Taxonomic and nomenclatural status of Iberian Algyroides (Lacertidae)

Abstract

The taxonomy and nomenclature of Iberian Algyroides are problematic. The first taxon described, A. hidalgoi Boscá, 1916, was based on a single specimen that was subsequently lost. The description of the second taxon, A. marchi Valverde, 1958, was based on the comparison of a newly discovered population with the original description of A. hidalgoi. However, A. hidalgoi specimens have never been recorded since for any locality. Therefore, three questions need to be addressed: Is A. hidalgoi Boscá, 1916 a morphologically diagnosable taxon different from all non-Iberian species of Algyroides? are A. hidalgoi and A. marchi conspecific? And if so, which is the correct name for the species? To clarify the taxonomic status of the Iberian Algyroides we (1) compare Boscá’s A. hidalgoi original description against the descriptions of all other species of Algyroides, (2) test the accuracy of Boscá’s A. hidalgoi by comparing it against 204 Iberian museum specimens, and (3) designate a neotype of A. hidalgoi that fits the head pholidosis described in the original description. We show that none of the diagnostic characters used by Valverde to differentiate between A. hidalgoi and A. marchi are actually diagnostic, as we found high levels of variability on those characters in the studied specimens. Our results validate Boscá’s description of A. hidalgoi, which fits within the morphological variability observed for southern Iberian Algyroides. As a result, we propose the strict synonymy of A. marchi Valverde, 1958 with A. hidalgoi Boscá, 1916.

Full text

Amphibia-Reptilia (2018) DOI:10.1163/15685381-20181016
Taxonomic and nomenclatural status of Iberian Algyroides
(Lacertidae)
Alberto Sánchez-Vialas1,∗, Marta Calvo-Revuelta1, José Luis Rubio2, Fernando Palacios1,
Mario García-París1
Abstract. The taxonomy and nomenclature of Iberian Algyroides are problematic. The first taxon described, A. hidalgoi
Boscá, 1916, was based on a single specimen that was subsequently lost. The description of the second taxon, A. marchi
Valverde, 1958, was based on the comparison of a newly discovered population with the original description of A. hidalgoi.
However, A. hidalgoi specimens have never been recorded since for any locality. Therefore, three questions need to be
addressed: Is A. hidalgoi Boscá, 1916 a morphologically diagnosable taxon different from all non-Iberian species of
Algyroides?areA. hidalgoi and A. marchi conspecific? And if so, which is the correct name for the species? To clarify the
taxonomic status of the Iberian Algyroides we (1) compare Boscá’s A. hidalgoi original description against the descriptions
of all other species of Algyroides, (2) test the accuracy of Boscá’s A. hidalgoi by comparing it against 204 Iberian museum
specimens, and (3) designate a neotype of A. hidalgoi that fits the head pholidosis described in the original description. We
show that none of the diagnostic characters used by Valverde to differentiate between A. hidalgoi and A. marchi are actually
diagnostic, as we found high levels of variability on those characters in the studied specimens. Our results validate Boscá’s
description of A. hidalgoi, which fits within the morphological variability observed for southern Iberian Algyroides.Asa
result, we propose the strict synonymy of A. marchi Valverde, 1958 with A. hidalgoi Boscá, 1916.
Keywords:Algyroides hidalgoi,Algyroides marchi, morphological variability, neotype designation, nomenclature, nov.
synonymy, taxonomy.
Introduction
The genus Algyroides Bibron and Bory de
Saint-Vincent, 1833 includes four distinct
Mediterranean species of lizards found along
the eastern coasts of the Adriatic and Ionian
seas (Algyroides nigropunctatus; Duméril and
Bibron, 1839), Peloponnesus and nearby is-
lands (Algyroides moreoticus Bibron and Bory
de Saint-Vincens, 1833), the Tyrrhenian Islands
(Algyroides fitzingeri; Wiegmann, 1834) and the
northern Betic mountains of the southeastern
Iberian Peninsula (Algyroides marchi Valverde,
1958). These Algyroides species are grouped in
two morphologically distinct clades: a western
clade formed by A. fitzingeri and A. marchi and
an eastern clade formed by A. moreoticus and
1 - Museo Nacional de Ciencias Naturales, MNCN-CSIC,
c/José Gutiérrez Abascal 2, 28006 Madrid, Spain
2 - Department of Ecology, Universidad Autónoma de
Madrid, Campus de Cantoblanco, 28049 Madrid, Spain
∗Corresponding author;
e-mail: albertosv@mncn.csic.es
A. nigropunctatus (Arnold et al., 2007; Pavlicev
and Mayer, 2009). Previous molecular studies
of the genus did not support the monophyletic
status of this genus as Dinarolacerta was in-
cluded within Algyroides (Pavlicev and Mayer,
2009; Pyron et al., 2013). However, in a re-
cent detailed molecular analysis, Mendes et al.
(2016) recovered Algyroides as a monophyletic
group, which is congruent with morphological
studies (Arnold et al., 2007; Arribas, 2012).
In 1916, Eduardo Boscá, one of the most in-
fluent Spanish herpetologists of his time (Fraga
Vázquez, 1989; Catalá Gorgues, 2004; Al-
baladejo et al., 2013), first noted the presence
of Algyroides in the Iberian Peninsula. In his
work, Boscá (1916a) provided a diagnosis and
a detailed description of an Iberian Algyroides
based on a juvenile specimen he received from
San Ildefonso, a village supposedly located in
the Sierra de Guadarrama mountain range in
the province of Segovia (central Spain) (Boscá,
©Koninklijke Brill NV, Leiden, 2018. DOI:10.1163/15685381-20181016

