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Terrestrial isopods from the Oued
Laou basin, north-eastern Morocco
(Crustacea: Oniscidea), with
descriptions of two new genera and
seven new species
Stefano Taitia & Claudia Rossanob
a Istituto per lo Studio degli Ecosistemi, Consiglio Nazionale delle
Ricerche, Florence, Italy
b Dipartimento di Biologia, University of Florence, Florence, Italy
Published online: 11 Mar 2015.
To cite this article: Stefano Taiti & Claudia Rossano (2015) Terrestrial isopods from the
Oued Laou basin, north-eastern Morocco (Crustacea: Oniscidea), with descriptions of two
new genera and seven new species, Journal of Natural History, 49:33-34, 2067-2138, DOI:
10.1080/00222933.2015.1009512
To link to this article: http://dx.doi.org/10.1080/00222933.2015.1009512
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Terrestrial isopods from the Oued Laou basin, north-eastern Morocco
(Crustacea: Oniscidea), with descriptions of two new genera and seven
new species
Stefano Taiti
a
*and Claudia Rossano
b
a
Istituto per lo Studio degli Ecosistemi, Consiglio Nazionale delle Ricerche, Florence, Italy;
b
Dipartimento di Biologia, University of Florence, Florence, Italy
(Received 5 June 2014; accepted 2 January 2015; first published online 11 March 2015)
Thirty-four species of terrestrial isopods (Crustacea: Oniscidea) from the Oued
Laou basin, in the Rif area of north-eastern Morocco, are recorded. One genus
(Paractenoscia) and seven species (Trichoniscus microphthalmus, Paractenoscia
cavernicola, Bathytropa rifensis, Soteriscus gibbosus, S. laouensis, Porcellio pseu-
dornatus, and Eluma praticola) are described as new. The genus Soteriscus, una-
vailable according to article 13.3 of the International Commission on Zoological
Nomenclature (ICZN), is here revalidated by choosing S. gaditanus as type
species. Six species (Graeconiscus thermophilus, Ctenoscia minima, Platyarthrus
parisii, Porcellio humberti, Porcellio flavocinctus and Eluma caelata) have been
fully illustrated to facilitate their identifications. Ctenoscia dorsalis Verhoeff is
considered to be a junior synonym of C. minima Dollfus. Porcellio ornatus from
southern Spain is also figured for comparison with P. pseudornatus sp. nov. The
composition and origin of the oniscidean fauna of the Rif region is briefly
discussed.
http://zoobank.org/urn:lsid:zoobank.org:pub:DCBF3103-1463-4A32-9BC0-
A4CFE8B762AE
Keywords: terrestrial Isopoda; taxonomy; new genera; new species; Morocco
Introduction
This paper deals with the terrestrial Isopoda collected in the Oued Laou valley in
Morocco during a biodiversity survey within the European Union project
MEDCORE. The area belongs to the Rif, the coastal mountain chain in the north-
eastern part of Morocco. In the Oligocene (c. 30 Ma), this chain together with the
Betic cordillera was connected to the Iberian peninsula and southern France, and
formed a continuous orogenic belt together with the Kabylies, Calabria, Corsica and
Sardinia (see the palaeogeographic reconstruction in Rosenbaum et al. 2002). During
the Miocene, the Rif–Betic belt moved westward and the formation of the Alboran
Sea split the mountain belt into the present Betic cordillera in the south-western
Iberian Peninsula and the Rif chain in north-eastern Morocco.
The Oniscidea of this area have been studied in the past by Vandel (1956a,1956b,
1958a,1958b,1958c), Schmalfuss (1987), Caruso and Di Maio (1990) and more
recently by Achouri et al. (2008a,2008c), Colombini et al. (2008), Charfi-
Cheikhrouha and El Gtari (2008) and Achouri and Charfi-Cheikhrouha (2009).
*Corresponding author. Email: stefano.taiti@ise.cnr.it
Journal of Natural History, 2015
Vol. 49, Nos. 33–34, 2067–2138, http://dx.doi.org/10.1080/00222933.2015.1009512
© 2015 Taylor & Francis
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Vandel (1958a,1958b) listed 17 species for the whole Rif region while Achouri
et al. (2008c) in a paper on the same area of this study (the Oued Laou basin)
recognized 20 species, some of which were only identified at genus level. In the
present paper 34 species of Oniscidea are recorded, of which seven are described as
new. A new genus (Paractenoscia) in the family Philosciidae is described and the
genus Soteriscus (Porcellionidae) is revalidated by choosing a type species.
Material and methods
The specimens were collected in 26 stations (St.) (Table 1) during two visits to the
Oued Laou valley in May 2004 and September 2005. The area investigated compre-
hends the Oued Laou basin from the river spring to the mouth in the Mediterranean,
including some minor tributaries. All different habitats present in the study area have
been investigated, i.e. sandy and rocky littoral shores, cultivated fields, meadows,
bushes, woods and some caves.
Table 1. List of sampling stations in the Oued Laou basin.
