Given the cold shoulder: a review of scientific literature for evidence of reptile sentience.
Animals 2019, 9(19): 821; doi.org/10.3390/ani9100821

Libourel, Paul-Antoine; Herrel, Anthony 2016:

Sleep in amphibians and reptiles: a review and a preliminary analysis of evolutionary patterns.
Biological Reviews 91(3): 833-866.

Mellor, David 2013:

Setting the scene. When coping is not enough: promoting positive welfare states in animals.
RSPCA Australia. Scientific Seminar 2013. Proceedings, p. 2-13.

Mohanty,, Nitya; Wagener, Carla; Herrel, Anthony; Thaker, Maria 2022:

26

Electronic copy available at: https://ssrn.com/abstract=4626177

The ecology of sleep in non-avian reptiles. Biological Reviews 97(2): 505-526.

Rose, Paul E.; Nash, Steve M.; Riley, Lisa M. 2017:

To pace or not to pace? A review of what abnormal repetitive behaviour tells us about zoo
animal management. Journal of Veterinary Behaviour 20: 11-21.
https://doi.org/10.1016/j.jveb.2017.02.007

Szydłowski, Pawel; Madej, Jan Pawel; Duda, Magdalena; Madej, Januz A; Sikorska-Kopyłowicz,
Agnieszka; Chemłmońska-Soyta, Anna; Ilnicka, Lucyna; Duda, Przemysław 2020:
Iridophoroma associated with the Lemon Frost coulor morph of the leopard gecko
(Eublepharis macularius). Scientific Reports 10:5734, doi.org/10.1038/s41598-020-62828-9.

Tremper, Ron 2020:

Leopard Geckos. The next generations. Self-published, 2nd Ed., 199 pp.

Warwick, Clifford 1990:

Reptilian ethology in captivity: observations of some problems and an evaluation of their
aetiology. Applied Animal Behaviour Science 26: 1-13.

Warwick, Clifford 2014:

The morality of the reptile pet trade. Journal of Animal Ethics 4(1): 74-94.

Whitham, Jessica; Wielebnowski, Nadja 2013:

New directions for zoo animal welfare science. Applied Animal Behaviour Science 147(3-4): 247-
260)

Young, Robert J. 2003:

Environmental Enrichment for captive animals. Universities Federation for Animal Welfare
(UFAW). Blackwell Publishing, 228p.

All internet sources considered between November 2022 and February 2023.

B) DIGITAL

B.1) HOMEPAGES

Bargen, Ben 2013 (Geckotime.com):

https://geckotime.com/enigma-syndrome/

Bundesamt für Lebensmittelsicherheit und Veterinärwesen (BLV) (Swizerland) 2014:

Schweizerische Eidgenossenschaft. Fedlex Die Publikationsplattform des Bundesrechts:
455.102.4 Verordnung des BLV über den Tierschutz beim Züchten.
https://www.fedlex.admin.ch/eli/cc/2014/747/de

27

Electronic copy available at: https://ssrn.com/abstract=4626177

Bundesministerium für Ernährung und Landwirtschaft (BMEL) (Germany) 2005:
Gutachten zur Auslegung von Paragraf 11b des Tierschutzgesetzes. Die Sachverständigen-
gruppe Tierschutz und Heimtierzucht diskutiert das Verbot von Qualzüchtungen.
https://www.bmel.de/DE/themen/tiere/tierschutz/gutachten-paragraf11b.html

Bundesministerium für Ernährung und Landwirtschaft (BMEL) (Germany) 1997:

Haltung von Reptilien. Gutachten der Sachverständigengruppe über die Mindestanforderungen
an die Haltung von Reptilien (10. Januar 1997).
https://www.bmel.de/DE/themen/tiere/tierschutz/haltung-reptilien.html

Bundesverband für Fachgerechten Natur-, Tier- und Artenschutz 2020:

Reptilienhandel in Deutschland. Ergänzende Daten zum BfN Skript 545. https://www.bna-
ev.de/downloads/f_e_publikation/2020_12_14_BNA-
Publikation_Reptilienhandel_in_Deutschland.pdf