2A. Sánchez-Vialas et al.
1916a). Boscá named this species Algiroides hi-
dalgoi Boscá, 1916 (sic, note the Latin “i” in-
stead of “y” in the genus name), stating that
it was closely related to A. fitzingeri. Unfortu-
nately, this specimen, now lost, was not illus-
trated and therefore, detailed drawings of it do
not exist.
No other specimens of Algyroides were re-
ported in the Iberian Peninsula until José An-
tonio Valverde discovered several individuals in
April 1958 in Piedra de Aguamula, in the Sierra
de Cazorla mountain range in the province of
Jaén (southeastern Spain) (Valverde, 1958), a
location far (about 320 km by air) from the
Sierra de Guadarrama. Valverde described the
Cazorla specimens as a new species, Algiroides
marchi Valverde, 1958 (sic, see above), that
according to him was morphologically distin-
guishable from A. hidalgoi. After Valverde’s
discovery, several authors tried to find A. hi-
dalgoi in the Sierra de Guadarrama, but all
attempts were unsuccessful (Klemmer, 1960;
García-París et al., 1989: 191-192), leading
some authors to question the validity of A.
hidalgoi (Klemmer, 1960; Mertens and Wer-
muth, 1960). However, other authors have con-
sidered A. hidalgoi a valid taxon (Valverde,
1958; Buchholz, 1964; Salvador, 1974; García-
París et al., 1989; Engelmann et al., 1993;
Pérez-Mellado, 1998; Fernández-Cardenete and
García-Cardenete, 2014).
The question of whether A. hidalgoi is a dis-
tinct species from A. marchi remains unsolved.
Some authors have treated it as a nomen du-
bium (Klemmer, 1960), while other have ques-
tionably included the name A. hidalgoi under
the synonymy of A. marchi (Mertens and Wer-
muth, 1960; Salvador, 1998; Alonso-Zarazaga,
2014). Unfortunately, the unique specimen of
A. hidalgoi is missing: it is not in Boscá’s per-
sonal collection in Valencia (Klemmer, 1960),
which was destroyed (Geniez et al., 2014)
nor at the Museo Nacional de Ciencias Nat-
urales (MNCN), which possesses some of
Boscá’s other type material (Pérez-Mellado,
1998; García-Díez and González-Fernández,
2013). Fortunately, Boscá’s description is very
detailed and complete (Valverde, 1958; Sal-
vador, 1974; Pérez-Mellado, 1998) and can be
compared with large series of A. marchi speci-
mens collected from different geographic areas
available from several Spanish scientific collec-
tions, and also with the remaining species of Al-
gyroides.
The problem is complex: first, whether A. hi-
dalgoi and A. marchi represent two different
taxonomic entities or a single one needs to be
determined, and second, the nomenclatural va-
lidity and status of A. hidalgoi needs to be clar-
ified. We address both issues by (1) compar-
ing Boscá’s original description of A. hidalgoi
against (a) the descriptions and preserved spec-
imens of all extant species of Algyroides, and
(b) 204 Iberian Algyroides specimens to deter-
mine if the enigmatic morphology of A. hidal-
goi can be recognized in any of them, and if so,
(2) by designating a neotype of A. hidalgoi that
corresponds to the head pholidosis described in
Boscá’s original description.
Material and methods
We studied preserved specimens of all species of Algy-
roides, plus a series of 198 specimens of Iberian Algy-
roides encompassing most of the described variability of the
species, collected from several localities and currently held
at the Museo Nacional de Ciencias Naturales (MNCN) (see
supplementary appendix S1). We also examined one para-
type of A. marchi (MNCN 7947). All other specimens of
the type series are currently located at the collection of the
Estación Biológica de Doñana (EBD-CSIC), and not, as
stated in Valverde (1958), in the collection of the Instituto
de Aclimatación de Almería (now Estación Experimental de
Zonas Áridas – EEZA-CSIC). All specimens are preserved
in 70% ethanol. Morphological characters were studied us-
ing a binocular microscope. Five quantitative morphological
measurements were taken using a digital caliper. Digital
photographs were taken with a reflex camera fitted with a
macro lens. For comparisons among Iberian specimens we
paid special attention to the characters used by both Boscá
(1916a) and Valverde (1958). These characters are as fol-
lows: (1) parietal shields separated or in contact; (2) occipi-
tal and interparietal shields separated or in contact; (3) posi-
tion of the tympanic scale: located directly over the upper
border of the tympanum opening (“transverse” following
cited authors) or at the upper-anterior corner of the tympa-
num opening (“oblique” op. cit.); (4) position of the mental
scale in relation to the rostral scale: both scales at the same

Taxonomy and nomenclature of Iberian Algyroides (Lacertidae) 3
vertical level or the rostral in a forward position; (5) ventral
coloration (bright yellow or bluish-grey); (6) shape and rel-
ative size of the dorsal and lateral trunk scales; (7) presence
of a central keel in the trunk scales; (8) shape of the scales in
the upper limb areas of the flanks; (9) shape of the external
scales of the hind limbs.
Results
Comparative diagnosis of A. hidalgoi
Boscá’s original description of A. hidalgoi reads
as follows: “cabeza pequeña, aplanada hacia
adelante y coincidiendo en el mismo punto
el extremo del hocico con el mentón. Escudo
occipital mínimo, triangular redondeado, sep-
arado del interparietal por los escudos pari-
etales que se tocan. Ojos grandes con cua-
tro escudos supralabiales anteriores al infra-
orbitario, que es rectangular y tan largo como
el diámetro transverso del ojo. Región parietal
con escudetes desiguales, existiendo en el lado
derecho uno que es mayor, representando el es-
cudo masetérico. Orificio auditivo grande, oval,
con la membrana timpánica convexa, y sobre
el borde superior, el escudo timpánico grande
y en sentido horizontal. Pliegue angular man-
ifiesto, separando la región en una parte ante-
rior en la que las escamas son alargadas y bas-
tante iguales entre sí, y otra parte posterior, en la
que las escamas son redondeadas, pequeñas las
de las primeras filas, aumentando gradualmente
de tamaño, estando limitadas por siete escamas
rectangulares, empizarradas y alternas sobre las
escamas que forman el collar, en número de
cuatro, que son algo mayores y con el borde
posterior libre. . . Tronco protegido por escamas
grandes, rombales, empizarradas, con una quilla
que termina en punta aguda, dispuestas en 17 fi-
las hacia la mitad del largo del cuerpo, siendo
de igual tamaño; sobre los lados y región su-
perior de los miembros aparecen algo pequeñas
y de forma más suave. Región inferior bril-
lante con 11 escudetes pectorales, de forma y
tamaño desiguales, yuxtapuestos siguiendo en
esa misma disposición seis filas de escudos ven-
trales, siendo más estrechas las de los de los la-
dos y las 2 del centro. Escudo anal hexágono,
bordeado en su parte anterior por otros 7 escu-
dos, de los que es mayor el del centro, siguiendo
a la abertura del ano tres filas de escamas granu-
jientas. La cola como uno y un quinto de la lon-
gitud del resto del animal, gruesa y cilindrácea
en su mitad anterior, terminando en punta afi-
lada, y el saliente de la quilla de sus escamas
forman una serie de verticilos ostensibles a sim-
ple vista. Patas posteriores extendidas hacia de-
lante sobre los flancos no alcanzan la axila, y
lo mismo que las extremidades anteriores, están
cubiertas por su cara inferior por escamas lisas
abrillantadas. Poros femorales en número de 9 a
cada lado; dedos finos, con una fila de escamas
gruesas en el borde de abajo, provistos de uña
corta y ganchuda. El color café acompaña in-
distintamente a las partes superiores, viéndose
machitas oscuras sobre los escudos cefálicos,
así como sobre las escamas lisas que protegen
las extremidades, pero en éstas las manchas son
mayores, redondeadas y distribuidas con cierto
orden. Partes inferiores de un gris azulado, tam-
bién uniforme, excepto sobre la cola, que lleva
el mismo color café toda ella. Longitud total, 59
mm.; cabeza, 6; cuello, 3; tronco, 18; cola, 32;
mayor anchura sobre el abdomen, 5 mm.”
This description differs in many traits from
all species of Algyroides with the exception
of A. marchi. For example, Algyroides hidal-
goi differs markedly from A. fitzingeri and A.
moreoticus because they present similar shaped
scales in the lateral and the dorsal areas of
the central portion of the trunk (markedly dif-
ferent in A. hidalgoi) (see for example Spey-
broek et al., 2016); and have highly keeled lat-
eral trunk scales, along the area comprised be-
tween the limbs (not keeled in A. hidalgoi). The
flank scales from the area located directly above
the limbs are small, rounded and un-keeled in
all species of Algyroides, quite similar to the
lateral trunk scales described for A. hidalgoi.
There are marked differences in the shape of the