Station Locality Altitude
(m)
Latitude N
(WGS84)
Longitude W
(WGS84)
1 West of Oued Laou 35°28ʹ32.2″5°06ʹ20.4″
2 West of Oued Laou 35°28ʹ29.2″5°06ʹ23.5″
3 Oued Laou estuary, left bank 35°26ʹ09.7″5°04ʹ56.4″
4 Oued Laou estuary, right bank 35°25ʹ40.6″5°04ʹ41.8″
5 Oued Laou bay, East of Tizgane 35°24ʹ18.6″5°03ʹ30.4″
6 Tizrharine 20 35°23ʹ42.5″5°09ʹ05.8″
7 Right side of Oued Tassikesté, near
Bnehassan
50–110 35°23ʹ08.6″5°12ʹ40.9″
8 Ghar-Knadel Cave 220 35°22ʹ18.4″5°10ʹ56.0″
9 Larvha 100 35°22ʹ13.9″5°10ʹ42.1″
10 Along gorges between Afertane and
Ghboula
50 35°20ʹ36.1″5°11ʹ11.7″
11 Tirinesse, along Oued Kefhammar 50 35°20ʹ32.8″5°10ʹ37.6″
12 Gorge near Ghboula spring 50 35°20ʹ25.0″5°11ʹ25.4″
13 Kefhammar Cave 100 35°20ʹ23.3″5°10ʹ34.6″
14 Tirinesse 260 35°19ʹ38.2″5°10ʹ18.5″
15 Tirinesse, Hama 330 35°19ʹ33.4″5°09ʹ36.2″
16 Tirinesse 200 35°19ʹ29.9″5°10ʹ49.9″
17 Left side of Oued Abyet 120 35°17ʹ56.0″5°12ʹ52.4″
18 Fahsa 210 35°17ʹ45.1″5°13ʹ40.7″
19 Tarhzoute 350 35°15ʹ52.6″5°13ʹ59.8″
20 Near Akchour 35°13ʹ33.6″5°08ʹ56.1″
21 South of Akchour, Oued Farda valley,
Taskala spring
817 35°12ʹ45.7″5°10ʹ54.5″
22 South of Akchour, along Oued Farda 775 35°11ʹ41.9″5°10ʹ44.5″
23 Oued Laou spring, North of Majjo 900 35°07ʹ26.0″5°11ʹ55.4″
24 Dardara 380 35°05ʹ51.0″5°15ʹ45.1″
25 Near Oued Ouara 670 35°03ʹ48.5″5°14ʹ06.8″
26 El Khizana 900 35°02ʹ42.9″5°13ʹ44.9″
2068 S. Taiti and C. Rossano
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The specimens were stored in 75% ethanol and identifications are based on
morphological characters. For each new species the material examined, description,
etymology and remarks are given. For each species already recorded from Morocco
their bibliographic references, material examined, distribution and remarks (when
necessary) are given. Some poorly known species have been illustrated to facilitate
their recognition. The taxa are illustrated with figures prepared with the aid of a
camera lucida mounted on Wild M5 and M20 microscopes. For some species pictures
were taken with a scanning electron microscope.
The material is deposited in the collections of the Museo Zoologico ‘La Specola’
of the University of Florence, Italy (MZUF) and in the Staatliches Museum für
Naturkunde, Stuttgart, Germany (SMNS).
Systematic account
Family LIGIIDAE Leach
Genus Ligia Fabricius, 1798
Ligia italica Fabricius, 1798
Material examined
1♂,2♀♀, St. 1, on rocks near sea, leg. S. Taiti and C. Rossano, 28 September 2005
(MZUF 9449).
Distribution
Coasts of Black Sea and Mediterranean Sea, Atlantic coasts of the Iberian Peninsula
and northern Africa down to Cape Vert, and Macaronesian islands. New record for
the Rif region.
Remarks
For diagnostic characters of L. italica see Vandel (1960a, p. 122, Figures 54 and 55).
Family TYLIDAE Milne-Edwards
Genus Tylos Audouin, 1826
Tylos europaeus Arcangeli, 1938
Tylos europaeus; Achouri et al., 2008c, pp. 76–78; Colombini et al., 2008, p. 86;
Charfi-Cheikhrouha and El Gtari, 2008, p. 95.
Material examined
12 ♂♂,12♀♀, 8 juvs, St. 1, under debris on beach, leg. S. Taiti and C. Rossano, 28
September 2005 (MZUF 9450); 14 ♂♂,14♀♀, St. 5, beach under stones, leg. S. Taiti
and C. Rossano, 1 May 2004 (MZUF 9451); 1 ♀, same locality and collectors, 1
October 2005 (MZUF 9452).
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Previous Rif records
Oued Laou region (Achouri et al. 2008c); Oued Laou (Colombini et al. 2008);
Aouchtane (Charfi-Cheikhrouha and El Gtari 2008).
Distribution
This littoral species is known with certainty from the whole Mediterranean Sea, Black
Sea, Atlantic coasts of Europe as far north as Brittany, and the Azores.
Remarks
The exact distribution of T. europaeus is difficult to be defined since in the past it has
been often recorded as Tylos latreillii Audouin, 1826 together with the other
Mediterranean species T. ponticus Grebnitzky, 1874. For comments on the taxonomy
and diagnostic characters of this species see Taiti and Ferrara (1996).
Family TRICHONISCIDAE Sars
Genus Trichoniscus Brandt, 1833
Trichoniscus microphthalmus sp. nov.
(Figures 1 and 2)
Material examined
Holotype: ♂, St. 8, leg. S. Taiti and C. Rossano, 28 September 2005 (MZUF 9453).
Paratypes: 25 ♂♂,75♀♀, same data as holotype (MZUF 9453); 3 ♂♂,5♀♀ same
locality and collectors, 28 April 2004 (MZUF 9454).
Description
Maximum length: ♂, 2.0 mm; ♀, 2.5 mm. Body colourless, ovoidal, with pleon
narrower than pereon (Figure 1A). Back almost smooth covered with numerous
cordiform scale-setae (Figure 1B). Cephalon (Figure 1C) with suprantennal line
bent downwards; antennal lobes rounded, distinctly visible in dorsal view; eye
reduced, visible as one to three small dots of dark pigment. Distal part of telson
with concave sides and truncate apex (Figure 1D). Antennule (Figure 1E) of three
articles; distal article longer than second, bearing three aesthetascs on apical margin.
