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Preface
1
MERTENSIELLA
Studies on Anomalies
in Natural Populations of Amphibians
Untersuchungen zu Anomalien
in natürlichen Populationen von Amphibien
Herausgegeben von
K H & A D
im Autrag der
Deutschen Gesellscha für Herpetologie und Terrarienkunde e.V.
Mertensiella
Supplement zu Salamandra
Mannheim, . August
9
Terminology and glossary
Commented glossary, terminology and synonymies of anomalies in
natural populations of amphibians
K H, A D & V V,
UFZ – Helmholtz Centre for Environmental Research,
Department of Conservation Biology, Permoserstr. , Leipzig, Germany; email: klaus.henle@ufz.de
Institut de Systématique, Évolution, Biodiversité ISYEB – UMR – CNRS, MNHN, UPMC, EPHE Muséum natio-
nal d’Histoire naturelle, Sorbonne Universités, rue Cuvier, CP , , Paris, France; email: adubois@mnhn.fr
Institute of Plant and Animal Ecology, Uralian Division, Russian Academy of Science,
Marta St , , Ekaterinburg, Russia; email: vol_de_mar@list.ru
Ural Federal University, Ekaterinburg, Mira St , , Russia
Abstract. To allow comparison of observations an unequivocal denition of terms is essential. While early scholars
of amphibian anomalies already introduced specic terms and provided extensive denitions, some of these terms
have changed their meaning over time, some are no longer used and others have been replaced more recently with al-
ternative terms by some but not all authors. Most of the more recently published glossaries are rather narrow in their
coverage, addressing primarily a limited set of skeletal anomalies, few provide synonyms and none of them discuss
inconsistencies among dierent terminologies. To facilitate an understanding of the contributions in this volume and
to facilitate comparison of publications in general, we provide an extensive glossary and synonymy of the terms used
to describe external anomalies of amphibians. is includes colour and pattern anomalies, morphological anomalies,
anomalies related to edema or tumours and anomalies of embryos. We discuss the inconsistencies among dierent
terminologies and make recommendations for a future standardized use of terms. Our most important recommen-
dation is to explicitly refer to a particular terminology and to describe the anomaly suciently to enable readers to
understand what has been observed even if they are accustomed to using another terminology.
Keywords. Amphibia, anomalies, colour anomalies, skeletal anomalies, synonymies, terminology
Zusammenfassung. Eine eindeutige Denition von Begrien ist für den Vergleich von Beobachtungen essentiell.
Zwar haben bereits die ersten Forscher, die sich mit Anomalien in natürlichen Populationen von Amphibien be-
schäigt haben, Begrie zur Dierenzierung verschiedener Typen von Anomalien eingeführt und dafür ausführliche
Denitionen gegeben, jedoch haben einige Begrie im Laufe der Zeit ihre Bedeutung geändert, andere elen in Ver-
gessenheit und wieder andere wurden von manchen, jedoch nicht allen neueren Autoren durch neue Begrie ersetzt.
Die meisten neueren publizierten Glossare sind im Umfang relativ beschränkt und beziehen sich vorwiegend auf eine
kleine Auswahl von Skelettanomalien. Die meisten von ihnen gehen weder auf Synonyme noch auf unterschiedliche
Denitionen von Begrien durch verschiedene Autoren ein. Um das Verständnis der folgenden Beiträge in diesem
Band und den Vergleich unterschiedlicher Publikationen zu erleichtern, stellen wir hier ein umfangreiches Glossar
inklusive Synonyme zusammen. Dies umfasst Farb- und Musteranomalien, morphologische Anomalien, ödem- so-
wie tumorähnliche Anomalien und Anomalien von Embryonen. Wir diskutieren abweichende Verwendungen von
Begrien durch verschiedene Autoren und machen Vorschläge für eine konsistente Begrisverwendung. Die wich-
tigste Empfehlung besteht darin, sich eindeutig auf eine bestimmte Terminologie zu beziehen und Beobachtungen
ausführlich genug zu beschreiben, dass Leser die Art der Anomalie eindeutig identizieren können, auch wenn sie
eine andere Terminologie gewöhnt sind.
Schlagwörter. Amphibia, Anomalien, Farbanomalien, Skelettanomalien, Synonyme, Terminologie
MERTENSIELLA | – |
10
Henle, Dubois & Vershinin
Introduction
Natural populations of amphibians show a large diver-
sity of deviations from the normal range of variation
(H et al. a). Various terms have been used for
such deviations. For deviations in morphology, for ex-
ample, the terms malformation, deformity, abnormality
or anomaly are used frequently. J et al. ()
suggested dening malformation as a “permanent struc-
tural defect resulting from abnormal development”, de-
formity as an “alteration in an organ or structure that
originally formed correctly” and abnormality as “any
gross deviation from the normal range in morphologi-
cal variation”. ese denitions were adopted by the US
Fish & Wildlife Service (USFWS ) for the standard
operating procedure of abnormal amphibian surveys in
the USA.
Malformations and deformities as dened here are
oen dicult to distinguish without a detailed study of
the case. For example, the absence of a digit (ectrodac-
tyly) could either result from perturbations of ontogen-
esis (e.g. M & S , D et al. )
or from amputation aer a normal ontogenesis (e.g.
D & D , M et al. ).
Some authors (e.g. MC & T ) do not
include injuries when reporting on abnormal amphibi-
ans. However, unless a wound is still open or bleeding,
it is oen very dicult to separate injuries from other
anomalies, and in the eld this is usually impossible
(H et al. a). erefore, we will consider both
kinds of abnormal phenotypes under the general term
“anomaly”, a practice that is followed by many authors.
We also include all non-morphological deviations from
the normal phenotype in this term.
e term “anomaly” is a relative one that has to do
with the taxonomic status of specimens. What is an
anomaly in a species or in a genus may not be so in an-
other one. us, “albinism” is not an anomaly in Proteus
anguinus or in other cavernicolous amphibians. White
eggs are normal in species that deposit their eggs in
places that are not exposed to sunlight, such as fast-run-
ning water, concealed sites on the ground or in vegeta-
tion above water (D & T ), but are an
anomaly in species in which eggs are exposed to sun-
light, such as most European species, the eggs of which
are deposited in water (e.g. R , B
). Likewise, black-eyed individuals are rare muta-
tions in some species (H et al. a), whereas they
are the normal condition in others (G & V
). In Plethodon cinereus, a red colour morph is fre-
quent in some regions but very rare in others (L
& S ) and thus might only classify as an anom-
aly in parts of the range.
Even gross morphological anomalies in a given spe-
cies may be the normal condition in another one: thus,
toads of the genus Brachycephalus have only two or three
ngers and three or four toes, a condition, which would
qualify as ectrodactyly in most other anuran genera. In
some salamanders, toe morphology is highly variable
and it is dicult to set the limit between the range of nor-
mal and abnormal variation, e.g. in Batrachuperus pin-
chonii (C & B ), Ambystoma maculatum
(W ) and Salamandrella keyserlingii
(B , V ). is variation may or
may not be described by one of the terms included in
the glossary below. us, the use of these terms does not
necessarily imply an anomaly. However, we recommend
using these terms only if such an implication is intended.
In any case, one needs to be precise about whether one
regards a particular phenotype as normal or abnormal.
It is impossible to provide a list of all possible anom-
alies in amphibians, as virtually all characters, in eggs,
embryos (see overview by B et al. ), larvae
and adults may be concerned. Some of these anomalies
are very spectacular, such as polymely, and have attract-
ed more attention than others (O ).
Early scholars of amphibian anomalies already pro-
vided extensive denitions of anomalies (e.g. G
S-H ) and several more recent glossa-
ries for amphibian anomalies also exist (e.g. D,
, T , M , M et al. ,
O , G & H , J-
et al. , , V , , S
, USFWS , L , , N
). Most of them address and are (almost) limited
to selected skeletal anomalies. e only relative compre-
hensive osteopathological glossary for amphibians and
reptiles was published recently by R et al.
(). D () provided denitions for terms
related to albinism and G & H ()
for terms related to tumours.
Unfortunately, the terminology of amphibian anom-
alies is not always consistent among the dierent glossa-
ries and among dierent scientists (B et al. ).
Moreover, terminology has changed over time; some
terms are no longer used, some have changed their
meaning and others have been replaced with new terms
by some but not all authors. Only B et al. ()
and R et al. () provided some synony-
mies and only the former authors discussed deviating
denitions used by dierent authors. erefore, it is not
rare that the type of anomaly reported remains unclear
even when a specic term was used.
To facilitate an understanding of the remaining pub-
lications in this issue, and to help identify the type of
anomalies reported in the literature, we provide a com-
mented glossary containing synonyms and explaining
the dierent denitions given to a particular term. We
also make recommendations towards future standard-
ization of the use of terms. Our recommendations are
guided by the following criteria: e recommended
terms should reduce as much as possible any existing
inconsistency and confusion. e terminology must be
applicable in the eld as in environmental monitoring
it is not feasible to collect hundreds of individuals for
detailed analyses in the lab. As a third criterion, with less
weighing, we strived for an internally and etymological-
ly consistent terminology.
11
Terminology and glossary
Fig. a: Albino-like white diurnal colouration of Chiromantis
xerampelina is attained physiologically and thus not called al-
binism. Road South of Lower Sabie, Kruger NP, South Africa.
Source: Wikipedia Commons. Photo: B. D.
We are aware that not everybody may agree with all of
our recommendations. In any case, we strongly urge sci-
entists to explicitly state which denition they followed
and to describe the anomaly in sucient details that
readers are able to follow what they were referring to.
An illustration of the observed anomalies using photo-
graphs should be considered, especially if they do not t
any of the given denitions very well. Anomalies may
be rather complex, e.g. supernumerary limbs that are
incompletely developed. For such cases we recommend
only using the term applicable for the primary anomaly,
in this instance polymely, and not to additionally call it
ectromely. If one believes it to be relevant, one may rath-
er provide a detailed description of the supernumerary
limb.
Terms, denitions and synonyms
. Anomalies of colouration and pattern
Terminology in this domain either follows external ap-
pearance or refer to the cells or pigments that were ab-
sent or modied. In most publications the presumed
modied or absent chromatophore type was not veried
by histological examination. However, inference from
external colouration is not always a reliable indicator of
the absence of a particular chromatophore type. ere-
fore, we recommend using a terminology based on the
absence of pigments or chromatophore types only if his-
tological examinations were made. Otherwise descrip-
tive terms and statements, such as “black eyes”, “blackish
frogs”, “blue frogs”, “erythristic”, “avistic”, “golden frogs”,
or “green frogs”, are preferable, ideally in combination
with colour photographs (compare N ).
D () provided a short glossary of terms
related to albinism. R & N () classied
dark variants in amphibians according to the presence
or absence of chromatophore types on dierent parts of
the body.
Colouration and pattern and thus deviations from
the normal phenotype are usually species-specic. Here
we limit our glossary to types of colour and pattern
anomalies that are not species-specic.
Albinism | Partial or complete absence of integumen-
tary pigmentation giving an individual a whitish, yel-
lowish to golden or pinkish to reddish appearance (Figs.
b, , , , , , & b). Individuals with transparent
skin are included, if the whole body is aected and if
a golden, yellowish, reddish or whitish tone is present;
however, cases in which only parts of the body show
abnormal ➝ translucent skin are never included de-
spite partially lacking integumentary pigmentation.