4A. Sánchez-Vialas et al.
limb scales; while keeled in A. fitzingeri and A.
moreoticus, they are smooth in A. hidalgoi.
Algyroides hidalgoi differs in many traits
from the two subspecies of A. nigropunctatus
(A. n. nigropunctatus and A. n. kephallithacius;
Keymar, 1986) but the most obvious characters
are that they have lateral scales and hind limb
external (inferior) scales markedly keeled (of-
ten asymmetrically keeled), while they are not
keeled in A. hidalgoi (“Patas posteriores. . . cu-
biertas por su cara inferior por escamas lisas
abrillantadas...” “... así como de las escamas
lisas que protegen las extemidades,...”).
The most similar species to A. hidalgoi is
A. marchi, which is the only species of Algy-
roides that presents lateral scales small and not-
keeled, and limb scales smooth and polished.
Algyroides marchi niethammeri was described
as subspecies of A. marchi by Buchholz (1964),
based on coloration of the throat and differences
in the number of dorsal scales. Algyroides m.
niethammeri was synonymized with typical A.
marchi by Palacios et al. (1974), after the study
of several specimens from its type locality: Bog-
arra (Albacete).
Most of the quantitative and qualitative char-
acters indicated in Boscá’s description of A. hi-
dalgoi are represented across the highly variable
species A. marchi. Valverde (1958) described
A. marchi, by directly comparing Boscá’s de-
scription with his newly collected materials.
In the following paragraphs we discuss one
by one those characters in a separate section.
In addition, other authors (Buchholz, 1964)
and colleagues (in litteris) added a few more
characters, not discussed by Valverde, but that
could be used to separate A. marchi from A.
hidalgoi. These characters are discussed be-
low.
One of these characters is the number of rows
of dorsal scales. Boscá’s (1916a) stated that
A. hidalgoi has 17 rows of dorsal scales (only
dorsal, and not total as indicated by Buchholz,
1964: 242), while A. marchi usually present
9 dorsal rows. This difference, although obvi-
ously clear, might however by just a conse-
quence of the counting process: in many spec-
imens of A. marchi dorsal scales are not ar-
ranged in a clear linear position, especially
along the broadest part of the dorsum (fig. 1A).
If Boscá counted scales, through an inverted V
line across dorsum (see numbers in fig. 1A), the
count will be between 16 to 18, including there-
fore the number indicated by Boscá (1916a). If
differences in the way of counting scales are
responsible for the differences described, this
character cannot be considered a diagnostic trait
for A. hidalgoi vs. A. marchi.
Another character used to separate A. marchi
from A. hidalgoi is the number of femoral pores,
stated as 9 in Boscá’s description of A. hi-
dalgoi, while represented by 11 to 16 in A.
marchi (Pérez-Mellado, 1998). Femoral pores
in young specimens of A. marchi are some-
times difficult to count. In fact, two young
specimens from the collection of the MNCN
present 9 poorly marked femoral pores in at
least one leg. So, if Boscá’s specimen was a ju-
venile, as it seems to be based on its size (59
mm length, tail included), it seems quite pos-
sible that the 9 femoral pores observed corre-
spond to the 9 pores sometimes observed in ju-
venile A. marchi. Consequently, this character
cannot be considered diagnostic for A. hidal-
goi vs. A. marchi comparisons either. A third
character considered distinctive of A. hidalgoi
vs. A. marchi is the dorsal/lateral contrasting
coloration shown by the latter, versus a dark
brown uniform coloration in the former. How-
ever, once again, the coloration in preservative
fades through time depending upon the qual-
ity and timing of the actual preservation. In the
MNCN collection, many old specimens of A.
marchi present the typical coloration of the in-
dividual described by Boscá (1916a) (fig. 1B),
making this character invalid for any diagnos-
tic usage between live and preserved speci-
mens.

Taxonomy and nomenclature of Iberian Algyroides (Lacertidae) 5
Figure 1. A. Dorsal row of scales fitting the number counted by Boscá’s description. B. Homogeneous dorsal body coloration
after years of fluid preservation.
There is a somewhat ambiguous phrase in
Boscá’s description: “Tronco protegido por es-
camas grandes, . . . igual tamaño; sobre los la-
dos y región superior de los miembros apare-
cen algo pequeñas y de forma más suave”. The
last portion of this sentence “on the sides and
upper region of the limbs they appeared some-
what smaller and of smoother shape” (transla-
tion ours) could be interpreted as if the species
presents the scales over the limbs smaller and
smooth, a character that would not be present
in any other species of Algyroides. However, we
are convinced that in this sentence Boscá refers
to the trunk scales located above the limbs (not
on the limbs), which are small and rounded in
all species of Algyroides, including A. marchi.
We believe this is the case because Boscá, as any
other trained taxonomist, will describe sepa-
rately the characteristics of the trunk from those
of the limbs, never mixing them in the same
sentence. In fact, Boscá described the scales on
the limbs a few sentences below: “Patas poste-
riores. . . cubiertas por su cara inferior por esca-
mas lisas abrillantadas...” “...así como sobre
las escamas lisas que protegen las extremidades,
...”
Morphological comparisons using Valverde’s
(1958) criteria
We found high morphological variability in
pholidosis, even among individuals of the same
population (as previously reported by Palacios
et al., 1974). Five morphological characters
were discussed in the original descriptions of
both A. hidalgoi and A. marchi (Boscá, 1916a;