Antenna (Figure 1F) fifth article as long as flagellum; flagellum of three articles with
one row of five aesthetascs on second article. Mandibles with one penicil in the right
(Figure 1G) and two penicils in the left (Figure 1H). Outer branch of maxillule with
5 + 5 teeth, apically entire, and three slender stalks; inner branch with three penicils
(Figure 2A). Maxilla with setose and bilobate apex, inner lobe smaller (Figure 2B).
Maxilliped endite narrow, with a large apical penicil (Figure 2C). Pereopods with an
ungual seta and a large, bifid and setose dactylar seta (Figure 2D,2E). Uropod
(Figure 1D) with protopod not grooved on outer margin; endopod slightly shorter
than exopod, inserted at the same level.
2070 S. Taiti and C. Rossano
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Figure 1. Trichoniscus microphthalmus sp. nov. from St. 8, paratype ♀: (A) adult specimen,
dorsal view; (B) dorsal scale-seta; (C) cephalon, frontal view; (D) pleonite 5, telson and left
uropod; (E) antennule; (F) antenna; (G) right mandible; (H) left mandible.
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Figure 2. Trichoniscus microphthalmus sp. nov. from St. 8, paratype ♀: (A) maxillule; (B)
maxilla; (C) maxilliped. Paratype ♂: (D) pereopod 1; (E) pereopod 7; (F) pleopod 1; (G)
pleopod 2.
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Male: Pereopods 1–4(Figure 2D) with carpus and merus bearing numerous short
scales on sternal margin. Pereopod 7 (Figure 2E) ischium with straight sternal
margin. Pleopod 1 (Figure 2F) exopod with straight medial margin and sinuous
outer margin; endopod of two articles, distal article tapering to a point, apical part
shagreened. Pleopod 2 (Figure 2G) exopod subrectangular with distal margin slightly
convex; endopod of two articles, distal article styliform, about three times as long as
basal one.
Etymology
From the Greek ‘mikrós’= small + ‘ophthalmós’= eye. The name refers to the
reduced eye visible as one to three small dots of dark pigment.
Remarks
The genus Trichoniscus was previously known in Morocco only for the widespread
species T. pygmaeus Sars, 1899 (Vandel 1959). A second species, Trichoniscus soli-
sensis Vandel, 1959, from a cave near Safi, western Morocco, has been proved to
belong to the genus Adoniscus Vandel, 1955c (Olibrinidae) (Taiti and Ferrara 2004).
In Algeria four species of Trichoniscus are known: T. gachassini (Giard, 1899), T.
fragilis Racovitza, 1908,T. provisorius Racovitza, 1908, and T. peyerimhoffi Vandel,
1955a (Schmalfuss 2003). Another species of Trichoniscus (T. gordoni Vandel, 1955a)
is known from several caves in southern Spain and Gibraltar (Vandel 1955a). The
new species T. microphthalmus is readily distinguishable from all these species by the
male pereopods 1–4 with sternal margin of carpus and merus bearing a fringe of
scales, and by the shape of the male pleopod 1 exopod.
Genus Graeconiscus Strouhal, 1940
Graeconiscus thermophilus (Çağlar, 1948)
(Figures 3–5)
Material examined
72 ♂♂,132♀♀, 3 juvs, St. 8, leg. S. Taiti and C. Rossano, 28 September 2005
(MZUF 9455); 2 ♂♂,2♀♀, same data (SMNS 15676); 2 ♂♂,1♀, same locality and
collectors, 28 April 2004 (MZUF 9456).
Distribution
Graeconiscus thermophilus was previously known from the southern Aegean islands
including Crete and western Turkey. New record for the Rif region.
Remarks
Haplophthalmus thermophilus was described by Çağlar (1948) on specimens from a
warm spring at Armutlu, near Gemlik (Turkey). Strouhal (1963) redescribed and
figured this species from the type locality. Schmalfuss et al. (2004) included the
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species in the genus Graeconiscus and considered Calconischellus aegeus Schmalfuss,
1972 recorded from Crete and several southern Aegean islands as a junior synonym
of G. thermophilus. The specimens from Ghar-Knadel Cave (maximum length ♂
3 mm, ♀3.7 mm) are tentatively identified as Graeconiscus thermophilus since they
correspond to the redescription provided by Strouhal (1963, p. 392, Figures 22–26); in
the disposition of dorsal ornamentation and the morphology of the male pereopod 7
and pleopod 1. They differ in lacking the eyes and in the shape of the tubercles on
pereonite 7 and pleonite 3. In G. thermophilus the pereonite 7 has 3 + 3 tubercles as in
the Moroccan specimens but the medial tubercle on each side is much smaller, and
the single tubercle on pleonite 3 is transversally elongated while it is rounded in the
Ghar-Knadel Cave specimens. The main characters of these specimens are illustrated
in Figures 3–5.
Figure 3. Graeconiscus thermophilus from St. 8, ♂: (A) adult specimen, dorsal view; (B)
pereonites 6, 7 and pleon, ventral view; (C) pereopod 7.
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Figure 4. Graeconiscus thermophilus from St. 8, ♀: (A) pleonite 5, telson and uropods; (B)
antennule; (C) antenna; (D) right mandible; (E) left mandible; (F) maxillule; (G) maxilla; (H)
maxilliped.