Likewise, individuals in which red colour replaces the
normal yellow colour and blue frogs are generally not
included in the term albinism. Sometimes albinism is
used in a more restrictive sense as a synonym of ➝
leucism, but most frequently it is either unspecied or
used in the broader sense given here. Since albinism has
been used in such a variable way, we recommend us-
ing a more specic term whenever possible. Terms for
specic types of albinism are based either on external
appearance [➝ albinistic pinto, ➝ complete albino, ➝
erythrism (partim), ➝ avism, ➝ leucism] or in rela-
tion to the presence/absence of pigments or chromato-
phores [➝ amelanism, ➝ axanthism (partim), ➝ hy-
pomelanism (partim), ➝ hypopigmentation (partim)].
e terms of these two types of classications cannot be
fully matched because the absence of a particular chro-
matophore type will lead to dierent external appear-
ance depending on the normal colouration of the spe-
cies involved. erefore, we strongly recommend using
the terminology based on appearance unless histologi-
cal examinations were made. Further subcategories of
albinism are ➝ partial albinism (only parts of the body
aected) and ➝ transient albinism (individuals increas-
ingly acquire pigmentation during development). Some
species can change physiologically to an albino-like
white colouration but are not called albino (Fig. a)
Albinistic pinto | Normal pigmentation only present as
small scattered spots (Fig. b); a subcategory of ➝ albi-
Fig. b: Two large albinistic and a normal tadpole of Bufotes
viridis; albinistic individuals may be dicult to classify: indi-
viduals, such as the one on the le, are oen called leucistic
despite of them still having a few slightly pigmented spots;
the middle one is closer to albinistic pinto with a few normal
coloured dark spots and normal coloured hindlegs; Roßwag,
Germany, September . Photo: K. R.
12
Henle, Dubois & Vershinin
nism. e term is not oen used in the herpetological
literature; instead, pinto individuals are oen described
under the more general term ➝ albinism or, when nor-
mal pigmentation covers larger spots, ➝ partial albi-
nism
Albino | Usually a synonym of ➝ true albino but some-
times additionally includes ➝ leucism
Amelanism | Melanin or melanophores absent (a sub-
category of ➝ albinism). If absence is not veried histo-
logically, individuals in which melanin or melanophores
are reduced but not completely absent may be called
amelanistic, but correctly speaking such cases fall into
the category ➝ hypomelanism. Sometimes amelanism
is used as a synonym of ➝ avism (e.g. W et
al. ) but this should be avoided because the absence
of melanin does not necessarily result in yellowish in-
dividuals
Anomaly N | ➝ Black eyed (Fig. )
Axanthism | Xanthophores or carotenoid vesicles in
xanthophores absent. If the normal phenotype is green
such individuals appear blue and are usually called ➝
blue (frogs). In species with another normal coloura-
tion, it may be a subcategory of ➝ albinism or may re-
sult in a species-specic abnormal colour pattern that is
not reminiscent of albinism
Black eyed | Iris black coloured (Fig. ); in some spe-
cies, such as various dendrobatids (Fig. ) and Brachy-
cephalus ephippium (Fig. ), black eye colouration is
the normal phenotype (G & V ). Unilat-
eral black-eyedness is a subcategory of ➝ heterochro-
mia
Blue frogs | Frogs with blue colouration (Fig. ). e
anomaly is due to a reduction or lack of xanthophores
or their yellow pigments in species in which green is the
normal phenotype. us, blue frogs are a special type of
➝ axanthism but usually not included in the term ➝
albinism. Notwithstanding, the term axanthism is sel-
dom applied to blue frogs. In a few frog species, such as
Aplastodiscus arildae (Fig. ), blue is part of the normal
phenotype and male Rana arvalis (Fig. a) may turn
blue during breeding
Complete albino | A subcategory of ➝ albinism, in
which all integumentary pigmentation is lacking (Fig.
), including in the eyes (eyes that completely lack pig-
mentation may appear red due to blood vessels or may
be vestigial; the body may appear pinkish due to blood
vessels). Sometimes, the term is also used for individu-
als that lack all integumentary pigmentation but whose
eyes have the normal colour. We recommend using the
term ➝ leucism for such cases and urge to use photos
or descriptions of the eye colour to avoid confusion. A
few cavernicolous species, notably Proteus anguinus an-
guinus (Fig. ), are complete albinos in the normal phe-
notype
Depigmentation | Normal colouration of the body, or
parts thereof, lost, usually due to exposure to chemicals
or disease
Depigmentation of beaks and/or denticles | Tadpoles
lacking pigmentation of the beak and/or denticles of the
tooth rows, which are usually dark brown to black due
to keratinization. Depigmentation may be due to pol-
lution or infection by Batrachochytrium dendrobatides,
i.e., an anomaly, but also due to cold temperature, i.e.,
not an anomaly (reviewed by H et al. a)
Erythema | Reddening of the skin due to increased
blood ow
Fig. 2: Adult Bufo bufo with a black eye (le), due to a reces-
sive mutation causing absence of iridophores as demonstrated
by a crossing experiment, and normal phenotype (right); Car-
nelle Forest near Paris, France; . Photo: A. D.
Fig. : Blue as part of the normal colour pattern is rare in
amphibians; in male Aplastodiscus arildae it is combined
with transparency on hidden surfaces; Teresopolis, Brazil,
... Photo: K. H.
13
Terminology and glossary
Erythrism | Red individuals. ere are two types of ab-
normally red individuals: ) xanthophores are replaced
by erythrophores but the remaining pigmentation is
normal (Fig. ); ) integumentary pigmentation is ab-
sent except for erythrophores. e latter case but not the
former is included in the term ➝ albinism; D
Fig. b: Rana arvalis, normal phenotype outside the breeding season; Tschekalin, Russia, ... Photo: K. H.
Fig. a: Rana arvalis, blue colour is the normal phenotype for males in some regions during the breeding season; Elbe ood-
plain, Dessau, ... Photo: A. K.
14
Henle, Dubois & Vershinin
Fig. b: Pelophylax esculentus normal phenotype; Tschekalin, Oka, Russia, July . Photo: K. H.
Fig. a: Blue, an abnormal phenotype of Pelophylax esculentus; Braunschweig, Germany. Photo: A. M.
15
Terminology and glossary
() called this category partial albino with erythro-
phores, but this terminology is rarely used. Some spe-
cies have a red colour as the normal phenotype (Fig.
); these are usually not called erythristic, with the red
e of Notophthalmus viridescens (Fig. ) and the red
morph of Plethodon cinereus being exceptions
Fig. : Normal phenotype and complete albino of Xenopus
laevis. Source: Wikipedia Commons. Photo: Nashville Zoo.
Fig. a: Salamandra salamandra: normal colouration of an adult;
Maourine pond, Toulouse, France, ... Source: Wiki-
pedia Commons. Photo: D. D.
Fig. : Oophaga pumilio, a species in which red colour and
black eyes are the normal phenotypes; Zoological Garden
Wilhelma, Stuttgart. Photo: K. H.
Fig. : e normal phenotype of Proteus a. anguinus is com-
plete albino; Baredine, Croatia, ... Photo: B. T.
Fig. b: Salamandra salamandra: erythristic adult, captive bred
individual; Germany. Photo: B. T.
Fig. : In immature Notophthalmus viridescens red is the nor-
mal phenotype; North Fork Mountain, USA. Source: Wikipe-
dia Commons. Photo: J. Q.
16
Henle, Dubois & Vershinin
Flavism | A subcategory of ➝ albinism that covers yel-
lowish or golden-to-yellowish individuals (integumen-
tary pigmentation absent except for xanthophores) (Fig.
& b). D () referred to this category as
partial albino with xanthophores, but this terminology
is rarely used. W et al. () used ➝ amelanism
as a synonym of avism but this should be avoided as
amelanism does not necessarily result in yellowish in-
Fig. a: Pelophylax esculentus; normal phenotype; adult individual; Oka, Tschekalin, Russia, August . Photo: K. H.
Fig. b: Pelophylax esculentus; avistic individual; Ballertasche, Germany. Photo: M. V.
17
Terminology and glossary
dividuals. Yellow or yellow-to-orange colouration of the
body as normal phenotype appears in aposematically-
coloured frogs (Fig. ) and the males of some frog
species, e.g. in the genus Dendropsophus (Fig. ), turn
completely yellow during the breeding season; usually,
they are not called avistic
Golden morphs | A subcategory of ➝ avism in which
individuals appear golden (Fig. b)
Hemorrhage | Reddening due to bleeding (e.g. M
et al. )
Heterochromia | e two eyes dier in their pigmenta-
tion (Fig. ); sometimes used for unilateral ➝ black-
eyed individuals. As many dierent types of heterochro-
mia exist, we recommend stating explicitly whether one
eye was black; otherwise it is not possible to assign them
to the black-eyed type of anomaly
Hyperpigmentation | Pigmentation (of body parts) in-
creased in intensity compared to the normal phenotype;
usually applied to cases that result from an increase in mel-
anophores or melanin expansion; R & N ()
called such individuals dark variants; if individuals are very
dark but not completely black, they are called ➝ melanoid
Fig. : Brachycephalus ephippium, a species in which black
eyes and a yellowish-orange colour belong to the normal
phenotype; Serra dos Orgãos National Park, Brazil, ...
Photo: K. H.
Fig. : Dendropsophus minutus, a species in which males turn yellow during the breeding season; near Ilhéus, Bahia state, Bra-
zil, ... Photo: A. K.
Fig. : Heterochromia in an adult female Bufo bufo; Glubo-
chenskoye pond, Ural, Russia, June . Photo: V. V.
18
Henle, Dubois & Vershinin
Hypomelanism | Melanin or melanophores substan-
tially reduced but present (Fig. ) or melanophores
permanently contracted; if the reduction is substantial,
individuals may be called ➝ albinistic
Hypopigmentation | Pigmentation (of body parts) re-
duced compared to normal individuals; if reduction is
substantial, individuals are oen called ➝ albinistic
Incomplete albinism | Occasionally used instead of ➝
leucism (e.g. S & I ) but this should
be avoided since all forms of albinism, except for ➝
complete albinism, are incomplete albinism
Leucism | (Almost) all integumentary pigmentation ab-
sent but eyes pigmented (Fig. & ); in the literature
sometimes confused with ➝ avism
Fig. : Hypomelanistic adult male Triturus cristatus. ose
body parts that are normally black show a faint grey pattern;
Nordstemmen, Germany. Source: M (). Photo: S.
M.
Fig. : Leucistic Epidalea calamita, ospring from a leucistic male, Germany. Photo: B. T.