6A. Sánchez-Vialas et al.
Figure 2. Dorsal variability in head scales (a red dot is placed on the left parietal scale). A-C represents parietals in contact
as noted by Boscá (1916a). D-F shows individuals with parietals separated as described by Valverde (1958). E represent a
distinctive individual with parietal plates fragmented, note also that its internasal is divided in two plates. Figure F shows
the interparietal fragmented. A: MNCN 32020; B: MNCN 32125; C: MNCN 32082; D: MNCN 32058; E: MNCN 32144; F:
MNCN 31976.
Valverde, 1958). Variation of characters 1 and
2 always follows a specific pattern: 8 speci-
mens (of the 198 examined) have the parietal
shields in contact and the occipital and inter-
parietal shields separated, as in the description
of A. hidalgoi (Boscá, 1916a) (fig. 2), while the
other 190 show separated parietals and occipital
and interparietal in contact, as in the description
of A. marchi (Valverde, 1958) (fig. 2). Charac-
ter 3, the tympanic scale, is variable in shape
and position (fig. 3). Of the 198 specimens, 53
present the scale over the upper edge of the tym-
panum opening as described by Boscá (1916a)
(fig. 3) and 145 show it at the upper-anterior
corner of the tympanic opening as recorded by
Valverde (1958) (fig. 3). Although the tympanic
scale is occasionally broken into smaller pieces,
its position can be unambiguously determined
(fig. 3). Although character 4, the mental scale,
is easily recorded, it is based on a moveable

Taxonomy and nomenclature of Iberian Algyroides (Lacertidae) 7
Figure 3. Lateral view of the head: scales variability. White lines point to tympanic scale (A-F) and masseteric plate (D).
A-B: Tympanic scale horizontally positioned as described by Boscá (1916a). C-E: Tympanic scale positioned as described by
Valverde (1958). F: Tympanic scale fragmented. Note the presence of masseteric plate in specimen of D. A: MNCN 32054;
B: 32074; C: 32058; D: 32139; E: 32013; F: 32098.
structure (lower jaw position with respect to the
upper jaw), so its value is highly relative: 41 of
the 198 specimens have the mental and rostral
scales at the same vertical level, as in Boscá’s
description (fig. 3), while the other 157 spec-
imens show the pattern described by Valverde
(fig. 3). With respect to character 5, ventral col-
oration, all specimens studied (which were pre-
viously preserved in ethanol), including a pa-
ratype of Valverde’s A. marchi (MNCN 7947),
present a bluish or, more rarely, brownish ven-
tral coloration as indicated by Boscá (1916a)
(fig. 4).
Discussion
We have determined that none of the five di-
agnostic characters used by Valverde (1958) to
differentiate between A. hidalgoi and A. marchi
are actually diagnostic. Four characters (all re-
lated to scales) are highly variable within popu-
lations, and both character states (Boscá’s and
Valverde’s) are observed within the same popu-
lations (e.g. Peal de Becerro, Cazorla, Riópar
and Bogarra). The fifth character, ventral col-
oration, is uniform in all specimens examined,
including material from the type series of A.
marchi. However, Valverde (1958) correctly in-
dicated the ventral color as “amarillo canario

8A. Sánchez-Vialas et al.
Figure 4. Differences in ventral coloration between pre-
served and living specimens of Iberian Algyroides.Ashows
the ventral grey-bluish coloration of a preserved specimen
(MNCN 32144) as described by Boscá (1916a). B depicts
the canary yellow ventral coloration of a live individual
from Río Mundo (Albacete) (not collected).
brillante” (“bright canary yellow”) as this is the
color observed in live specimens. The yellow
color faded to bluish-grey in the type mate-
rial of A. marchi and in other preserved Al-
gyroides specimens. Therefore, the bluish-grey
ventral coloration described by Boscá, and also
observed for Valverde’s specimens, is an artifact
of preservation in ethanol.
Valverde (1958) stated that “Esto parece in-
dicar que se trata de una especie diferente o
bien que la descripción, basada sobre un joven,
es errónea” (it seems to indicate that [Boscá’s
specimen] represent a different species or oth-
erwise, that its description, based on a juvenile,
is erroneous). Based on our morphological anal-
ysis, both assumptions seem incorrect: Boscá’s
description was accurate, and it indicates in-
traspecific variability rather than specific dif-
ferentiation. Valverde (1958) only studied five
specimens, and likely did not consider the pos-
sibility of large population variability. Addition-
ally, none of the characters suggested by other
authors (Buchholz, 1964) or our colleagues to
differentiate between A. marchi and A. hidal-
goi are diagnostic, in fact most of them possi-
bly correspond to erroneous interpretations of
Boscá’s writing.
Based on the evidence provided here, we
demonstrate that the specimens used to describe
both A. hidalgoi and A. marchi represent a sin-
gle taxonomic and evolutionary unit.
Nomenclature of Iberian Algyroides
Klemmer (1960), Buchholz (1964) and Pérez-
Mellado (1998) have commented on the nomen-
clatural problems associated with the name A.
hidalgoi. Klemmer (1960) suggested that the
name A. hidalgoi was a nomen dubium:“so
läßt sich doch hidalgoi nach der unzureichen-
den Beschreibung nicht mit der notigen Exak-
theit deuten, so dag dieser Name als ein Nomen
dubium einstweiIen ruhen muß” (hidalgoi, ac-
cording to the inadequate description, cannot be
interpreted with the necessary exactness, so that
this name must rest, as a Nomen dubium,forthe
time being).
Buchholz (1964: 246) argued in favor of the
validity of the name A. hidalgoi: “1. Der Name
Algyroides hidalgoi kann nicht als nomen du-
bium aufgefaßt werden. Nach der detaillierten
Beschreibung von Boscá ist das Taxon ohne
weiteres identifizierbar.” (The name Algyroides
hidalgoi cannot be treated as a nomen dubium.
According to the detailed description of Boscá,
the taxon is readily identifiable.).
We agree with Buchholz (1964) as the de-
scription of A. hidalgoi is complete and can-
not be confused with any other Algyroides
species. However, the type of A. hidalgoi is
lost so cannot be used to inform on this is-
sue. In 1914, Boscá’s type collection was in-
corporated into the collections at the MNCN in
Madrid (García-Díez and González-Fernández,
2013). Notably though, A. hidalgoi was de-
scribed after this event. In 1958, Francisco
Bernis, then head of the MNCN vertebrate
collections, searched for but was unable to
find the type of A. hidalgoi in the collections