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Family HALOPHILOSCIIDAE Verhoeff
Genus Halophiloscia Verhoeff, 1908
Halophiloscia couchii (Kinahan, 1858)
Philoscia couchii; Dollfus, 1896, p. 548
Philoscia Couchii; Paulian de Félice, 1939, p. 192
Halophiloscia couchi; Vandel, 1958a p. 128
Halophiloscia couchii; Taiti and López, 2008, p. 44, Figures 1A–C,8A,B,9
Material examined
10 ♂♂,21♀♀, St. 1, under debris on beach, leg. S. Taiti and C. Rossano, 28
September 2005 (MZUF 9457, 9458); 7 ♂♂,6♀♀, St. 5, beach under stones, leg.
S. Taiti and C. Rossano, 1 May 2004 (MZUF 9459).
Figure 5. Graeconiscus thermophilus from St. 8, ♂: (A) pereopod 1; (B) pereopod 7; (C)
pleopod 1; (D) pleopod 2.
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Previous Rif records
Melilla, Mar Chica (Vandel 1958a); Oued Laou (Taiti and López 2008).
Distribution
Coasts of the Atlantic Ocean from Senegal to the British Isles, Azores, Madeira,
Canary Islands, Cape Verde, Mediterranean and Black Sea. Introduced to North and
South America, Hawaii and Australia.
Remarks
For diagnostic characters of H. couchii see Schmidt (2003, Figures 52–57) and Taiti
and López (2008, p. 44, Figures 1A–C,8A,B).
Genus Stenophiloscia Verhoeff, 1908
Stenophiloscia glarearum Verhoeff, 1908
Stenophiloscia glarearum; Colombini et al., 2008,p.86
Previous Rif records
Oued Laou (Colombini et al. 2008).
Material examined
16 ♂♂,8♀♀, 2 juvs, St. 1, under debris on beach, leg. S. Taiti and C. Rossano, 28
September 2005 (MZUF 9460, 9461); 2 ♂♂,1♀, St. 5, beach under stones, leg. S.
Taiti and C. Rossano, 1 May 2004 (MZUF 9462); 11 ♂♂,10♀♀, 1 juv., same
locality and collectors, 1 October 2005 (MZUF 9463).
Distribution
Southern England, Canary Islands, Morocco, eastern Spain, Balearic Islands, south-
east France, Italy, Malta, Croatia, and Ionian coasts of Greece (Schmalfuss 2003).
Remarks
For diagnostic characters of S. glarearum see Vandel (1962, p. 488, Figures 244 and
245) and Ferrara and Taiti (1978, p. 22, Figure 5) as Halophiloscia (Stenophiloscia)
zosterae Verhoeff, 1928, junior synonym of S. glarearum (see Schmalfuss 2003).
Family PHILOSCIIDAE Kinahan
Genus Ctenoscia Verhoeff, 1928
Ctenoscia minima (Dollfus, 1892)
(Figures 6–8)
Chaetophiloscia dorsalis Verhoeff, 1928, p. 137, Figures 34,37,38, syn. nov.
Ctenoscia minima; Vandel, 1961, p. 256
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Figure 6. Ctenoscia minima from St. 14, ♀: (A) adult specimen, dorsal view; (B) dorsal scale-
seta; (C) co-ordinates of noduli laterals; (D) cephalon, dorsal view; (E) cephalon, frontal view;
(F) left side of pereon showing disposition of noduli laterales; (G) pereonite 1, left side; (H)
pleonites 4, 5, telson and uropods; (I) antennule.
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Figure 7. Ctenoscia minima from St. 14, ♀: (A) antenna; (B) left mandible; (C) right mandible;
(D) maxillule; (E) maxilla; (F) maxilliped.
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Figure 8. Ctenoscia minima from St. 14, ♂: (A) pereopod 1; (B) pereopod 7; (C) pleopod 1
(specimen 4.5 mm long); (D) genital papilla and pleopod 1 (specimen 4.0 mm long); (E)
pleopod 2; (F) pleopod 3 exopod; (G) pleopod 4 exopod; (H) pleopod 5 exopod.
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Material examined
2♂♂,16♀♀, 1 juv., St. 8, leg. S. Taiti and C. Rossano, 28 April 2004 (MZUF
9464); 4 ♂♂,14♀♀, St. 10, leg. S. Taiti and C. Rossano, 26 April 2004 (MZUF
9465); 1 ♀, St. 12, leg. C. Rossano, 26 April 2004 (MZUF 9466); 12 ♂♂,20♀♀,
St. 14, Phillyrea wood, leg. S. Taiti, 27 April 2004 (MZUF 9467); many ♂♂ and
♀♀, St. 15, sieved ground near stream, leg. S. Taiti, 27 April 2004 (MZUF 9468); 1
♀, St. 17, leg. S. Taiti, 29 April 2004 (MZUF 9469); 1 ♀, St. 18, cork-oak wood,
leg. S. Taiti, 29 April 2004 (MZUF 9470); 1 ♂,2♀♀, St. 21, leg. S. Taiti and C.
Rossano, 30.IX.2005 (MZUF 9471); 2 ♀♀, St. 24, cork-oak wood, leg. S. Taiti, 29
April 2004 (MZUF 9472).
Remarks
Ctenoscia minima was originally described by Dollfus (1892) from Grenada, Spain,
and later recorded for several localities in Spain, including the Canary and Balearic
islands, Portugal (see Schmalfuss 2003 for previous citations), and Morocco (Vandel
1961). This species has been illustrated by Vandel (1946) and Rodríguez and
Barrientos (1993). The genus Ctenoscia also includes another species, C. dorsalis
(Verhoeff, 1928), originally described from San Remo and Grimaldi (Liguria,
Italy), and later recorded from several localities in mainland Spain and the Balearic
Islands, Corsica, Italy including Sardinia and Sicily, Malta (see Schmalfuss 2003 for
previous citations), and Tunisia (Achouri et al. 2008b).