19
Terminology and glossary
Melanism | A recent replacement term for ➝ nigrin-
ism; body or parts thereof completely black (Fig. );
oen used more broadly to include ➝ melanoid indi-
viduals and ➝ hyperpigmentation. If only parts of the
body are aected, oen called partial ➝ melanism. A
few species are completely black as normal phenotype,
e.g. Salamandra atra (Fig. ), and some, such as Sala-
mandrella keyserlingii (H et al. a), may change
colour physiologically and become completely black
Melanization | Individuals gain darker pigmentation
over time from melanin synthesis, melanin expansion
or an increase in melanophore numbers. Phenotypi-
cally, individuals may either become darker on larger
parts of the body or dark spots may increase in size
(V , ); may or may not be an abnor-
mal condition
Melanoid | Very dark but not completely black individ-
uals; a few species, e.g. Salamandrella keyserlingii, may
change colour physiologically and become almost black
(Fig. b)
Mid-dorsal stripe absent | Abnormal in species, in
which the normal pattern generally includes a mid-dor-
sal stripe
Mid-dorsal stripe abnormal | Mid-dorsal stripe bro-
ken, bent or irregular in shape (Fig. )
Mid-dorsal stripe present | Abnormal in species in
which a mid-dorsal stripe does not belong to the normal
range of variation
Nigrinism | Old term for ➝ melanism (e.g. K-
)
Novel colours | Parts of the body show colours that do
not belong to the normal range of variation (e.g. yellow
patches in Bufotes viridis: H et al. a)
Partial albinism | e term is used inconsistently; most
commonly, it is used for individuals, in which integu-
mentary pigmentation is only absent on parts of the
body (Fig. ) but some authors also applied the term
to individuals, in which melanophores are absent or
strongly reduced but which have iridophores (then a
synonym of ➝ amelanism respectively ➝ hypomela-
nism); others apply it to ➝ avistic individuals and
K & H () used the term instead
of ➝ transient albinism. To reduce ambiguity and con-
fusion, we recommend restricting the term to the de-
nition given by us and using other subcategories of ➝
albinism for other types of reduced pigmentation; this
also makes it consistent with the use of the term ➝ par-
tial melanism
Partial black | ➝ Partial melanism
Partial melanism | Individuals in which parts of the
body but not the entire body are completely black; the
remaining body shows normal colouration; a subcat-
egory of ➝ melanism
Pattern lacking | ➝ Unicolor
Periodic albinism | ➝ Transient albinism
Semi-albino | Rarely used; if used, then either as a syno-
nym of ➝ avism (e.g. C ) or ➝ leucism
(e.g. S , T ); we discourage
the use of this term because this inconsistency may cre-
ate confusion and established widely used terms are
available for these two types of colour anomaly
Transient albinism | An anomaly in which the eggs are
white but the tadpoles increasingly acquire pigmenta-
tion aer hatching; in rare cases, pigmentation does
not appear before metamorphosis; the light coloured
tadpoles in early stages are called depigmented by some
authors (V ) but ➝ depigmentation is usu-
ally applied to cases in which individuals initially were
pigmented and later lost pigmentation. K &
Fig. : Melanistic Salamandra salamandra almanzoris; born
in captivity. Photo: U. S.
Fig. : Black is the normal phenotype in Salamandra atra;
adult near Bichlbach, Austria, ... Photo: A. G.
20
Henle, Dubois & Vershinin
H () called transient albinism ➝ partial
albinism, which is inconsistent with the usual denition
of the latter term. Many amphibians that deposit their
eggs at concealed sites have white eggs but the larvae or
froglets that hatch from them are pigmented; these cases
are usually not called transient albinism
Translucent | Pigmentation lacking and skin transpar-
ent (Figs. , b & ) (N ). In some spe-
cies, notably in the family Centrolenidae, parts of the
skin are translucent as the normal phenotype (Fig. )
Transparent | ➝ Translucent
True albino | ➝ Complete albino
Unicolor | Individuals that lack the dark or light pattern
typical for the species, e.g. uniform green backs in the
hylid Acris crepitans (G )
Xanthism | Yellowish individuals in which melanin is
strongly reduced or absent (i.e., a synonym of ➝ av-
ism, e.g. P , W et al. )
. Morphological anomalies
Several glossaries of skeletal anomalies have been pub-
lished (e.g. G S-H , T ,
M , M et al. , O ,
Fig. 20: Middorsal stripe abnormally bent, subadult Pelophy-
lax lessonae; Tschekalin, Russia, 9.9.2002. Photo: K. H.
Fig. 19b: Colour variation in Salamandrella keyserlingii: al-
most black subadult taken in a very dark and cold place. is
specimen looks like an abnormal melanoid individual but
changed to normal colouration in a warmer environment.
It also shows oligodactyly on the right hand; Ekaterinburg,
Russia, ... Photo: V. V.
Fig. a: Colour variation in Salamandrella keyserlingii: normal phenotype; Nishni Angarsk, Lake Baikal, Russia, ...
Photo: K. H.
21
Terminology and glossary
J et al. , , S , USFWS
, L , , N , R-
et al. ). Most of them also provided deni-
tions of a few non-skeletal morphological anomalies but
no comprehensive glossary exists for non-skeletal mor-
phological anomalies in amphibians. Moreover, recent
glossaries oen ignored the earlier terminology that
already existed and most did not provide synonymies,
thus creating considerable potential for confusion. Only
B et al. () discussed deviating use of some
terms by dierent authors.
2.2.1 Anomalies of limbs
Acheiria | A synonym of ➝ apody that is rarely used in
studies of amphibian anomalies
Acheiropodia | A rarely used synonym of ➝ apody in
the herpetological literature (R et al. );
in the medical literature (e.g. I et al. ), it
is used as a synonym of ➝ hemimely; to reduce confu-
sion, we recommend using the historically established
term, i.e. ➝ apody
Fig. a: Normal phenotype in an adult male Triturus carnifex, Stanjel, Slowenia, ... Photo: A. G.
Fig. b: Partial albinism in Triturus carnifex; in this male, the
normal colouration is maintained on the ventral parts of the
body; thus, it is not a leucistic individual (but one might call it
partial leucism); born in captivity. Photo: S. M.
Fig. : Translucent skin is the normal phenotype in many
centrolenid frogs, here Hyalinobatrachium eischmanni; Gua-
yabo, Costa Rica, ... Photo: A. K.
22
Henle, Dubois & Vershinin
Fig. a: Scinax fuscovarius: Normal phenotype of an adult; Intervales National Park, Brazil, ... Photo: K. H.
Fig. b: Scinax fuscovarius: abnormal transparency combined with avism in a recently metamorphosed individual. Transpar-
ency is most obvious on the head and digits and most of the body is of a golden yellow colour. Such individuals are oen called
avistic or are described under the generic term albinism. It is best, however, to refer to such individuals as “golden morph” and
to provide a colour photo; Pro Mata Reserve, São Francisco do Sul, Brazil, ... Photo: A. K.
23
Terminology and glossary
Adactyly | Absence of all digits (Fig. a; O
, R et al. ); thus a specic form of
➝ ectrodactyly; not dened in most recent glossaries
of amphibian malformations; some authors use it as a
synonym of ➝ oligodactyly (e.g. J et al. )
Amely | One or several limbs completely lacking (Fig.
; T , M , USFWS , L
, , N , J et al. ,
R et al. ); a specic form of ➝ ectrome-
ly; some recent glossaries (e.g. J et al. , S-
) dene ➝ ectromely in the same way as we
dene here amely; they were followed e.g. by R
et al. (); presumably, this resulted from an incon-
sistency in the denition of ectromely by T ()
(see: ectromely); this denition of ectromely creates
confusion as it deviates from historically established
denitions that are still commonly used; therefore, we
strongly recommend to keep the historically established
denitions in which amely is treated as a subcategory
of ectromely. Absence of limbs is the normal phenotype
only in the order Gymnophiona (Fig. )
Ankylodactyly | A rarely used synonym of ➝ syndactyly
Ankylosis | Various denitions exist: growing together
(R et al. ), stiness of a joint (http://
en.wikipedia.org/wiki/Ankylosis), fusion of two bones
into one (http://www.merriam-webster.com/dictionary/
ankylosis); not commonly used in studies of amphib-
ian anomalies, and if so, mainly for the fusion of bones
of a limb across a joint, sometimes combined with ➝
taumely; we recommend limiting its use in the literature
of anomalies of amphibians to the fusion of bones across
a joint
Anomaly E | A dominant mutation in Rana temporaria
that causes ➝ clinodactyly, ➝ ectrodactyly, ➝ syndac-
tyly, ➝ ectromely and ➝ brachymely; it may further
involve ➝ ectropolydactyly (Fig. ; R ,
D )
Anomaly No | A dominant mutation in Rana tempo-
raria that causes ➝ symphalangy, ➝ brachyphalangy,
➝ clinodactyly and ➝ hypophalangy but aected toes
are dierent from those aected in ➝ Anomaly Ro
(D )
Anomaly P | A syndrome of anomalies characterized
by ➝ polydactyly in benign forms, either only in the
hindlimbs or both in the hind- and forelimbs, whereas
in its severe forms (Fig. ) it includes ➝ brachymely,
inguinal ➝ tumours, bony excrescences and even ➝
polymely; it shows a postero-anterior gradient and a
good, although not always perfect, bilateral symmetry
(R , D )
Anomaly Ro | A dominant mutation in Rana tempo-
raria that causes ➝ symphalangy, ➝ brachyphalangy,
➝ clinodactyly and ➝ hypophalangy but aected toes
are dierent from those aected in ➝ Anomaly No
(D )
Anteversion | A joint of (long) bones oriented in a for-
ward direction (O ), leading to an abnor-
mal positioning of the foot or hand (Figs. & ); thus,
Fig : Complete absence of limbs is the natural phenotype in
the order Gymnophiona, here a female Herpele squalostoma
with its litter; Yaoundé, Cameroon, ... Source: K
et al. (). Photo: M.T. K.
Fig. : Amely, a special form of ectromely, in Rana arvalis;
Ekaterinburg, Russia, ... Photo: V. V.
24
Henle, Dubois & Vershinin
Fig. : Severe form of anomaly P in Pelophylax synkl. esculentus, France. e bottom gure shows particularly well
the bilateral symmetry that is very frequently observed in anomaly P. Source: R ().
25
Terminology and glossary
anteversion is a specic form of ➝ rotation (a term not
used by O ). H () applied antever-
sion more broadly, including ➝ bony triangles, which
is a specic form of ➝ taumely; L () de-
ned anteversion very dierently: as cork-screw shaped
bones (which according to him usually is combined with
taumely); based on the gure given by him, we assume
that it is the same as what we call ➝ taumely; because
of these divergent and oen unclear usages of the term,
we discourage its further use, unless accompanied by a
detailed description and then preferentially restricted to
the denition given by O ()
Aphalangy | Absence of all phalanges of a digit (Figs.
b & a,c); R et al. () dened it as the
absence of some phalanges of ngers, which is incorrect
etymologically and for which another term exists: ➝ hy-
pophalangy; aphalangy and ➝ oligodactyly may be used
for the same type of anomaly, but the reference diers:
the digit in the case of aphalangy and the hand or foot
in the case of oligodactyly; both are subcategories of ➝
ectrodactyly
Apody | Foot (hand) partially or completely missing
(Fig. ); a specic form of ➝ ectromely
Arthrogryposis | It literally means curved joint, imply-
ing that it is xed or stuck in the curved position (S-
et al. ); in the herpetological literature, it is
occasionally used in the same sense, especially when
combined with an underdeveloped musculature (in
German ➝ “Streichholzbeinchen”) (e.g. Z )
and rarely also for ➝ bent bones (L ); most
frequently, however, the English term ➝ sti limbs is
used instead of the technical term. We suggest restrict-
ing the use of the term to its original meaning
Bent bones | ➝ Curved bones
Bidy | A rarely used synonym of ➝ schizodactyly (e.g.