Taxonomy and nomenclature of Iberian Algyroides (Lacertidae) 9
(Valverde, 1958). Klemmer (1960) tried to lo-
cate the specimen at Boscá’s work place in
Valencia but was also unsuccessful. The type
has been considered lost since then (García-
Díez and González-Fernández, 2013; M. Calvo
herpetological collection data). Recently, Ge-
niez et al. (2014) indicated that Bosca´s con-
temporary types of Podarcis guadarramae were
destroyed. Boscá probably sent specimens of
his collection, in exchange or for discussion,
to contemporary specialists. Fernand Lataste,
George Albert Boulenger, and Jacques von
Bedriaga, described new taxa using Boscá’s ma-
terials, discussed his identifications, or were
recognized with new taxa dedicated to them
(see for example Lataste, 1879; Boscá, 1880;
Boulenger, 1919), so it could have been pos-
sible that any of them received the holo-
type of A. hidalgoi for examination. The col-
lections of all these three herpetologists are
held at the Natural History Museum (Lon-
don), which do not include any specimen that
could be associated with Algyroides hidalgoi
(http://www.nhm.ac.uk/our-science/collections/
zoology-collections/herpetology-collections.
html).
As discussed in the previous section, Iberian
Algyroides are represented by a single taxo-
nomic unit; however, two valid names are avail-
able for this unit: A. hidalgoi and A. marchi.
In order to address the two-name nomenclatural
problem, exacerbated by the loss of the A. hidal-
goi type material, and to avoid future taxonomic
problems and uncertainties, we deem necessary
to designate a neotype for A. hidalgoi (ICZN,
2000: Article 75.3).
Designation of neotype for Algyroides hidalgoi
An adult specimen (MNCN 32128) mostly con-
sistent with Boscá’s original description, in-
cluding parietal scales in contact and occipi-
tal and interparietal scales separated, was se-
lected as the neotype (fig. 4). The locality of
the selected neotype (Los Rasos, Peal de Be-
cerro) is relatively far from the one indicated by
Boscá’s. However, Boscá’s locality was prob-
ably erroneous or the consequence of acci-
dental transport with plants or other materials
to the San Ildefonso Royal Gardens (O. Ar-
ribas in http://www.montesdevalsain.es/index_
reptiles.html; consulted: 21/2/2017), making it
impossible to select a neotype from that local-
ity. Boscá (1916b) described Lacerta muralis
guadarramae (=Podarcis guadarramae)also
from specimens, he did not collect, from San
Ildefonso. He found these materials at the col-
lections of the Botanical Garden of the Univer-
sity of Valencia (Boscá, 1916b; Geniez et al.,
2014). The description of the coloration pattern
of L. m. guadarramae, described as presenting
six longitudinal stripes, fits precisely with the
coloration pattern of the specimens of Podar-
cis from Sierra de Cazorla (the area selected for
the neotype of A. hidalgoi), and not at all with
the coloration pattern of female specimens from
the Guadarrama mountains (always presenting
less than six stripes). This additional observa-
tion supports either an erroneous labelling of
both Boscá’s Podarcis and Algyroides,ormore
likely the existence of a local toponym named
“San Ildefonso” in the mountains of southeast-
ern Spain, then and now forgotten. The name
“San Ildefonso” is a common one all over south-
ern Spain and, for example, it is currently held
by a chapel and a “cortijo” in the province of
Jaén (Google search: “San Ildefonso, Jaén”).
The mentioned “Cortijo of San Ildefonso” is lo-
cated in Montizón, a village relatively close to
Sierra de Cazorla.
Description of neotype: adult male (fig. 5);
SVL 42.86 mm; total length 78.94 mm; belly
color bluish (fig. 5B); 24 femoral pores on both
hind limbs; head length (snout-collar) 15.81
mm; head width: 7.08 mm; anal scale width
3.13 mm; 6 ventral scale rows; 8 collar scales.
Masseteric plate large. Tail regenerated.
The neotype, MNCN 32128, which bears the
labels “Los Rasos-Cazorla-Cauce del
Guadalquivir” “7.6.1981, 13h, 1.6 gr” // “7539”
[Fernando Palacios leg.] // Neotypus des. by

10 A. Sánchez-Vialas et al.
Figure 5. Habitus of the neotype of Algyroides hidalgoi (MNCN 32128) (Los Rasos, Peal de Becerro, Jaén). A: dorsal view;
B: ventral view.
Sánchez-Vialas, Calvo-Revuelta and García-
París, 2018 //, is held at the MNCN Herpetolog-
ical Collection and is preserved in 70% ethanol.
Variability of Cazorlan populations has been
previously described in detail by Palacios et
al. (1974); variability, including extreme val-
ues for key characters, can be further evalu-
ated based on the following specimens from
Los Rasos-Cazorla: MNCN 32125: male, SVL
44.59 mm; total length 127.16 mm; 26 femoral
pores on both hind limbs; head length (snout-
collar): 17.31 mm; head width: 7.84 mm; anal
scale width: 3.34 mm; 6 ventral scale rows; 9
collar scales; masseteric plate present. MNCN
32126: female, SVL 45.67 mm; head length
(snout-collar): 14.70 mm; head width: 6.79 mm;
anal scale width: 2.54 mm; 6 ventral scale rows;
9 collar scales; no masseteric plate; both parietal
shields fragmented into two plates situated on
both sides of the occipital scale. MNCN 32127:
female, SVL 46.81 mm; head length (snout-
collar): 15.18 mm; head width: 6.68 mm; anal
scale width: 2.63 mm; 6 ventral scale rows; 8
collar scales; no masseteric plate.
By designating a neotype for A. hidalgoi,us-
ing a specimen collected close to the type lo-
cality of A. marchi, we definitively and unam-
biguously solve both the taxonomic and the
nomenclatural problems, resulting in A. hidal-
goi and A. marchi becoming “objective” syn-
onyms. Following the “precedence” rule of the
ICZN (2000: 23.1), the name Algyroides hidal-
goi must be retained over Algyroides marchi,
which becomes a junior synonym of A. hidal-
goi.
The conservation of the widely used name A.
marchi is problematic. Reversal of precedence
for A. marchi is not immediate. ICZN (2000:
23.9.1.1) establishes that reversion of prece-
dence cannot be applied when the oldest name
was published after 1899. The name A. hidal-
goi was used for the first time in 1916 and has
been used repeatedly in recent times (Buchholz,
1964; Salvador, 1974; García-París et al., 1989;
Engelmann et al., 1993, among others) and con-
sequently establishing precedence of A. marchi
over A. hidalgoi would require an application to
the ICZN.