According to the descriptions, C. minima differs from C. dorsalis in having the
distal part of the male pleopod 1 endopod bent outwards instead of straight, while all
the other characters are identical in the two species. The examination of many
specimens of Ctenoscia from the Oued Laou valley showed that C. dorsalis has to
be considered as a junior synonym of C. minima. In fact, the distal part of the male
pleopod 1 endopod is straight or bent outwards according to the size of the speci-
mens. In the population from St. 14 (Tirinesse), 4 mm long males show the endopod
bent outwards as in C. minima (Figure 8D), while in 4.5 mm long males this
appendage is straight as in C. dorsalis (Figure 8C), showing that this character varies
according to the age and size of the specimens. For the same reason also the distal
margin of the male pleopod 1 exopod is more or less convex.
Ctenoscia minima is here fully illustrated to facilitate its identification (Figures 6–8).
Distribution
Portugal, Spain (including Canary and Balearic islands), Corsica, Italy (including
Sardinia, Sicily and surrounding islands), Malta, Morocco and Tunisia. New record
for the Rif region.
Genus Paractenoscia gen. nov.
Diagnosis
Pereonites with one line of noduli laterales per side, noduli inserted more or less at
the same distance from the lateral margins and at the same distance from the
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posterior margins of the pereonites. No visible gland pores on lateral margins of
pereonites. Cephalon with suprantennal line and without frontal line. Pleon dis-
tinctly narrower than pereon; epimera of pleonites 3–5 adpressed, ventrally with
small posterior points not visible in dorsal view. Antennule with petaliform aesthe-
tascs. Molar process of mandible dichotomized (i.e. consisting of some plumose
setae each arising separately). Outer branch of maxillule with some pectinate teeth;
inner branch with short and stout penicils and without posterior point. Inner lobe of
maxilla with verruca-like protuberances and a tuft of short segmented sticks.
Maxilliped endite without penicil. Pereopods with inner claw of dactylus enlarged.
Pleopodal exopod without respiratory structures. Uropods with protopod and exo-
pod grooved on outer margin; insertion of endopod and exopod almost at the same
level.
Etymology
From the Greek ‘pará’= beside, near + Ctenoscia. The name refers to the similarities
with the genus Ctenoscia.
Remarks
Paractenoscia is very similar to the genera Ctenoscia and Anaphiloscia Racovitza,
1907 from the Mediterranean area. It is readily distinguished from both in the maxilla
with verruca-like protuberances and a tuft of short sticks on outer lobe; from
Ctenoscia also in having the noduli laterales all close to the posterior margin of the
pereonites, antennule with petaliform instead of thin aesthetascs, molar process of the
mandible dichotomized instead of semidichotomized (i.e. consisting of some plumose
setae arising from a common stem, see Ferrara et al. 1995), and maxillular inner
branch with short and stout penicils and without posterior point; from Anaphiloscia
also in having pointed instead of fan-shaped dorsal scale-setae and dichotomized
instead of simple molar penicil. Pectinate maxillular teeth are present also in the
genera Benthanops Barnard, 1932 from South Africa, Benthana Budde-Lund, 1908,
Benthanoides Lemos de Castro, 1958 and Alboscia Schultz, 1995 from South
America. Paractenoscia differs from Benthanops in the number and position of the
noduli laterales (see Taiti and Ferrara 1982), from Benthana, Benthanoides and
Alboscia in the inner branch of the maxillule with short and stout instead of long
and thin penicils and in the peculiar shape of the maxilla with the inner lobe covered
with verruca-like protuberances and with a tuft of short segmented sticks near the
medial margin, features not present in any other genera of Philosciidae.
Paractenoscia cavernicola sp. nov.
(Figures 9–11)
Material examined
Holotype: ♂, St. 8, leg. S. Taiti and C. Rossano, 28 April 2004 (MZUF 9473).
Paratypes: 3 ♂,9♀♀, same data as holotype (MZUF 9473).
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Figure 9. Paractenoscia cavernicola sp. nov. from St. 8, paratype ♀: (A) adult specimen, dorsal
view; (B) dorsal scale-seta; (C) co-ordinates of noduli laterals; (D) cephalon, dorsal view; (E)
cephalon, frontal view; (F) right side of pereon showing disposition of noduli laterales; (G)
pleonites 3–5, telson and uropods; (H) antennule; (I) antenna.
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Description
Maximum length: ♂, 3.5 mm; ♀, 4.8 mm. Body outline as in Figure 9A. Colourless
body. Back smooth with some scattered pointed scale-setae (Figure 9B); noduli
laterales with b/c and d/c co-ordinates as in Figure 9C. Cephalon (Figure 9D,9E)
with suprantennal line slightly bent downwards in the middle; eyes absent. Pereonites
1–3 with convex distal margins and rounded posterior corners; pereonites 4–7with
Figure 10. Paractenoscia cavernicola sp. nov. from St. 8, paratype ♀: (A) left mandible; (B)
right mandible; (C) maxillule; (D) maxilla; (E) maxilliped.
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Figure 11. Paractenoscia cavernicola sp. nov. from St. 8, paratype ♂: (A) antenna; (B)
pereopod 1; (C) pereopod 7; (D) genital papilla and pleopod 1; (E) pleopod 2; (F) pleopod 3
exopod; (G) pleopod 4 exopod; (H) pleopod 5 exopod.