D’A et al. )
Bony bridge | ➝ Bony triangle
Bony extension | ➝ Bony projection
Bony projection | Digit-like microappendices project-
ing from the region of a (limb) bone (M );
these microappendices can be small rudimentary ele-
ments of a supernumerary limb (i.e., ➝ polymely) or
rudiments of distal parts of an incomplete limb (➝
hypomorphic limb); in the latter case, bony projection
is either part of ➝ ectromely (if the microappendix is
Fig. : Anteversion combined with brachymely in a Rana ar-
valis; Ekaterinburg, Russia, ... Photo: V. V.
Fig. : Anteversion of the right hindleg combined with a
shortened foot (brachypody); unidentied species. Photo:
USFWS.
Fig. : Apody in Indosylvirana temporalis; Morningside, Sri
Lanka, . Photo: P. J.
26
Henle, Dubois & Vershinin
a rudiment of the next distal bone, and thus should be
classied as ectromely) or a form of ➝ phocomely (if the
microappendix originates from the region of a proximal
long bone and is a complete or rudimentary digit) and
thus should be scored as phocomely; ➝ femoral projec-
tion is a subcategory of bony projection in which the
microappendices originate from the region of the femur
Bony protuberance | ➝ Bony projection
Bony triangle | ➝ Bent long bone(s) forming a bony
triangle or pyramid (Fig. ; O , R et
al. ), also called ➝ bony bridge (e.g. M
, M et al. , N ); it is a
subcategory of ➝ taumely; J et al. (, )
regarded it as the main form of ➝ taumely and used the
latter term when referring to bony triangles
Brachydactyly | Abnormally short digits; may be due
to a reduced number (➝ hypophalangy; Fig. b,d)
or length (➝ brachyphalangy) of phalanges or a com-
bination thereof (Fig. b). A range of dierent de-
nitions exists. M () restricted the deni-
tion to the one given here for brachyphalangy but did
not include hypophalangy in any of the terms dened
by her; however, USFWS (), which builds on her
denitions, simply dened brachydactyly as short dig-
its, which is equivalent to the denition given here;
N () dened brachydactyly as we dene
hypophalangy. L (, ) dened brachy-
dactyly as normal number of metatarsals but an abnor-
mal number of phalanges, which would also include
➝ schizodactyly and ➝ polyphalangy (and thus ➝
polydactyly according to our denition); this deni-
tion is discouraged as it creates considerable confusion
of disparate patterns; besides, it is dicult to diagnose
in the eld whether the number of metatarsals is nor-
mal or not; V () dened brachydactyly as
symmetrically shortened digits. Z () used the
term ➝ microdactyly as a synonym of brachydactyly;
according to B et al. (), Z also used the
term ➝ ectromely for the anomaly we dene as brachy-
dactyly, but this may be a lapsus calami; brachydactyly
is a specic form of ➝ ectrodactyly; note that several
species, especially within Urodela, have a highly vari-
able number of phalanges in their digits; thus brachy-
dactyly is part of the normal range of variability in these
species and one needs to know this normal range to
decide whether a particular phenotype is abnormal or
not; dierent authors may dier in their opinion for a
particular species
Brachymely | Proportionally shortened limb (Figs. ,
& ), i.e., all long bones of the limbs are present but
at least one is abnormally short (T , O-
, K et al. , R et al.
, V ); in the recent North American
literature, and especially in the nation-wide surveys, the
term ➝ hemimely is oen used for this type of anom-
aly (e.g. M et al. , USFWS , L
, N , W et al. ). While
this denition is also commonly used in the literature
on humans (e.g. C & B ) it deviates from
the historically established denitions that we provide
here and that is widely used in the literature on anoma-
lies in amphibians. As this recent change in the deni-
tion of hemimely can introduce considerable confu-
sion, we discourage its use and recommend sticking to
the original use of the term, which is also etymologi-
cally correct (“shortened limb”). Hemimely on the other
hand means “half limb” and is thus inappropriate for a
shortened limb; J et al. (, ) dened bra-
chymely as “abnormal shortness of one or more limbs”;
while the denition is not explicit enough, comparing
it with the denition they provided for ectromely and
micromely suggests that they used it in the sense given
here for brachymely; S () dened the term
➝ micromely in the same way as we dene brachyme-
ly; R et al. () regarded the two terms as
synonyms; here we regard micromely as a subcategory
of brachymely in which all limb bones are shortened
(as well as being proportionally smaller in diameter);
some Russian authors apply the term ectrodactyly to the
anomaly dened by us a brachymely (de B et al.
); ➝ brachypody is a further subcategory in which
the tarsal or carpal bones are shortened
Fig. : Brachymely in an adult Rana dalmatina; Waldstein-
berg, Germany, May . Photo: K. H.
Fig. : Clinodactyly and oligodactyly of the right hindleg
in a juvenile Rana temporaria; Tschekalin, Russia, ...
Photo: K. H.
27
Terminology and glossary
Fig. b: Specic forms of ectrodactyly: two sub-
categories of brachydactyly: hypophalangy (dig-
its and ) and brachyphalangy (digit ) of the
right foot compared to a normal foot (le); Rana
arvalis; Ekaterinburg, Russia, ... Photo: V.
V.
Brachyphalangy | A bony element of a digit
reduced in length (Fig. b), thus, a specic
form of ➝ brachydactyly; E-P et
al. () also included missing proximal or
middle phalanges in this category, which is
inconsistent with the established use in the
literature; we recommend sticking to its es-
tablished use and to refer to such cases as
➝ hypophalangy or to include them in the
more inclusive term ➝ brachydactyly
Brachypody | Hand or foot completely de-
veloped but some or all bones of the hand or
foot shortened (Fig. ). Not all authors con-
sider this term and include such cases in the
more inclusive term brachymely (V
)
Carpalia abnormal | Bones of the hand
fused, absent or additional bones present;
oen dicult to assess without radiographs
Fig. a: Specic forms of ectrodactyly: adactyly (right arm), adult Bufotes viridis; Magnitogorsk, Russia, ...
Photo: V. V.
28
Henle, Dubois & Vershinin
Cartilaginous spike | A cartilaginous outgrowth from
the end of a limb with missing distal bones (Fig. ; S-
); only individuals that show ectromely can
have this type of anomaly; thus we do not recommend
counting it as an additional anomaly when totalling the
number of anomalies or types of anomalies per indi-
vidual
Clinodactyly | Curvature of digit caused by the pres-
ence of an intercalary little asymmetrical bone between
two phalanges (Fig. ). e denitions provided by T-
() and R et al. () do not include
the intercalary bone, thus also including ➝ curvature of
bones in this category; however, the explanation given
by T () excludes such cases from clinodactyly.
Note that when digits are split (➝ schizodactyly) one
part of the duplicated digit oen bends outwards (Fig.
b); therefore, and because the gure in T ()
can be misunderstood as a schizodactylous individual,
some authors (e.g. V ; note that since
his group follows the denition given here – V
) used the term clinodactyly for cases of schizodac-
tyly; as this diers from the established use of schizo-
dactyly and can create confusion, we discourage this
usage of the term clinodactyly and recommend always
referring to such cases as schizodactyly. Note that clino-
dactyly is dicult to diagnose in preserved specimens
unless radiographs are used
Clinomely | Curvature of limb (F & P );
judging from a poorly reproduced photograph they
seem to use the term as a synonym of ➝ taumely. We
strongly advice not to replace the established term
taumely by clinomely. e term clinomely has rarely
been used; an appropriate use could be as a technical
term for ➝ curved long bones as dened here
Curved long bones | Bones are curved; ➝ torsion of
limbs is sometimes used as a synonym for this type of
anomaly but includes other types of anomalies as well.
Cases in which the curvature involves a bending back
of a bone on itself by > ° or a misalignment with the
rest of the bones are called ➝ taumely; if the abnor-
mal bones have the shape of a triangle or a pyramid,
the terms ➝ bony triangle or ➝ bony bridge are oen
used interchangeably to describe this subcategory of ➝
taumely (J et al. ); we suggest using these
latter terms to specify these types of bent bones or to
describe the type of bending explicitly
Cutaneous fusion | ➝ A band of skin crosses a joint of
long limb bones restricting the motion of that limb; ➝
skin webbing (M , L ) and ➝
skin fold (V ) are commonly used syno-
nyms
Digits swollen | ickening of the epidermis or the
musculature of (parts of) a digit
Diplopody | Complete symmetrical duplication of the
hand or foot including all digits (a specic form of ➝
polypody)
Dipygus parasiticus | A specic form of ➝ polymely
in which the extra limb originates from the distal end of
the pelvic girdle (G ); supercially, it may be
misidentied as a retained tail; the recent literature rare-
ly dierentiates between the dierent types of polymely
Ectrodactyly | Partial or complete absence of one or
more digits (Figs. b, -) or digits shortened (G-
S-H , T , O
). Some recent North American glossaries (e.g. M-
et al. , J et al. , S ,
USFWS , L , ) restricted the term
to the complete absence of a digit (including the meta-
tarsal bone), making it a synonym of ➝ oligodactyly as
dened here. is contradicts with most of the earlier
literature and the denition given here that is also the
common usage outside of North America. We recom-
mend adhering to the established terminology as given
here using ➝ oliogodactyly for a complete absence of at
least one digit (e.g. T ) and treating ➝ adactyly
(Fig. a), ➝ oligodactyly (Fig. c) and ➝ brachydac-
tyly (Fig. b,d) and their subcategories as special cases
Fig. 32c: Specic forms of ectrodactyly: aphalangy (when re-
ferred to a single digit) or oligodactyly (when referred to the
foot), juvenile Rana temporaria; Samarowo, Russia, 18.8.2012.
Photo: K. H.
Fig. d: Specic forms of ectrodactyly: brachydactyly, le
hind limb of a Rana arvalis; Elizavet, Russia, ... Photo:
V. V.
29
Terminology and glossary
of ectrodactyly; R et al. () recommend-
ed the spelling ectodactyly instead of ectrodactyly; we
discourage this, as we never found this spelling in the
literature on amphibian anomalies nor in the medical
literature. Note that some species, e.g. Proteus anguinus
(Fig. ), have less than four ngers and ve toes – the
most common numbers in amphibians – as the normal
phenotype
Ectromely | Partial or complete absence of a limb (Figs.