Taxonomy and nomenclature of Iberian Algyroides (Lacertidae) 11
Synonymic list
Algyroides hidalgoi Boscá, 1916
Algiroides hidalgoi Boscá, 1916a: 295. Terra
typica: “San Ildefonso (Segovia)”
Algiroides marchi Valverde, 1958: 127. Terra
typica: “Piedra de Aguamula, en la Sierra de
Cazorla (Jaén)”. syn. nov.
Algyroides marchi niethammeri Buchholz 1964:
244. Terra typica: “Sierra de Agua (Jaén),
SO-Spanien, nahe dem Paß (1480 m) an
der Straße Alcaráz-Riópar, bei ca. 1430 m”.
Synonymy with A. marchi established by
Palacios et al. (1974).
Boscá (1916a) and Valverde (1958) used the
name Algiroides Duméril and Bibron, 1839 in-
stead of Algyroides Bibron and Bory de Saint-
Vincent, 1833. Algiroides is an incorrect subse-
quent spelling of Algyroides and, according to
the code, incorrect spellings or emendations at
the genus level do not have to be treated as new
combinations (ICNZ 2000: 51.3.1). Following
this logic, the combinations recorded in the syn-
onymic list of Salvador (1997, 2014) based on
incorrect subsequent spellings were removed
from the list presented here. Boulenger (1920:
339) misspelled A. hidalgoi as “Algiroides hi-
dalgi”. Boulenger (1921) also proposed erro-
neously that A. hidalgoi was a synonym of
Ophisops occidentalis Boulenger, 1887. How-
ever, it seems clear that Boulenger did not study
the original specimen, and its attribution was
speculative.
Acknowledgements. We thank Fernando Palacios for his
help with the labelling data of MNCN specimens; Ernesto
Recuero and especially Oscar Arribas for their extremely
valuable comments on an earlier version of the manuscript;
an anonymous reviewer by his critical comments that really
improved the manuscript. We are grateful to Judit Vörös,
curator of the Magyar Természettudományi Múzeum (Bu-
dapest) for providing Algyroides specimens and her wel-
come at the Museum. We thank Melinda Modrell for a thor-
ough language revision. Also, we are grateful to Miguel Án-
gel Alonso-Zarazaga for his help with some nomenclatural
issues. This study was supported by grant CGL2015-66571-
P (MINECO-FEDER).
References
Albaladejo, C.M., Prieto, R.M.G., Sanchiz, B. (2013): Dic-
tamen de Mariano de la Paz Graells sobre la obra her-
petológica de Eduardo Boscá: análisis de una evaluación
científica del siglo XIX. Asclepio. 65 (2): p023.
Alonso-Zarazaga, M.A. (2014): Algiroides hidalgoi Boscá,
1916. In: Fauna Ibérica, Vol. 10. Reptiles, 2aedición
aumentada y corregida, p. 1339-1340. Salvador, A., Ed.,
Museo Nacional de Ciencias Naturales, CSIC, Madrid.
Arnold, E.N., Arribas, O., Carranza, S. (2007): Systematics
of the Palaearctic and Oriental lizard tribe Lacertini
(Squamata: Lacertidae: Lacertinae), with descriptions of
eight new genera. Zootaxa. 1430: 1-86.
Arribas, O.J. (2012): Hemipenial microornamentation in
Algyroides marchi Valverde, 1958 (Squamata: Sauria:
Lacertidae). Herpetozoa. 25 (1-2): 21-24.
Boscá, E. (1880): Gongylus bedriagai, nueva sub-especie de
la Península Ibérica. An. Soc. Esp. Hist. Nat. 9: 495-503.
Boscá, E. (1916a): Un género nuevo para la fauna her-
petológica de España y especie nueva o poco conocida.
Bol. R. Soc. Esp. Hist. Nat. 16: 294-297.
Boscá, E. (1916b): Dos observaciones a propósito de Lac-
erta muralis en España. Bol. R. Soc. Esp. Hist. Nat. 16:
327-330.
Boulenger, A.G. (1919): Sur le Pelobates Wilsoni, Boscá.
Bol. R. Soc. Esp. Hist. Nat. 19: 293-294.
Boulenger, G.A. (1920): Monograph of the Lacertidae,
Vol. I. British Museum of Natural History, London.
Boulenger, G.A. (1921): Monograph of the Lacertidae.
Vol. 2. British Museum of Natural History, London.
Buchholz, K.F. (1964): Zur Kenntnis des Genus Algyroides
(Reptilia: Lacertidae) in Spanien. Bonn. zool. Beitr. 15
(3/4): 239-246.
Catalá Gorgues, J.I. (2004): El desarrollo de una carrera
científica en un contexto institucional precario: el caso
del naturalista Eduardo Boscá y Casanoves (1843-1924).
Cronos. 7(1): 3-60.
Engelmann, W.E., Scholz, J. (1993): Lurche und Kriechtiere
Europas. Neumann, Radebeu.
Fernández-Cardenete, J.R., García-Cardenete, L. (2014):
Algyroides marchi (Valverde, 1958). In: Fauna Ibérica,
Vol. 10. Reptiles, 2aedición revisada y aumentada,
p. 368-381. Salvador, A., Ed., Museo Nacional de Cien-
cias Naturales, CSIC, Madrid.
Fraga Vázquez, X.A. (1989): A modernización da Tax-
onomía herpetológica a fins do XIX no Estado español:
as aportacións de Boscá e López Seoane. Treb. Soc. Cat.
Ictiol. Herpet. 2: 26-43.
García-Díez, T., González-Fernández, J.E. (2013): The rep-
tile type specimens preserved in the Museo Nacional de
Ciencias Naturales (CSIC) of Madrid, Spain. Zootaxa.
3619 (1): 46-58.
García-París, M., Martín, C., Dorda, J., Esteban, M. (1989):
Los anfibios y reptiles de Madrid. Ministerio de Agricul-
tura, Pesca y Alimentación, Madrid.
Geniez, P., Sa-Sousa, P., Guillaume, C.P., Cluchier, A., Cro-
chet, P.A. (2014): Systematics of the Podarcis hispani-
cus complex (Sauria, Lacertidae) III: valid nomina of the
western and central Iberian forms. Zootaxa. 3794 (1):
1-51.