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progressively more acute corners (Figure 9F). Telson triangular with slightly convex
sides (Figure 9G). Antennule (Figure 9H) with third article longer than first and
second; third article with two petaliform apical aesthetascs, two petaliform aesthe-
tascs in the middle and one pointed aesthetasc closer to proximal margin. Antenna
(Figures 9I,11A) with flagellum slightly longer than fifth article of peduncle; flagellar
articles subequal in length; a row of three aesthetascs on second and four aesthetascs
on third flagellar article. Mandible with molar penicil consisting of three long and
two short setae; left mandible (Figure 10A) with 2 + 1 free penicils and two lines of
scales between the two groups of free penicils; right mandible (Figure 10B)with1+1
free penicils. Maxillule outer branch with 4 + 6 (4 pectinate) teeth; inner branch with
two short stout penicils (Figure 10C). Maxilla (Figure 10D) deeply bilobate with
setose apex, inner lobe slightly larger than outer one, bearing verruca-like protuber-
ances at the base and with tuft of short segmented sticks near medial margin.
Maxilliped (Figure 10E) endite with a line of setae in the middle and no penicil;
first article of palp with two strong setae. Pereopods with a row of scales on distal
margins of ischium, merus and carpus.
Male: Antenna with fifth article of the peduncle distinctly enlarged (Figure 11A).
Pereopod 1 (Figure 11B) with no distinct sexual modifications. Pereopod 7
(Figure 11C) ischium with straight sternal margin. Pleopod 1 (Figure 11D) exopod
subtriangular; endopod with pointed apical part slightly bent outwards. Pleopod 2
(Figure 11E) endopod with apical part flagelliform, distinctly longer than exopod.
Pleopod 3–5 exopods as in Figure 11F–H.
Etymology
From the Latin ‘cavernicolus’= cavernicolous, cave-dwelling. The name refers to the
locality of collection: Ghar-Knadel Cave.
Family BATHYTROPIDAE Vandel
Genus Bathytropa Budde-Lund, 1885
Bathytropa rifensis sp. nov.
(Figures 12–14)
Material examined
Holotype: ♂, St. 15, sieved ground near stream, leg. S. Taiti, 27 April 2004 (MZUF
9474). Paratypes: 2 ♂♂,5♀♀, same data as holotype (MZUF 9474).
Description
Maximum length: ♂, 2.8 mm; ♀, 4.0 mm. Colour pale. Body strongly convex with
epimera of pereon and pleon enlarged, obliquely directed, clinger type (Schmalfuss
1984). Dorsal surface of cephalon, pereon and pleon with large tubercles and ribs
arranged as in Figure 12A. Back covered with triangular scale-setae (Figure 12B);
posterior margins of the body segments with rectangular scale-setae (Figure 12C); one
line of noduli laterales per side arranged on top of the outmost tubercle of the
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Figure 12. Bathytropa rifensis sp. nov. from St. 15, paratype ♀: (A) adult specimen, dorsal
view; (B) dorsal scale-seta; (C) scale-seta on posterior margins of segments; (D) cephalon,
dorsal view; (E) cephalon, frontal view; (F) pleonite 5, telson and right uropod; (G) antennule;
(H) antenna.
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posterior row of each pereonite. Cephalon (Figure 12D,E) with median lobe raised
up, broadly rounded in dorsal view and rectangular in frontal view; quadrangular
lateral lobes, obliquely directed and slightly more protruding frontwards than median
Figure 13. Bathytropa rifensis sp. nov. from St. 15, paratype ♂: (A) left mandible; (B) right
mandible; (C) maxillule; (D) maxilla; (E) maxilliped; (F) pereopod 1.
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Figure 14. Bathytropa rifensis sp. nov. from St. 8, paratype ♂: (A) pereopod 7; (B) genital
papilla and pleopod 1; (C) pleopod 2; (D) pleopod 3 exopod; (E) pleopod 4 exopod; (F)
pleopod 5 exopod.
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lobe; no suprantennal line; eye small with five ommatidia. Pereon with quadrangular
epimera progressively pointing backwards from first to seventh; posterior margin of
the first pereonite slightly sinuous at sides. Pleonites 3–5 with subrectangular epimera
continuing the outline of the pereon. Telson slightly wider than long, triangular with
straight sides and broadly rounded apex (Figure 12F). Antennule (Figure 12G) with
second article much shorter than first and third; third article with an apical tuft of
four aesthetascs. Antenna (Figure 12H) with flagellum about as long as fifth article of
peduncle; second flagellar article about four times as long as first, bearing a row of
four aesthetascs in the middle. Mandibles with molar penicil consisting of four or five
hairy setae; left mandible (Figure 13A) with 2 + 1 free penicils; right mandible
(Figure 13B) with 1 + 1 free penicils. Maxillule (Figure 13C) outer branch with
4 + 7 (3 cleft) teeth; inner branch with two short stout penicils and a distinct apical
point. Maxilla (Figure 13D) distally bilobate with setose apex; inner lobe much wider
than outer one. Maxilliped (Figure 13E) endite with three triangular stout teeth on
distal margin and no penicil; first article of palp with two setae, the medial one much
longer than the outer one. Pleopodal exopods with no respiratory structures as in
most species of the genus.
Male: Pereopod 1 (Figure 13F) with no distinct sexual modifications. Pereopod 7
(Figure 14A) ischium enlarged in the distal part, with straight sternal margin.
Pleopod 1 (Figure 14B) exopod subtriangular with rounded distal part; endopod
with pointed apical part slightly bent outwards. Pleopod 2 (Figure 14C) endopod
with apical part flagelliform, slightly longer than exopod. Pleopod 3–5 exopods as in
Figure 14D–F.