, , ) (T , N ), except for
cases where proximal bones are missing but the foot or
digits are present (which is called ➝ phocomely); ec-
tromely thus includes ➝ amely, ➝ apody and ➝ hem-
imely. A few recent authors (e.g. M , US-
FWS ) include phocomely in the term ectromely
but most authors do not; a few authors use the term
more restrictively or dierently: () for cases where
the femur is present but the distal parts of the limb are
missing (L ), which we dene as ➝ hem-
imely; () for cases where a limb is completely absent
(e.g. H et al. , S , P et al. ,
R et al. ), which we dene as ➝ amely. We
recommend using the term amely for this specic sub-
category as otherwise most readers will not realize that a
specic subcategory is meant; moreover, etymologically
amely is the correct term for a completely missing limb;
note that while T () also dened ectromely as
the complete absence of a limb, he explained that ec-
tromely can also be partial and included ➝ hemimely
as a subcategory under ectromely; () R et
al. () recommended using the term “ectomely” and
regarded “ectromely” as a synonym of ➝ phocomely; to
our knowledge ectromely has never been used as a syno-
nym of phocomely and the term ectomely is never used
at all. ey also included ➝ adactyly as a subcategory of
ectromely, which again contradicts historical and cur-
rent usage and thus should be avoided; () Z ()
used the term ectromely for what we call ➝ brachydac-
tyly (de B et al. )
Ectropolydactyly | One or several digits are absent
while another digit is duplicated on the same hand or
foot (Fig. ; D ); it may be part of the ➝
anomaly E syndrome. Supercially the hand or foot
may look normal in this type of anomaly; word combi-
nations are not oen used to dene the co-occurrence of
two dierent types of anomalies on the same body part;
instead, most authors would call it ➝ ectrodactyly plus
➝ polydactyly
Femoral projection | A specic form of ➝ bony projec-
tion in which the digit-like microappendages originate
from the tissue of the femur; they may contain ossied ele-
ments (J et al. , K et al. ); only
those individuals exhibiting ➝ ectromely, ➝ phoco
mely,
or ➝ polymely can have this type of anomaly; thus we
recommend not counting it as an additional anomaly
when counting the number of anomalies per individual
Fracture of bone | Broken bone
Front leg(s) remaining covered by the opercular fold |
e front legs of anurans usually remain covered by the
opercular fold until the nal stages of metamorphosis,
when they break through the skin; in this rare anomaly
one or both arms do not break through the skin (Fig. )
Fig. : Bilaterally symmetrical ectropolydactyly (anomaly E) in an adult female Rana temporaria (see D ); the lo-
cation of the metatarsal shows that toe I is duplicated; toe II is absent. At a glance, the individual might be scored as normal;
Nointel near Paris, France, . Photo: A. D.
30
Henle, Dubois & Vershinin
Gastromely | A supernumerary limb inserted at the
venter between the front and hind legs (originating
from the anterior part of the pelvis) (G S-
H ); a specic form of ➝ polymely; in the
recent literature the term is rarely used
Hemimely | Complete loss of the distal half of a limb
(Fig ; T , O , J et al. ,
, S ); hemimely is a subcategory of ➝
ectromely. V () dened it as the absence
of the distal parts of a limb, which may also include ➝
apody; R et al. () dened the term as
“defective limbs, especially distal components”, which is
rather ambiguous and could refer to a range of dierent
types of anomalies. Some recent North American and
Russian glossaries (e.g. M , USFWS ,
L , , N ) dened it as we
dene ➝ brachymely here; this is inconsistent with all
of the earlier herpetological literature and usage outside
of North America but follows the denition of some
authors working with other vertebrates (e.g. C &
B ). e term is also inconsistently dened
in medicine: older inuential lexica (e.g. D
) dened it as we do here; more recently, a wider
denition is oen used that includes ➝ brachymely, but
then usually only in combination with the aected bone.
As the denition provided by us here is traditional, long-
standing and well-established and still most frequently
used in the literature on anomalies in amphibians, we
discourage recent deviations from the established use
of the term as this only creates considerable confusion.
Etymologically this is also preferable as “hemi” and
“brachy” literally mean “half” and “short”, respectively;
thus, applying hemimely to shortened limbs is etymo-
logically incorrect
Hyperdactyly | A specic form of ➝ polydactyly in
which there is an entire extra digit present, with or with-
out the duplication of a metacarpal or metatarsal bone
(Fig. a); the term is rarely used (e.g. W &
G ). N () restricted the term
polydactyly to this type of anomaly and R
et al. () regarded polydactyly and hyperdactyly as
synonyms but the traditional use of the term polydac-
tyly also includes incompletely duplicated digits
Hyperphalangy | Extra phalange present in a digit
(F & H , R et al. ); thus
it is a synonym of ➝ polyphalangy; sometimes, it is
used more broadly as a synonym of ➝ polydactyly (e.g.
S ); for a consistent terminology we suggest
limiting it to cases in which additional phalanges are
linearly arranged within a digit without the splitting of
a digit, which is covered by the term ➝ schizodactyly,
and without a complete extra digit (➝ polydactyly)
Hypodactyly | A synonym of ➝ oligodactyly
Fig. : Right frontleg remaining covered by the opercular fold in a very large tadpole of Bufotes viridis; Roßwag, Germany,
September . Photo: K. H.
31
Terminology and glossary
Hypomorphic limb | e presence of one or more non-
articulating microappendices from the so tissue of a
limb (S ); contrary to the opinion of S-
() this is not diagnostic of amputation (see
H et al. a). In the eld, it may be dicult to
separate from ➝ bony projection. Also, ethymological-
ly, the denition is incorrect as “hypomorphic” means
that a limb is incompletely formed and will therefore
include other types of limb anomalies, such as ➝ ec-
tromely, ➝ phocomely and ➝ brachymely, as well as
their subcategories
Hypophalangy | One or several but not all bones of a
digit are absent (Fig. b,d), i.e. a specic form of ➝
brachydactyly, which in turn is a specic form of ➝ ec-
trodactyly; it includes ➝ monophalangy as a special case
Interdigital webbing | Webbing between digits in spe-
cies that normally do not have such webbing
Joint dislocation | ➝ Luxation
Limb hyperextension | Excessive or rigid exure of a
limb joint (J et al. ); L et al. ()
and L () used the term for an anomaly where
the legs are immobile and straight, with the anomaly be-
ing caused by the knee being locked in extension and
the hip joint also being immobile; thus, his denition is
a subcategory of ➝ sti limbs
Luxation | Displacement of the joint between two limb
bones causing an abnormal articulation of bones; if dis-
location is incomplete, it is called ➝ subluxation
Melomely | ➝ Polymely of the front limbs (Fig. )
(Georoy Saint-Hilaire ); the term is rarely used in
the recent literature
Meromely | Denitions vary; R et al. ()
regarded it as a synonym of ➝ adactyly, however, we
could not nd any publication that used it in that re-
strictive sense. In the medical literature and for other
vertebrates oen dened as partial absence of the limb
skeleton (C & B ); this denition in-
cludes ➝ ectromely, ➝ phocomely and ➝ ectrodactyly
and all subcategories thereof; further denitions exist.
erefore and as it has rarely, if at all, been used in the
literature on anomalies in natural populations of am-
phibians, we recommend using other existing terms; if
it is used, it must be explicitly dened
Metacarpal/metatarsal tubercle lacking | An anomaly
only found in species that usually have a metacarpal or
a metatarsal tubercle, respectively
Metatarsal tubercle abnormally enlarged | Metatarsal
tubercle larger than within the normal range of variation
Microdactyly | Z () used it as a synonym of ➝
brachydactyly
Micromely | Limbs complete but all elements propor-
tionally smaller than normal (M , O
, USFWS , N , L ).
R et al. () treated it as a synonym of ➝
brachymely, which they dened as we do; J et
al. (, ) and S () dened micromely
as “abnormal smallness of a limb”, which may or may
not include brachymely as dened by us; S
() did not include brachymely in his glossary and
thus might have regarded both terms as synonyms; in
contrast, J et al. () also provided a deni-
tion of brachymely: “shorter limb or limbs” and thus it
is likely that they did not regard the terms as synonyms.
In many publications, it remains unclear as to whether
brachymely or micromely as we dene these terms is
what is actually meant; we recommend using both terms
as dened here, thus regarding micromely as a specic
form of ➝ brachymely and describing the anomaly in
sucient detail to unequivocally identify the type of
anomaly observed
Monodactyly | A special case of ➝ oligodactyly, in
which only one digit is present
Monophalangy | A special case of ➝ hypophalangy, in
which only one phalange is present
Nanomely | A rarely used synonym of ➝ micromely
(B et al. , R et al. )
Notomely | A special case of ➝ polymely, in which the
supernumerary limb originates from the dorsum
Fig. : Hemimely in a metamorph of Bufotes viridis with a
short cartilaginous spike; September , Roßwag, Germany,
MNHN .. Photo: H. S.
32
Henle, Dubois & Vershinin
Fig. : Phocomely – le tarsus attached to the femur – in a
juvenile Bufo bufo; Sagra, Russia, . Photo: V. V.
Oligodactyly | Less than the normal number of digits
but at least one digit present (Figs. b, , c) (N-
, R et al. , V ).
T () is not explicit enough to know whether
he regarded oligodactyly as a synonym of ectrodactyly
or as a special case of it as we do; oligodactyly and ➝
aphalangy may be used for the same type of anomaly, but
the reference diers: the digit in the case of aphalangy
and the hand or foot in the case of oligodactyly
Phocomely | An incomplete limb with a proximal bone
missing but distal bones present (Fig. ) (M
, N , R et al. ); oen
the foot (hand) or toes (ngers) are attached directly to
the shoulder or the pelvic girdle; M () re-
garded this anomaly as a special case of ➝ ectromely
but most authors treat the two types of anomalies as
separate categories; therefore, we recommend following
the established use regarding them as separate catego-
ries; B et al. () used phocomely for complete
but malformed limbs, which is inconsistent with the
usual denition of phocomely
Pleuromelophorus tetrachirus | A specic case of ➝
melomely, with four arms; melomely in turn is a sub-
category of ➝ polymely; the term is rarely used in the
recent literature
Pleuromelochirus tetrascelus | A specic case of ➝
pygomely, with four hind legs; pygomely in turn is a
subcategory of ➝ polymely; the term is rarely used in
the recent literature
Polydactyly | Duplication of digit(s) (Figs. & ) or
parts thereof (Figs. a,b) (F , T ,
J et al. , S ); this denition in-
cludes ➝ schizodactyly (Figs. a,b) in which only parts
of a digit are duplicated, ➝ synpolydactyly (Fig. ) in
which the proximal parts of the duplicated digit are
fused, and ➝ hyperdactyly, in which the whole digit is
duplicated (Fig. ). J et al. () dened poly-
dactyly as “supernumerary digit(s)”, which presumably
but not denitely includes schizodactyly. V
() followed the denition given here but earlier pub-
lications of him and some other Russian authors used
the term as a synonym of hyperdactyly (e.g. N
); some authors (e.g. M , M et
al. , L , , R et al. )
and presumably O () restricted the term
polydactyly further to cases in which more than the nor-
mal number of metatarsal/metacarpal bones is present;
R et al. () further regarded polydactyly
and ➝ hyperdactyly as synonyms but the denition of
hyperdactyly generally does not include the presence of
an additional metatarsal/metacarpal; they referred to a
complete duplication of digits without an extra meta-
tarsal/metacarpal bone as ➝ polyphalangy, which also
deviates from the commonly used denition of poly-
phalangy, which is used for linearly arranged duplicated
phalanges; ➝ schizodactyly was not included in the
denition of the above authors nor did they provide a
term for this type of anomaly. Whether a metacarpal or
metatarsal bone is duplicated is very dicult to assess
in the eld and therefore this distinction has rarely been
made for eld samples; therefore, and because the de-
nition given by M () deviates from the more
widely used denition, we recommend adhering to the
traditional denition given here
Polymely | Duplication of a complete limb or parts
thereof (if more than the digits are duplicated) (Figs.