12 A. Sánchez-Vialas et al.
Klemmer, K. (1960): Zur Kenntnis der Gattung Algyroides
(Rept., Lacertidae) auf der Iberischen Halbinsel. Senck-
enb. Biol. 41 (1/2): 1-7.
Lataste, F. (1879): Sur une nouvelle forme de Batracien
anoure d’Europe (Alytes obstetricans Boscai, n. subsp.).
Rev. Int. Sci. 4: 543-545.
Mendes, J., Harris, D.J., Carranza, S., Salvi, D. (2016):
Evaluating the phylogenetic signal limit from mi-
togenomes, slow evolving nuclear genes, and the con-
catenation approach. New insights into the Lacertini ra-
diation using fast evolving nuclear genes and species
trees. Mol. Phylogenet. Evol. 100: 254-267.
Mertens, R., Wermuth, H. (1960): Die Amphibien und
Reptilien Europas. Waldemar Kramer Verlag, Frankfurt
am Main.
Palacios, F., Ayarzagüena, J., Ibáñez, C., Escudero, J.
(1974): Estudio sobre la lagartija de Valverde Algyroides
marchi (Reptilia, Lacertidae). Doñana Act. Vert. 1(2):
5-31.
Pavlicev, M., Mayer, W. (2009): Fast radiation of the sub-
family Lacertinae (Reptilia: Lacertidae): history or me-
thodical artefact? Mol. Phylogenet. Evol. 52 (3): 727-
734.
Pérez-Mellado, V. (1998): Algyroides marchi (Valverde,
1958). In: Fauna Ibérica, Vol. 10. Reptiles, p. 175-
181. Salvador, A., Ed., Museo Nacional de Ciencias
Naturales, CSIC, Madrid.
Pyron, R.A., Burbrink, F.T., Wiens, J.J. (2013): A phy-
logeny and revised classification of Squamata, including
4161 species of lizards and snakes. BMC Evol. Biol. 13
(1): 1.
Salvador, A. (1974): Guía de los anfibios y reptiles es-
pañoles. Instituto Nacional para la Conservación de la
Naturaleza, Madrid.
Salvador, A., Ed. (1998): Fauna Ibérica, Vol. 10. Reptiles.
Museo Nacional de Ciencias Naturales, CSIC, Madrid.
Salvador, A., Ed. (2014): Fauna Ibérica, Vol. 10. Reptiles,
2aedición revisada y aumentada. Museo Nacional de
Ciencias Naturales, CSIC, Madrid.
Speybroeck, J., Beukema, W., Bok, B., Van Der Voort, J.
(2016): Field Guide to the Amphibians and Reptiles of
Britain and Europe. Bloomsbury Publishing, London.
Valverde, J.A. (1958): Una nueva lagartija del género Al-
giroides Bribron procedente de la Sierra de Cazorla (Sur
de España). Arch. Inst. Aclim. 7: 127-134.
Submitted: February 20, 2018. Final revision received:
August 14, 2018. Accepted: August 21, 2018.
Associate Editor: Miguel Angel Carretero.

1
Amphibia-Reptilia
Taxonomic and nomenclatural status of Iberian Algyroides (Lacertidae)
Alberto Sánchez-Vialas*, Marta Calvo-Revuelta, Mario García-París
Abstract. The taxonomy and nomenclature of Iberian Algyroides are problematic. The first taxon
described, A. hidalgoi Boscá, 1916, was based on a single specimen that was subsequently lost. The
description of the second taxon, A. marchi Valverde, 1958, was based on the comparison of a newly
discovered population with the original description of A. hidalgoi. However, A. hidalgoi specimens have
never been recorded since for any locality. Therefore, three questions need to be addressed: Is A. hidalgoi
Boscá, 1916 a morphologically diagnosable taxon different from all non-Iberian species of Algyroides?
are A. hidalgoi and A. marchi conspecific? And if so, which is the correct name for the species? To clarify
the taxonomic status of the Iberian Algyroides we (1) compare Boscá’s A. hidalgoi original description
against the descriptions of all other species of Algyroides, (2) test the accuracy of Boscá’s A. hidalgoi by
comparing it against 204 Iberian museum specimens, and (3) designate a neotype of A. hidalgoi that fits
the head pholidosis described in the original description. We show that none of the diagnostic characters
used by Valverde to differentiate between A. hidalgoi and A. marchi are actually diagnostic, as we found
high levels of variability on those characters in the studied specimens. Our results validate Boscá’s
description of A. hidalgoi, which fits within the morphological variability observed for southern Iberian
Algyroides. As a result, we propose the strict synonymy of A. marchi Valverde, 1958 with A. hidalgoi
Boscá, 1916.
Keywords: Algyroides hidalgoi, Algyroides marchi, morphological variability, neotype designation,
nomenclature, nov. synonymy, taxonomy.
Museo Nacional de Ciencias Naturales, MNCN-CSIC, c/ José Gutiérrez Abascal 2, 28006 Madrid,
España
*Corresponding author; e-mail: albertosv@mncn.csic.es
Supplementary material

2
Appendix S1: Material examined
Specimens of non-Iberian Algyroides examined (MNCN = Museo Nacional de Ciencias
Naturales, Madrid. HNHM = Hungarian Natural History Museum / Magyar
Természettudományi Múzeum, Budapest).
Algyroides nigropunctatus
Croatia: Dobrinj: Krk Island: 06/09/1965, 8 specimens, Fernbach J. (HNHM:
2011.211.1-2011.211.8, 65.153.1). Greece: Corfu: Messonghi: 1 specimen, 07/1993, Tartaly
Á. (HNHM: 2003.17.1); Perumia: 2 specimens, 10/1995, Tóth, T. (HNHM: 2011.89.1,
98.41.1). “Illiria”: Balkan peninsula: 07/1881, 1 specimen (MNCN 7960); “Illiria”: Balkan
peninsula: 1 specimen, 1881, Sr. Sohneiber (MNCN 7961).
Algyroides fitzingeri
Italy: Sardinia: Alghero: 1 specimen (HNHM: 62.295.1.).
Algyroides moreoticus
Greece: Kefalonia: Argostoli: ruins of Krane: 1 specimen (HNHM: 62.294.1.).
Specimens of Iberian Algyroides examined. MNCN = Museo Nacional de Ciencias Naturales
(Madrid).
Spain: Castilla la Mancha: Albacete: Riópar: Cueva de los Chorros del Río Mundo:
17/08/1975, 1 specimen (MNCN 7948); Riópar: Chorros del Río Mundo: 02/07/1978, 1
specimen, F. Palacios y J.L. Rubio (MNCN 9473); Riópar: Chorros del Río Mundo: 3
specimens (MNCN 14376-14378); Riópar: Chorros del Río Mundo: 23/04/1973, 3 specimens,
F. Palacios (MNCN 31537-31539); Riópar: Arroyo de la Celadilla: 15/08/1973, F. Palacios
(MNCN 31974); Riópar: Chorros del Río Mundo: 20/07/1982, 3 specimens, F. Palacios, J.
Gisbert, B. Elvira (MNCN 31975-31977); Riópar: Chorros del Río Mundo: 1 specimen, F.