Etymology
The name refers to Rif where the specimens have been collected.
Remarks
At present the genus Bathytropa includes 10 species distributed in the Mediterranean
area (Schmalfuss 2003). According to Vandel (1962), B. meinertii Budde-Lund, 1885
includes two subspecies: B. m. meinertii and B. m. costata Budde-Lund, 1885. The
new species is readily distinguishable from B. meinertii costata, B. tuberculata
Racovitza, 1908, and B. schembrii Caruso and Lombardo, 1982 in the presence of
two paramedian tubercles instead of a single median tubercle on pleonites; from B.
granulata Aubert and Dollfus, 1890 and B. graevei (Verhoeff, 1940) in having distinct
dorsal ribs and tubercles instead of granulations; from B. wahrmani Strouhal, 1968 in
the telson triangular with rounded apex, instead of hour-glass-shaped, and the
absence of respiratory structures on the pleopods; from B. meinertii meinertii, B.
colasi Vandel, 1954,B. dollfusi Strouhal, 1936,B. patanei Caruso, 1973a and B. ruffoi
Caruso, 1973b in the number and disposition of dorsal ornamentation. In northern
Africa three species of Bathytropa were previously recorded: B. tuberculata and B.
meinertii, with both subspecies B. m. meinertii and B. m. costata, from Algeria, and B.
colasi Vandel, 1954 from ‘Berberie’(Vandel 1955b).
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Family STENONISCIDAE Budde-Lund
Genus Stenoniscus Aubert and Dollfus, 1890
Stenoniscus carinatus Silvestri, 1897
Material examined
1♀, St. 5, beach under stones, leg. S. Taiti and C. Rossano, 1 May 2004 (MZUF 9475).
Distribution
This littoral species is common along the coasts of Canary Islands, Portugal, Corsica,
Italy, Malta and Croatia. It is also recorded from Florida (Taiti and Ferrara 1996),
where it is most probably introduced. New record for the Rif region and northern
Africa.
Remarks
For diagnostic characters of S. carinatus see Schmidt (2003, Figures 67–72).
Family PLATYARTHRIDAE Verhoeff
Genus Platyarthrus Brandt, 1833
Platyarthrus caudatus Aubert and Dollfus, 1890
Material examined
9♂♂,12♀♀, St. 20, under stones along path to waterfall, leg. S. Taiti and C.
Rossano, 2 May 2004 (MZUF 9476).
Distribution
Known from most lands of the western Mediterranean. New record for the Rif
region.
Remarks
For diagnostic characters of P. caudatus see Vandel (1962, p. 457, Figures 228, 229).
Platyarthrus parisii Arcangeli, 1930
(Figure 15)
Platyarthrus schöbli parisii; Vandel, 1946, p. 219
Platyarthrus Parisii; Arcangeli, 1952, p. 137
Material examined
1♂,1♀, St. 25, under stones in meadow, leg. S. Taiti, 30 April 2004 (MZUF 9477).
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Distribution
Canary Islands and Morocco. New record for the Rif region.
Remarks
Platyarthrus parisii was described by Arcangeli (1930) on female specimens from the
Canary Islands (Gran Canaria and Tenerife). Vandel (1946) considers this form as a
Figure 15. Platyarthrus parisii from St. 25, ♂: (A) dorsal scale-seta; (B) cephalon and pereonite
1, dorsal view; (C) pleonite 5, telson and uropods; (D) pereopod 1; (E) pereopod 7; (F) pleopod 1.
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subspecies of P. schoblii Budde-Lund, 1885 and adds a record for Morocco (Mamora
Forest, between Rabat and Meknès). Arcangeli (1952) in a discussion on the various
subspecies of P. schoblii definitely considers this form as a species (see also
Schmalfuss 2003). The specimens from the St. 25 (near Oued Ouara) examined by
us show the same large lateral lobes of the cepalon and the same disposition of the
dorsal ribs as P. parisii, but differ in the more protruding median frontal lobe. It is
difficult to say whether this character can be enough to distinguish a distinct species
or it falls within the variability of P. parisii. In order to facilitate a future comparison
with the species from the Canary Islands the main characters of the specimens from
Oued Ouara are illustrated in Figure 15.
Family PORCELLIONIDAE Brandt
Genus Leptotrichus Budde-Lund, 1885
Leptotrichus panzerii (Audouin, 1826)
Leptotrichus Panzeri; Dollfus, 1896, p. 542; Paulian de Félice, 1939, p. 212
Leptotrichus panzeri; Achouri et al., 2008c,p.76–78
Leptotrichus panzerii; Colombini et al., 2008,p.86
Material examined
2♂♂,5♀♀, St. 3, leg. C. Rossano, 28 April 2004 (MZUF 9478); 1 ♀, same locality,
under stones, leg. S. Taiti and C. Rossano, 1 October 2005 (MZUF 9479); 3 ♂♂,5
♀♀, St. 4, dune under stones, leg. S. Taiti and C. Rossano, 1 May 2004 (MZUF
9480).
Previous Rif records
Oued Laou region (Achouri et al. 2008c); Oued Laou (Colombini et al. 2008).
Distribution
Countries encompassed by the Mediterranean Sea, Azores, Madeira, Canary Islands,
Cape Verde, Saint Helena and Bermuda.
Remarks
For diagnostic characters of L. panzerii see Vandel (1962, p. 645, Figures 317, 318).
Genus Agabiformius Verhoeff, 1908
Agabiformius lentus (Budde-Lund, 1885)
Agabiformius lentus; Achouri et al., 2008c,p.76–78; Achouri et al., 2008a, p. 129
Material examined
2♂♂,1♀, St. 7, leg. S. Taiti and C. Rossano, 29 September 2005 (MZUF 9481).