& ; G S-H , T ,
J et al. , S , N ,
V ); the denition includes ➝ schizomely
and ➝ polypody as subcategories. M () pro-
vided a similar denition but excluded polypody; L-
(, ) went one step further and restricted
the term to complete extra limbs. We suggest following
the more widely used broader denition given here. In
the early herpetological literature, but rarely in the re-
cent one, the following terms and their subcategories
were used to dierentiate between the dierent types or
the position of the supernumerary limb: ➝ gastromely,
➝ melomely, ➝ notomely, ➝ pygomely
Polyphalangy | An extra bone inserted into a digit with-
out splitting the digit (i.e. in a linear arrangement). Vari-
mal number of metatarsal/metacarpal bones is present;
R et al. () further regarded polydactyly
and ➝ hyperdactyly as synonyms but the denition of
hyperdactyly generally does not include the presence of
an additional metatarsal/metacarpal; they referred to a
complete duplication of digits without an extra meta-
tarsal/metacarpal bone as ➝ polyphalangy, which also
deviates from the commonly used denition of poly-
phalangy, which is used for linearly arranged duplicated
Polymely | Duplication of a complete limb or parts
thereof (if more than the digits are duplicated) (Figs.
& ; G S-H , T ,
J et al. , S , N ,
V ); the denition includes ➝ schizomely
and ➝ polypody as subcategories. M () pro-
vided a similar denition but excluded polypody; L-
(, ) went one step further and restricted
the term to complete extra limbs. We suggest following
the more widely used broader denition given here. In
the early herpetological literature, but rarely in the re-
cent one, the following terms and their subcategories
were used to dierentiate between the dierent types or
the position of the supernumerary limb: ➝ gastromely,
➝ melomely, ➝ notomely, ➝ pygomely
Polyphalangy | An extra bone inserted into a digit with-
out splitting the digit (i.e. in a linear arrangement). Vari-
33
Terminology and glossary
ous other denitions exist: N () included
➝ schizodactyly as dened by us in her denition of
polyphalangy and H () applied the term to any
extra phalanges, whether linearly arranged (i.e. poly-
phalangy in our denition) or with splitting (➝ schizo-
dactyly in our denition) or a complete duplication of
digits (➝ hyperdactyly in our denition); thus he used
it as a synonym of ➝ polydactyly. M () and
M et al. () used the term for another type
of ➝ polydactyly, in which there is (are) extra digit(s)
present without a duplication of the metatarsal bone(s);
except for in the recent North American literature (e.g.
MD et al. ), this denition is rarely applied
in the literature on anomalies of amphibians (and very
Fig. : Polydactyly in Pelophylax ridibundus; Oka, Tschekalin, Russia, ... Photo: K. H.
Fig. : Polydactyly, with proximal fusion of the duplicated
ngers, in Pelophylax esculentus. Although one could call this
anomaly polydactyly plus syndactyly or synpolydactyly, we
recommend counting only the primary anomaly, in this case
polydactyly; Seelenhofer Ried, Germany (compare M
et al. ). Photo: C. M.
Fig. b: Schizodactyly in the foot of an adult Rana arvalis.
Note: the supernumerary toe bends away from the normal
toe; although one could call it polydactyly plus clinodactyly
we recommend not counting it as clinodactyly because the
bending is a consequence of the primary anomaly schizodac-
tyly; Kalinovsky forest park, Ekaterinburg, Russia, ...
Photo: V. V.
Fig. a: Schizodactyly, a specic form of polydactyly, in the
right hind limb of a Lissotriton vulgaris; Ekaterinburg, .
Photo: V. V.
34
Henle, Dubois & Vershinin
dicult to apply in the eld). e denition provided by
R et al. () is not very precise but presum-
ably means the same as the denition provided by M-
(). To reduce ambiguity, to make terminol-
ogy consistent and in particular because the distinction
between the dierent types of duplicated digits made by
M () is dicult in the eld and therefore
has rarely been made for eld samples, we recommend
using the denition given here for ➝ polyphalangy, ➝
schizodactyly and ➝ hyperdactyly if one can identify
the nature of the duplication and otherwise using the
more inclusive term ➝ polydactyly
Polypody | A limb with one or more extra feet (hands)
or parts thereof (J et al. , , S
) but more than the duplication of digits (Fig. ); a
specic form of ➝ polymely
Pseudomely | V et al. () used the term as a
synonym of ➝ polymely
Pygomely | A specic form of ➝ polymely, in which the
hind limb(s) are duplicated (Fig. )
Rotation (of limbs) | Limbs appear twisted and the foot
(hand) is not at with the surface; as dened here, this
type of anomaly can either be caused by a distortion of
the direction of the bone growth (➝ curved bones) or
an abnormal articulation of two bones (➝ anteversion;
Figs. & ); M () and N ()
restricted the term to the rst case; a rotation of limbs is
always combined with ➝ sti limbs
Schizodactyly | Forked digits (Figs. a,b); this is a spe-
cic form of ➝ polydactyly; rarely called ➝ bidy (e.g.
D’A et al. ). e denition given by N-
() for ➝ polyphalangy includes schizodac-
tyly but not our denition. Note that the supplementary
phalanges oen bent away from the base of the toe (Fig.
b); therefore and because the gure of ➝ clinodactyly
in T () – but not his denition of it – is mis-
leading, some authors (e.g. V ) referred
to cases of schizodactyly as clinodactyly. As incomplete
duplication of a digit is the primary type of anomaly,
we discourage this usage and recommend to call such
cases schizodactyly to distinguish them from cases of ➝
clinodactyly that do not involve the partial duplication
of a digit. Since V () follows the de-
nition given here
Schizomely | Completely duplicated limbs in which the
proximal parts of the limbs are fused; thus it is a specic
form of ➝ polymely
Skin fold | A synonym of ➝ cutaneous fusion (V-
)
Fig. : Melomely, a specic case of polymely in a captive o-
spring of Dendrobates tinctorius. Photo: S. K.
Fig. : An unusual form of polypody, which is a specic form of polymely, in an adult female Lissotriton vulgaris; the two super-
numerary feet show dierent degrees of duplication, the lower one would not be regarded as duplication of the foot but rather
as a bony protuberance following the classication of M (); Bayreuth, Germany, spring . Photo: K. S.
Pygomely | A specic form of ➝ polymely, in which the
hind limb(s) are duplicated (Fig. )
Rotation (of limbs) | Limbs appear twisted and the foot
(hand) is not at with the surface; as dened here, this
type of anomaly can either be caused by a distortion of
the direction of the bone growth (➝ curved bones) or
an abnormal articulation of two bones (➝ anteversion;
Figs. & ); M () and N ()
restricted the term to the rst case; a rotation of limbs is
always combined with ➝ sti limbs
Schizodactyly | Forked digits (Figs. a,b); this is a spe-
cic form of ➝ polydactyly; rarely called ➝ bidy (e.g.
D’A et al. ). e denition given by N-
() for ➝ polyphalangy includes schizodac-
tyly but not our denition. Note that the supplementary
phalanges oen bent away from the base of the toe (Fig.
b); therefore and because the gure of ➝ clinodactyly
in T () – but not his denition of it – is mis-
leading, some authors (e.g. V ) referred
to cases of schizodactyly as clinodactyly. As incomplete
duplication of a digit is the primary type of anomaly,
we discourage this usage and recommend to call such
cases schizodactyly to distinguish them from cases of ➝
clinodactyly that do not involve the partial duplication
of a digit. Since V () follows the de-
nition given here
Schizomely | Completely duplicated limbs in which the
proximal parts of the limbs are fused; thus it is a specic
form of ➝ polymely
Skin fold | A synonym of ➝ cutaneous fusion (V-
)
35
Terminology and glossary
Skin webbing | A synonym of ➝ cutaneous fusion (US-
FWS , N )
Spindly limbs | English term for ➝ Streichholzbeinchen
Sti limbs | Limbs are immovable; stiness is linked to
other anomalies of bones, such as ➝ polymely (Fig. ),
➝ rotation of limbs (Figs. & ) or ➝ taumely (Fig.
), to ➝ cutaneous fusion or the reduction of muscu-
lature; the latter is called ➝ Streichholzbeinchen in the
German amphibian husbandry literature
Streichholzbeinchen | e German term for ➝ sti limbs
with degenerative musculature, giving the limbs a spindly
appearance. is anomaly is called the xed limb syndrome
by K (). It is a subcategory of sti limbs
Subluxation | Incomplete or partial ➝ joint dislocation
(O )
Symmely | Partial or complete fusion of two limbs (P
& F , R et al. ). We only know
this type of anomaly from supernumerary limbs that
are partially fused with the normal limb, from Bombina
bombina in Hungary (P & F ) and from a
single individual of Pelophylax esculentus described by
B () – he called it ➝ sympody – in which the
hindlegs were fused at the level of the femur and not
just the feet. In the case of the fusion of supernumer-
ary limbs, the primary anomaly is polymely; therefore,
we recommend counting such cases as polymely as op-
posed to listing them additionally as symmely. O
() and J et al. () provided a dierent
denition for symmely: fusion of a limb or parts of a
limb to a body part; we have not found any publication
on anomalies in natural populations of amphibians that
described such an anomaly; symmely is a specic ex-
pression of ➝ synostosis
Symphalangy | Two phalanges of the same digit are
fused (D ); it is a subcategory of ➝ syndactyly
Sympody | Only the distal parts of two limbs are fused
(B ); a very rare type of anomaly; the individ-
ual described by B (), however, would rather
classify as ➝ symmely to which sympody belongs as a
subcategory; sympody is a specic expression of ➝ syn-
ostosis
Syndactyly | e partial or complete fusion of two or
more digits due to failure of the so tissue to break
down between digits or due to the fusion of bones of
dierent digits (T , O , J
et al. , , S , USFWS , L
, , R et al. , V ),
the latter being termed ➝ symphalangy; syndactyly is a
specic expression of ➝ synostosis
Synostosis | Fusion of bones
Synpolydactyly | Duplication of digits, with digits
fused by so tissue or with proximal phalanges partially
fused (Fig. ; C & B ); ➝ schizodactyly
is similar but the proximal phalanges are single rather
than duplicated and fused; synpolydactyly is a subcat-
egory of ➝ polydactyly
Tarsalia abnormal | Bones of the foot are either fused or
absent or additional bones are present
Taumely | Gross disturbance of the limb plan, whereby
the misalignment of a long bone is more than ° (Fig.
; T , V , ). e degree to
which the bone bends back may vary and some authors
(e.g. G & H ) included any degree of
bending back within the term taumely, whereas oth-
ers (e.g. T , M , M et al.
, J et al. ) restricted it to the denition
given here. We suggest including also cases with a lesser
degree of bending if the bauplan of the limb is grossly
disturbed. e abnormal bone may have the shape of
a pyramid or triangle; such cases are oen called ➝
bony triangle or ➝ bony bridge (e.g. M et al.