3
Palacios (MNCN 31978); Riópar: Pollo del Fraile: 1 specimen, 23-08-1973 F. Palacios (MNCN
31979); Riópar: 1 specimen, 25-08-1973, F. Palacios (MNCN 31980); Riópar: Línea de Puertas
en el Calar: 1 specimen, 25-08-1973, F. Palacios (MNCN 31980); Riópar: 5 specimens, 27-04-
1973, F. Palacios (MNCN 31981- 31985); Riópar: Los Chorros del Río Mundo: 2 specimens,
18-07-1981, F. Palacios, J. Gisbert, B. Elvira (MNCN 31986-31987); Riópar: Los Chorros del
Río Mundo: 11 specimens, 24-07-1981, F. Palacios, J. Gisbert, B. Elvira (MNCN 31988-
31998); Riópar: Boquera del Calar: 1 specimen, 28-07-1973, F. Palacios (MNCN 31999);
Riópar: 1 specimen, 27-04-1973, F. Palacios (MNCN 32000); Riópar: Los Chorros del Río
Mundo, El Charco de las Truchas: 2 specimens, 27-08-1973, F. Palacios (MNCN 32001-
32002); Riópar: Los Chorros del Río Mundo: 1 specimen, 07-05-1973, F. Palacios (MNCN
32003); Riópar: Los Chorros del Río Mundo, El Charco de las Truchas: 4 specimens, 22-08-
1973, F. Palacios (MNCN 32004-32007); Riópar: 2 specimens, 09-07-1973, F. Palacios
(MNCN 32008-32009); Riópar: Los Chorros del Río Mundo: 6 specimens, 08-07-1973, F.
Palacios (MNCN 32010-32015); Riópar: Los Chorros del Río Mundo: 13 specimens, 10-07-
1973, F. Palacios (MNCN 32016-32028); Riópar: Los Chorros del Río Mundo: 3 specimens,
28-04-1973 F. Palacios (MNCN 32029-32031); Riópar: Los Chorros: 10 specimens, 07-07-
1981, F. Palacios, J. Gisbert, B. Elvira (MNCN 32032-32041); Riópar: Los Chorros del Río
Mundo: 33 specimens, 26-04-1973, F. Palacios (MNCN 32042-32074); Riópar: Los Chorros del
Río Mundo: 8 specimens, 25-08-1973, F. Palacios (MNCN 32075-32082); Riópar: Los Chorros
del Río Mundo: 3 specimens, 29-04-1973, F. Palacios (MNCN 32083-32085); Bogarra: El
Toril-Cuesta de las Carboneras: 7 specimens, 19-04-1974, F. Palacios y J. Escudero (MNCN
32086-32092); Riópar: Cantera en Sierra del Agua: 1 specimen, 12-08-1973, F. Palacios
(MNCN 32093); Bogarra: Cuesta de las Carboneras: 2 specimens, 25-07-1973, F. Palacios
(MNCN 32094-32095); Bogarra: Cuesta de las Carboneras: 10 specimens, 10-08-1973, F.
Palacios (MNCN 32096-32105); Riópar: Los Chorros del Río Mundo: 3 specimens, 05-08-
1978, F. Palacios (MNCN 32106-32108); Riópar: Nacimiento del Arroyo de la Celadilla: 4
specimens, 09-07-1973, F. Palacios (MNCN 32109-32112); Riópar: Los Chorros del Río
Mundo: El Charco de las Truchas: 6 specimens, 06-07-1973 F. Palacios (MNCN 32113-32118);

4
Bogarra: Cuesta de las Carboneras: 2 specimens, 08-06-1974, F. Palacios (MNCN 32119-
32120); Riópar: Los Chorros del Río Mundo: 1 specimen, 07-07-1981, F. Palacios, J. Gisbert,
B. Elvira (MNCN 39290). Andalucía: Jaén: Sierra de Cazorla, Peña de Aguamula, 1450 m, 1
specimen, juvenile, 19-04-1958, J. A. Valverde (MNCN 7947) (Paratype of Algiroides marchi
Valverde, 1958); Peal de Becerro: Los Rasos: 24-03-1970, 1 specimen, F. Palacios y J.
Escudero (MNCN 32121); Peal de Becerro: Campamento de los Rasos: 08-07-1981, 3
specimens, F. Palacios (MNCN 32122-32124); Peal de Becerro: Campamento de los Rasos: 07-
06-1981, 4 specimens, F. Palacios (MNCN 32125-32127, 32129); Los Rasos, Cazorla, Cauce
del Guadalquivir: 07-06-1981, 13 h, 1.6 gr., 1 specimen, Fernando Palacios (MNCN 32128)
(Neotype of Algiroides hidalgoi Boscá, 1916); Cazorla: Parador de Cazorla: 15-07-1980, 1
specimen, F. Palacios (MNCN 32130); Peal de Becerro: Campamento de los Rasos: 08-06-
1981, 8 specimens, F. Palacios (MNCN 32131-32138); Peal de Becerro: Los Rasos, rio
Guadalquivir: 10-06-1981, 4 specimens, F. Palacios (MNCN 32139-32142); Peal de Becerro:
Campamento de los Rasos: 15-06-1981, 2 specimens, F. Palacios (MNCN 32143-32144); La
Hiruela: Roblehondo: Arroyo de las Gracias: 15-07-1980, 7 specimens, F. Palacios (MNCN
32145-32151); La Hiruela: Roblehondo: 20-07-1980, 3 specimens, F. Palacios (MNCN 32152-
32154); La Hiruela: Roblehondo: 22-08-1981, 2 specimens, F. Palacios (MNCN 32155-32156);
La Hiruela: Roblehondo: 14-04-1981, 4 specimens, F. Palacios (MNCN 32157-32160); Sierra
de Cazorla: Peña de Aguamula: 06-1968, 1 specimen, Palau (MNCN 32161); Peal de Becerro:
Sierra de Cazorla: Los Llanos de las Cuerdas - Gilillo: 08-06-1981, 1 specimen, F. Palacios
(MNCN 32162); Sierra de Cazorla: Nacimiento del río Aguamula: 02-04-1980, 8 specimens, F.
Palacios y J. Ayarzagüena (MNCN 32183-32190); La Hiruela: Roblehondo: 14-04-1981, 1
specimen, F. Palacios (MNCN 39291). No data available: 1 specimen (MNCN 31973).