Care Guides for Pet Lizards:

https://www.lizards101.com/enigma-syndrome-in-a-leopard-gecko/

Der Leopardgecko.de:
https://www.der-leopardgecko.de/haltung-und-mehr/farbformen-von-a-z/enigma-combos/

Geckoboa.com (John Scarbrough):

1 https://www.geckoboa.com/available-leopard-geckos.html
2 https://www.geckoboa.com/leopard-gecko-genetics.html
3 Damuth, Dillon (without year): Leopard Geckos: A hobby developing to an artistic scene.
Www.geckoboa.com

Geckosetc.com (Steve Sykes):

https://geckosetc.com/leopard-geckos/

Imperial Reptiles and Exotics:

https://imperialreptiles.com/products/enigma-leopard-gecko-for-sale

Leopard Gecko Wiki:

http://www.leopardgeckowiki.com/index.php/Enigma

Ledopardgecko. Com (Rom Tremper)

http://leopardgecko.com/

pogona.ch
https://testrecht.pogona.ch/9.1_Qualzucht

Reptilefiles:
https://reptifiles.com/leopard-gecko-care/leopard-gecko-diseases-health/leopard-gecko-
enigma-syndrome/

28

Electronic copy available at: https://ssrn.com/abstract=4626177

Schuppli, Catherine; Fraser, David; Bacon, H. 2014:
Welfare of non-traditional pets. In: Wild and exotic animals as pets collection.
https://www.wellbeingintlstudiesrepository.org/wileapet/2

The Urban Reptile (Craig and Lori Stewart):

https://theurbanreptile.com/511-2/leopard-geckos-for-sale/

The Reptile Guide:

thereptileguide.com

B.2) FORUMS, SELLING PALTFORMS

All About Leopard Geckos (Facebook)

https://www.facebook.com/groups/435984710284620/search/?q=enigma%20syndrome

Enigma Syndrome & neurological issues in Leopard geckos (Facebook)

https://www.facebook.com/groups/1556752061339811

Faszination Leopardgecko (Facebook)

https://www.facebook.com/groups/1294006437475039/search/?q=enigma

Leopard Gecko Life (Facebook)

https://www.facebook.com/groups/2054010221490441/search/?q=enigma%20syndrome

Morphmarket
https://www.morphmarket.com/

Quoka:
https://www.quoka.de/tiermarkt/reptilien-terraristik/kleinanzeigen.html

B.3) VIDEOS

Denby Charl:
https://www.youtube.com/watch?v=EKP1x1rrWCM

Forsaken December:
Enigma leopard gecko with balance disorder. https://www.youtube.com/watch?
v=7nvMUq6HBp8

GeckoLand:
Enigma syndrome in Leopard geckos | Neurological disorders. https://www.youtube.com

Happy Gecko Farm:

https://www.youtube.com/watch?v=jxQSuuS8kO8

29

Electronic copy available at: https://ssrn.com/abstract=4626177

Ice Cream:
Enigma Syndrome 101 Leopard Gecko Basics. https://www.youtube.com/watch?v=fcaWfO906lE

Jessica's Animal Friends:

1 What is Enigma Syndrome? https://www.youtube.com/watch?v=ylyif01srlg
2 Enigma Syndrome in Leopard Geckos. Everything You Need to Know.
https://www.youtube.com/watch?v=Cuin3N43Sqc
3 Death Rolling. The Worst Symptom of Enigma Syndrome. https://www.youtube.com/watch?
v=E-5KNJMhHww
4 Neurological Disorders in Leopard Geckos. Symptoms, Causes & Treatment.
https://www.youtube.com›watch?v=R1lFeb_RpJE
5 White and Yellow Syndrome in Leopard Geckos. https://www.youtube.com/watch?
v=kC8M3ZE02EA
6 Problematic Leopard Gecko Morphs. https://www.youtube.com/watch?v=w_0nKnSnJ8g
7 Another Leopard Gecko with Enigma Syndrome. Meet Ros.
https://www.youtube.com/watch?v=5AfNMhpALrs

SaSobek Reptiles:
A little about Enigmas. https://www.youtube.com/watch?v=s8skyxXZmgM

Wildfire Geckos:
Leopardgecko Enigma Syndrome. https://www.youtube.com/watch?v=mw9LYsweiE8

30

Electronic copy available at: https://ssrn.com/abstract=4626177