); however, it is not entirely clear whether M-
() and J et al. () regarded bony
triangles as a synonym or as a subcategory of taumely;
we recommend the latter to separate these anomalies
from other types of taumely; we further suggest calling
slightly bent bones ➝ curved bones to separate them
from grossly misaligned bones
Fig. : Particularly grossly malformed le hindlimb that
looks like a bony triangle, which is a special case of taume-
ly. e malformation is, however, complex, since the tibia is
either missing or completely synostosed with the femur and
the foot is reduced in size and the number of digits; in such
cases it is best to describe and provide a gure of the observed
individual and not simply allocate them to a particular term
of anomaly; juvenile Rana arvalis; Ekaterinburg, Russia,
... Photo: V. V.
36
Henle, Dubois & Vershinin
Toe pad swollen | Epidermis of toe pad thickened
Torsion of limbs | A synonym of ➝ rotation of long
limb elements
Triphalangy | A specic form of ➝ oligodactyly, in
which three digits are present (Fig. b)
Twisted limbs | A synonym of ➝ rotation of limbs
2.2.2 Anomalies of the head, body or tail
Aglossia | Tongue absent (N )
Agnathia | Jaw completely missing (N )
Anadydimus | Two tails (W )
Anal tube closed | Anal tube distally covered by skin
Anophthalmy | Eye(s) lacking (blindness) (Fig. ;
T , M , J et al. , S-
, USFWS , N , L
, V ); L () called this type
of anomaly microphthalmy, which was a lapsus calami;
unilateral anophthalmy is sometimes called ➝ cyclopy
by Russian authors (e.g. Z , F ) but
cyclopy usually is dened dierently
Asyntaxia caudalis | Duplication of tail due to a failure
of the anal fold to close; it is a specic expression of ➝
anadydimus
Axial exure | Torsion of the vertebral column
Axial incurvation | ➝ Axial exure
Beak lacking keratinization | e jaw sheaths of the
mouths in tadpoles are developed but not keratinized or
the keratin was destroyed by pollution or disease; there-
fore beaks are not pigmented; note that lack of keratini-
zation may also be due to low temperature (R
) and thus not an abnormal condition
Bent tail | e tail is bent instead of being straight; ➝
kinky tail is a synonym
Bicephaly | Head duplication; a synonym of ➝ diceph-
aly and ➝ catadydimus
Brachycephaly | Head abnormally short but of normal
width; in post-embryonic stages, this type of anomaly is
rare (V , H et al. a)
Brachygnathia | Abnormal shortness of the lower jaw
(M , L ); usually either used
Fig. 43: Anophthalmy in a captive bred Cruziohyla cf. calcari-
fer, ... Photo: J. K.
Fig. 44: Cauda bida in a tadpole of Hyla arborea; riptis
Mountains, Crete, Greece, ... Source: H et al.
(); Photo: K. H.
as a subcategory or a synonym of ➝ mandibular hy-
poplasia
Brachyuria | Shortened tail (R et al. );
note that the term is used only for inherited shortness
of tails and not for lost parts of a tail; this term is rarely
used in the literature on amphibian anomalies
Catadydimus | Two headed (W ); a syno-
nym of ➝ dicephaly and ➝ bicephaly that is frequently
used for snakes but rarely for amphibians
Cataract | Clouding of the eye lens, which leads to a de-
crease in vision
Cauda bida | Terminal part of tail duplicated (Fig. )
Cle lip | Lip with a ssure (Fig. )
Curvature of tail | ➝ Bent tail
Cyclopia | Eyes completely fused into one single median
eye (A ). Note that some Russian authors
(e.g. F ) call unilateral ➝ anophthalmy cy-
clopy
Dicephaly | Two heads (Fig. ; O ); ➝ bi-
cephaly and ➝ catadydimus are synonyms
37
Terminology and glossary
Duplicitas anterior | Duplication of anterior parts of the
body; ➝ Janus twins is a specic form of it; S
() restricted the term to duplication of the head, i.e.
used it as a synonym of ➝ dicephaly; K ()
included ➝ polymely of the forelimbs (i.e. ➝ melome-
ly) under this term
Duplicitas posterior | Duplication of posterior parts of
the body; K () included ➝ polymely of the
hindlimbs (i.e. ➝ pygomely) under this category
Exophthalmy | Eyes protruding abnormally
Eye displacement | Eye displaced laterally, medially,
cranially or caudally (L , , N
); called abnormal eye position by V ()
Gigantism | Body size much larger than the normal
maximum size
Gut miscoiling | Abnormal coiling of the gut; most
common is a loose coiling
Hump | ➝ Kyphosis of the spine
Hypognathia | A synonym of ➝ mandibular hypoplas-
ia and ➝ mandibular dysplasia
Janus twins | Duplication of dorso-anterior structures
in the absence of duplication of posterior structures; i.e.
a specic form of ➝ duplicitas anterior
Jaw shape abnormal | Jaw fully developed but upper
and lower jaw dier in shape and mouth does not close
completely
Jaw sheaths abnormal | A specic form of abnormal
➝ mouthparts in larval amphibians, in which there are
unnatural breaks, gaps or other deformities in the jaw
sheaths or the jaw sheaths may lack keratinization
Keratinized denticles absent | e labial tooth rows
lack denticles or denticles are not keratinized in species
that normally have keratinized denticles; a specic form
of ➝ tooth rows abnormal
Fig. 46: Dicephaly in a captive born larval Salamandra sala-
mandra (G & G ). Photo: C. G.
Fig. 45: Cle lip, presumably due to injury, in an adult Bufo bankorensis; Taroko, Taiwan, ... Photo: K. H.
38
Henle, Dubois & Vershinin
Kinking | Torsion of notochord, vertebral column or
tail; ➝ kyphosis, ➝ lordosis and ➝ scoliosis are spe-
cic expressions of kinking
Kinky tail | ➝ Bent tails (T et al. ); a specic
form of ➝ kinking, in which the tail is aected
Kyphosis | Abnormal backward curvature of the spine
(tail or body) (M , O , J-
et al. ); a specic expression of ➝ kinking;
note that M () does not include ➝ lordo-
sis in her glossary, while USFWS () included it but
not kyphosis; they seem to use the two terms as syno-
nyms, as is also indicated by photos on the website
http://www.fws.gov/contaminants/amphibian/Pictureumbs.html
Labial papillae abnormal | Labial papillae underdevel-
oped, interrupted or absent
Laceration | Skin wound (Fig. )
Lesion | Pathological area of an organ (R et
al. ); may be due to injury, disease or ➝ tumour
Lordosis | Abnormal forward curvature of the spine (tail
or body) (O , J et al. , R-
et al. , V ); a specic expression
of ➝ kinking; note that USFWS () does not include
➝ kyphosis in its glossary, while M () in-
cluded it but not lordosis; they seem to use the two terms
as synonyms, as is also indicated by photos on the website
http://www.fws.gov/contaminants/amphibian/Pictureumbs.html
Macrophthalmia | Abnormally large eyes (F &
P )
Mandibular dysplasia | A synonym of ➝ mandibular
hypoplasia (J et al. , S ), the
latter being more commonly used
Mandibular hypoplasia | Lower jaw underdeveloped or
completely missing (T , J et al. )
but the term may be used also in a more restrictive
way that excludes completely missing jaws. ➝ Brachy-
gnathia and ➝ agnathia are specic types of mandibular
hypoplasia as dened here. e following terms are less
commonly used synonyms: ➝ mandibular dysplasia, ➝
hypognathia and ➝ micrognathia
Maxillary hypoplasia | Upper jaw underdeveloped or
completely missing
Microcephaly | Abnormally small head (M
, USFWS , N , R et
al. ); either the whole head or parts thereof may be
reduced in size or shortened. L (, ) de-
ned the term dierently: “blunt nose; shortened upper
jaw”; while such an anomaly is a specic form of micro-
cephaly, most authors dene the term more broadly; for a
shortened upper jaw a dierent technical term exists: ➝
maxillary hypoplasia; moreover, etymologically, “micro”
means “small” and “cephaly” means “head” Microcephaly
is a rare type of anomaly with the exception of embryos
Micrognathia | A synonym of ➝ mandibular hypoplas-
ia (N )
Microphthalmy | Eye(s) abnormally small (Fig. )
(M , O , N ,
L , J et al. , V );
sometimes also spelled microophthalmy; as a lapsus
Fig. 47: Laceration (skin wound) on the head of a juvenile Rana arvalis; in early stages of healing, the skin is transparent;
Samarowo, Russia, ... Photo: K. H.
39
Terminology and glossary
calami, L () called this type of anomaly an-
ophthalmy
Monorhyny | Having a single nostril (R ,
O , R et al. , V
); so far only cases are known that are caused by a
fusion of the nostrils, i.e. ➝ synrhyny
Mouthparts abnormal in larvae | Beaks, labial teeth
rows or labial papillae reduced in size, number or form,
misplaced or abnormal in shape
Mydriasis | Excessive dilation of the eye (S et
al. )
Myiasis | Lesions caused by the maggots of parasitic ies
that deposit eggs on living individuals; in severe cases,
the nasal openings may be destroyed (Fig. )
Nanism | Body size much smaller than normal; in natu-
ral populations this type of anomaly is very dicult to
distinguish from small individuals that are still in the
process of growing
Nares closed | Failure of nostrils to open
Nasal opening destroyed | is anomaly is caused by
maggots of the parasitic y Lucilia bufonivora that depos-
its eggs on the nostrils or other body parts (Fig. ); the
larvae migrate to the nostrils and destroy them killing the
infected individual; usually Bufo bufo is the host but rare-
ly other species are also parasitized (H et al. a)
Nose blunt | Nose truncated in species that usually have
a rounded snout
Opening underneath opercular fold unusually wide |
Resorption of the opercular fold was abnormally large
Ophistocony | Used as a synonym of ➝ lordosis by
F & ()
Oral labial papillae swollen | A specic form of the
category ➝ mouthparts abnormal, in which the labial
papillae are expanded in size
Osteolathyrism | Decreased connective tissue strength
(S & C ); this anomaly leads to
notochord and tail deformities in embryos and to ➝
joint dislocation and ➝ bent long bones in metamor-
phosing anurans
Otocephaly | Absent or underdeveloped lower jaw
(R et al. ); this denition includes ➝
mandibular hypoplasia and ➝ agnathia; both terms
were not included in the glossary of R et al.
(); whereas we did not nd any use of the term oto-
cephaly in the literature on amphibian anomalies, the
latter terms are commonly used
Palatine eye | Eye in the mouth (O )
Panophthalmitis | Infection of the whole eye
Polyophthalmy | More than two eyes (F & P
)
Sacrum asymmetric | Sacral processes inserted on dif-
ferent vertebrae
Scars | Skin wound, scratches; may not be abnormal in
territorial species that ght with each other
Scoliosis | Abnormal lateral curvature of the spine (tail
or body) (M , O , L
, J et al. , R et al. ,
V ); a subcategory of ➝ kinking
Siamese twins | Two almost complete individuals that
are fused to each other and share most body parts
Skeletal kinking | ➝ kinking involving the skeleton
Snout pointed | Snout is pointed instead of being round
in species that normally have a round snout
Spiraculum number abnormal | More or fewer than
the normal number of spiracula (H-R );
most tadpoles have a single spiraculum as the normal
phenotype but tadpoles of rhynophrynids, pipids and
Lepidobatrachus have two spiracula (MD &
A )
Spiraculum misplaced | Spiraculum on the wrong side
of the body or ventrally in species in which the normal
position is lateral; the most common normal phenotype
is a single spiraculum on the le side of the body (M-
D & A )
Synrhyny | Fused nostrils; so far this type of anomaly is
the only known form of ➝ monorhyny
Tail bifurcation | ➝ Cauda bida
Tail duplication | Tail is partially or completely dupli-
cated; includes ➝ Asyntaxia caudalis and ➝ Cauda bi-
da as subcategories
Tail n reduced | Parts of the tail n lacking or reduced
in height
Tail retention in metamorphosed anurans | Tail is not
completely reabsorbed during metamorphosis; note
that in some species tail resorption may be completed
only aer individuals have le the water (V
, H et al. a)
Tail stunted | Tail shorter than normal (T )
40
Henle, Dubois & Vershinin
Fig. 48: Microphthalmy (top) in an adult male Rana arvalis; for comparison the normal eye of the le side is shown (bottom);
Ekaterinburg, Russia, ... Photo: V. V.
41
Terminology and glossary
Tooth rows abnormal | Tooth rows in tadpoles mis-
shaped, interrupted, lacking denticles or ➝ keratiniza-
tion of denticles absent
Torsion of body | ➝ kinking
Torsion of tail | ➝ Bent tails
Twinning | Duplication of body parts; ➝ catadydimus,
➝ dicephaly, ➝ duplicitas anterior, ➝ duplicitas pos-
terior, ➝ Janus twins, ➝ Siamese twins are specic
forms of twinning; duplications that involve only the
limbs, the tail or parts thereof are usually not called
twinning
Tympanum absent | Abnormal in species whose nor-
mal phenotype is an externally visible tympanum; note
that in some species this is the normal condition
Urostyle bent | Curved bones of the urostyle (V-
)
Vertebral column sti | Vertebral column immovable,
usually due to ➝ vertebral fusion
Vertebral fusion | Two or several vertebrae are fused
Vertebral column truncated | Vertebral column con-
siderably reduced in length (H et al. b: Fig. )
. Edema, tumour and similar types of anomalies
G & H () provided a terminol-
ogy for anomalies related to tumours and B et
al. () and G () explained terms related to
edema. Here we only include terms for externally vis-
ible anomalies. Many of these types of edema occur fre-
quently in embryos but are rarely observed in the eld
at later stages of development.
Ascites | ➝ Edema of the peritoneal cavity (belly) (Fig.
; O ); ➝ hydrocoelom is a synonym;
since amphibians do not have separate thoracic and
abdominal cavities, the term is regarded as inappropri-
ate for amphibians by G (); some authors (e.g.
F & B ) used the term more
broadly as a synonym of edema
Blister | A vesicle of the skin containing uid (B
et al. ); a specic type of ➝ edema
Bloatedness | Swelling of the entire body; usually no dif-
ferentiation is made as to whether gas or liquid accumu-
lation caused bloatedness (e.g. L ), and it is
not always easy to make this distinction in the eld; we
recommend using either ➝ gas accumulation or ➝ ede-
ma when the cause (gas / liquid) of the swelling is known
Carcinoma | Malignant form of ➝ neoplasia; contains
➝ melanoma as a subcategory
Cephalic edema | ➝ Edema of the head; ➝ hydroceph-
aly and ➝ hydroencephaly are generally used as syno-
nyms, though the later does not necessarily involve the
whole head in tadpoles or later stages
Cyst | Swelling caused by abnormal tissue growth to
encapsulate parasites; also used for swellings that con-
tain mushy material; the denition for cysts makes it a
subcategory of ➝ tumour but usually the term cyst is
regarded as a separate category
Fig. 49: Nasal openings destroyed by maggots of Lucilia bu-
fonivora in an adult Bufo bufo; Bialogard, Poland. Source:
Wikipedia Commons. Photo: R. A
Fig. 50: Edema in a juvenile Lissotriton vulgaris; Ekaterinburg,
Russia, ... Photo: V. V.
42
Henle, Dubois & Vershinin
Dysplasia | Abnormal development of structures, such
as muscles or organs, conventionally used for non-ma-
lignant abnormal structures (R et al. )
Edema | Subcutaneous accumulation of uid (Fig. ;
J et al. , , S , V
); ➝ hygroma is a rarely used synonym; ➝ hydrops
is used as another synonym by some authors but regard-
ed as a subcategory by others; edema is a subcategory
of ➝ bloatedness and contains the following subcatego-
ries: ➝ ascites, ➝ hydrocephaly, ➝ hydrocoelom and
➝ hydroencephaly
Epithelioma | A synonym of ➝ papilloma (P
et al. )
Gas accumulation | A subcategory of ➝ bloatedness, in
which the swelling of the body is caused by gas
Granuloma | ➝ Tumour caused by an inammatory
swelling (G & H )
Gut miscoiling | Abnormal coiling of the gut; most
common is a loose coiling
Hydrocephaly | ➝ Edema of the head (e.g. H
); ➝ cephalic edema is a synonym; because the most
common form is an enlargement of the brain ventricles
due to an excessive amount of uid; hydrocephaly and
➝ hydroencephaly are generally used interchangeably
(e.g. P-Z et al. )
Hydrocoelom | Fluid accumulation in the body cavity
(Fig. ); a synonym of ➝ ascites, if ascites is dened
as it is here
Hydroencephaly | ➝ Edema of the brain (B et al.
, V ); ➝ hydrocephaly is oen used as
a synonym, although strictly speaking hydroencephaly
is a specic form of hydrocephaly in which the brain is
aected
Hydrops | A synonym of ➝ edema; G () pre-
ferred to use this term only for cases in which the sub-
cutis, tissues and coelomic cavity of larval amphibians
are jointly aected; E () used it for edema of
the subcutaneous lymph sacs in larval and adult am-
phibians
Hygroma | Used as a synonym of ➝ edema by L
()
Hyperplasia | Abnormally strong development of struc-
tures, such as muscles or organs; a subcategory of ➝ tumour
Melanoma | Malignant ➝ neoplasia arising from the
melanocytic system of the skin (R et al.
) or other organs; melanomas are characterised by
black pigmentation
Neoplasia | Development of abnormal new structures
(G & H , R et al. ,
V ); a subcategory of ➝ tumours; contains
➝ carcinoma as a subcategory
Nodule | ➝ Swelling or lump; may be lled with lipids,
viral particles, spores of fungi or parasitic microorgan-
isms; oen an initial stage in the formation of tumours
(e.g. L & S )
Papilloma | ➝ Neoplasia in which the ectoderm shows
an abnormal outgrowth in the form of a horny epitheli-
um or warts (Fig. ); plural: papillomas or papillomata;
➝ epithelioma is a synonym
Ulceration | Sore or lump; may originate from infection
or tumour (e.g. L & S )
Tumour | Abnormal mass of tissue resulting from ex-
cessive cell division (Fig. ); a tumour may be inam-
matory (➝ granulomata), parasitic (encysted immature
trematodes), ➝ hyperplastic or ➝ neoplastic (G
& H ); malignant tumours are called
➝ carcinoma; note that abnormal tissue mass produced
to encapsulate parasites is most commonly referred to
as a cyst and not as a tumour, though the denition of
tumours also includes cysts
. Anomalies of eggs and early embryonic stages
As most of these anomalies are only studied in the labo-
ratory, here we only list a few types of anomalies that
can easily be detected in the eld with a hand lens. See
B et al. () for a more detailed description of
various embryonic anomalies
Acephaly | Head structures lacking
Asyntaxia medullaris | A synonym of ➝ spina bida
Axial duplication | ➝ Spina bida
Clutch abnormal | Common clutch anomalies are
clutches lacking eggs (Fig. ), eggs without embryos,
twin embryos, membrane lacking between embryos
(Fig. ) and abnormally white eggs (see ➝ albinism
and ➝ transient albinism in section .) (V
, )
Cyclopia | Eyes completely fused into one single medi-
an eye (A , S ); Z ()
and F () used the term as a synonym of uni-
lateral ➝ anophthalmy
Microcephaly | Head reduced in size (N )
Spina bida | Duplication of (parts of) the vertebral col-
umn, usually combined with duplications of major parts
of the body (H ); ontogenetically, the
43
Terminology and glossary
correct name should be ➝ asyntaxia medullaris (B-
) but this name has not become established
(see also ➝ twinning; ➝ Siamese twins)
Supernumerary appendages | Duplication of embry-
onic structures
Synophthalmy | Eyes are close-set, can adhere more
or less to each other but remain distinct, which distin-
guishes it from ➝ cyclopy (S )
. Other terms
Abnormality | Gross deviation from the normal range
in morphological variation (J et al. , US-
FWS )
Aneuchrony | e speed of development is accelerated
or delayed compared to the normal condition (D
); ➝ heterochronic and ➝ homochronic aneuchro-
ny are subcategories
Anomaly | Any deviation of the phenotype (morpho-
logical and non-morphological) from the range of vari-
ation of the phenotype considered to be normal and ir-
respective of its cause
Deformity | Alteration of an organ or structure that
originally formed correctly (J et al. , US-
FWS )
Heterochronic aneuchrony | Dissociation of the devel-
opment rate of characters, some being either accelerated
or decelerated compared to the normal development
rate of other characters (D ); ➝ neoteny is a
common form of heterochronic aneuchrony
Homochronic aneuchrony | e speed of development de-
viates from the normal rate for all characters (D )
Fig. 51: Papilloma in Cynops pyrrhogaster; Japan. Photo: M. A & V.B. M-R.
44
Henle, Dubois & Vershinin
Malformation | A permanent structural defect result-
ing from abnormal development (J et al. ,
USFWS )
Neoteny | A specic form of ➝ heterochronic aneu-
chrony, in which development of the somatic charac-
ters is delayed or arrested but not that of gonads, so that
the resulting giant larvae are able to reproduce (Fig. )
(D )
Paedomorphosis | Retention of juvenile traits in adults
(D )
Acknowledgments
We thank M U, Leipzig, and the UFZ library
team, for their assistance in compiling the literature.
Please note that copyrights may apply to all those photos
published with this article. Photos retrieved from Wiki-
media Commons (http://commons.wikimedia.org) were
published under the creative commons licence (https://
creativecommons.org/licenses/by-nc-sa/2.0/). See the cit-
ed webpage for conditions of using these photos. Pho-
tos of the United States Fish and Wildlife Service are
in the public domain and were retrieved from http://
www.fws.gov/contaminants/amphibian/Pictureumbs.
html. We owe thanks to all photographers who made
photos available via these platforms or who provided
us with their photos directly. VV was supported by Act
Fig. 52: Abnormal clutches of Salamandrella keyserlingii: le: eggs arranged in strings lacking individual membranes; right: ab-
normally low number of eggs; Ekaterinburg, Russia, ... Photo: V. V.
45
Terminology and glossary
Government of the Russian Federation, agreement
.A... Above all, we would like to thank
J-C B, Toulouse, and A C-
, Paris, for carefully reviewing and commenting
on the glossary, which helped us to clarify synonyms
and deviating denitions. KH thanks N R-
for her help in translating Russian literature
and for her patience during many hours of work on the
manuscript.
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