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Previous Rif records
Oued Laou region (Achouri et al. 2008a,2008c).
Distribution
This species is common in all the lands of the Mediterranean basin where it is native.
It has been introduced with human activities in many parts of the world.
Agabiformius obtusus (Budde-Lund, 1909)
Agabiformius obtusus; Colombini et al., 2008,p.86
Material examined
4♀♀, St. 4, dune under stones, leg. S. Taiti and C. Rossano, 1 May 2004 (MZUF
9482); 1 ♀, St. 5, beach under stones, leg. S. Taiti and C. Rossano, 1 May 2004
(MZUF 9483).
Previous Rif records
Oued Laou (Colombini et al. 2008).
Distribution
This species occurs in the lands of the Mediterranean, where it is very common along
sandy beaches. It is also recorded from Kuwait and Sudan.
Remarks
For diagnostic characters of A. lentus see Vandel (1962, p. 640, Figures 315, 316).
Genus Lucasius Kinahan, 1859
Lucasius pallidus (Budde-Lund, 1885)
Material examined
5♂♂,2♀♀, St. 8, 220 m, leg. S. Taiti and C. Rossano, 28 April 2004 (MZUF 9484);
26 ♂♂,18♀♀, 2 juvs, same locality and collectors, 28 September 2005 (MZUF
9485); 11 ♂♂,21♀♀, St. 25, under stones in meadow, leg. S. Taiti, 30 April 2004
(MZUF 9486); 2 ♀♀, St. 26, cork-oak wood, leg. S. Taiti, 30 April 2004 (MZUF
9487).
Distribution
Morocco, southern Spain, southern France, Corsica, Sardinia and Tuscany. New
record for the Rif region.
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Remarks
In Morocco two species of Lucasius have been recorded: L. myrmecophilus Kinahan,
1859, originally described from Algeria (Kinahan 1859); and L. pallidus, originally
described from southern France, Sicily, southern Spain and Algeria. In Sicily this
species was confused with Mica tardus (Budde-Lund, 1885) and, according to Caruso
and Di Maio (1996), it is not present. Achouri et al. (2008c) cite L. myrmecophilus
from the Oued Laou basin. The specimens from the same area examined by us
definitely fit the description of L. pallidus, but the differences of this species with L.
myrmecophilus are still unclear. According to Vandel (1962) and Schmölzer (1965)L.
myrmecophilus differs from L. pallidus in having a smooth instead of granulated
dorsum, but in the original description by Kinahan (1859) and in the redescription
by Budde-Lund (1885, p. 135) it is clearly stated that the dorsum of the body is
granulated. Therefore there is a possibility that the two species are synonymous, but a
re-examination of the type material of L. myrmecophilus is necessary to confirm this
hypothesis.
For diagnostic characters of L. pallidus see Vandel (1962, p. 651, Figure 321).
Genus Porcellionides Miers, 1877
Porcellionides pruinosus (Brandt, 1833)
Metoponorthus (Metoponorthus) pruinosus; Vandel, 1958a, p. 129
Porcellionides pruinosus; Achouri et al., 2008c,p.76–78; Achouri et al., 2008a, p. 129
Material examined
1♂,1♀, St. 13, cave entrance, leg. C. Rossano, 28 April 2004 (MZUF 9488).
Previous Rif records
Peñón de Alhucemas (Vandel 1958a); Oued Laou region (Achouri et al. 2008a,
2008c).
Distribution
Species of Mediterranean origin with a cosmopolitan distribution.
Remarks
For diagnostic characters of P. pruinosus see Vandel (1962, p. 618, Figures 306, 307)
and Gruner (1966, p. 251, Figures 190–193).
Porcellionides sexfasciatus lusitanus (Vandel, 1946)
Metoponorthus sexfasciatus lusitanus Vandel, 1946, p. 269, Figures 80B–D, 84, 85
?Porcellionides sexfasciatus; Achouri and Charfi-Cheikhrouha, 2009, p. 1048
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Material examined
1♂, St. 2, under stones at base of cliff, leg. S. Taiti and C. Rossano, 1 May 2004
(MZUF 9489); 1 ♂,1♀, St. 3, under stones, leg. S. Taiti and C. Rossano, 1 October
2005 (MZUF 9490); 1 ♀, St. 4, beach under logs, leg. S. Taiti and C. Rossano, 1 May
2004 (MZUF 9491); 1 ♂,2♀♀, St. 6, near well and wheat field, leg. S. Taiti and C.
Rossano, 26 April 2004 (MZUF 9492); 1 ♂, St. 7, leg. S. Taiti, 29 April 2004 (MZUF
9493); 1 ♂, same locality, leg. S. Taiti and C. Rossano, 29 September 2005 (MZUF
9494); 1 ♀, St. 9, under stones in meadow, leg. S. Taiti, 28 April 2004 (MZUF 9495);
3♂♂,9♀♀, St. 10, leg. S. Taiti and C. Rossano, 26 April 2004 (MZUF 9496); 1 ♂,2
♀♀, St. 14, Phillyrea wood, leg. S. Taiti, 27 April 2004 (MZUF 9497); 5 ♂♂,3♀♀,
same locality, leg. S. Taiti and C. Rossano, 27 September 2005 (MZUF 9498); 4 ♂♂,
6♀♀, St. 19, leg. S. Taiti, 29 April 2004 (MZUF 9499); 3 ♀♀, St. 23, under stones,
leg. S. Taiti, 30 April 2004 (MZUF 9500); 5 ♂♂,11♀♀, St. 25, under stones in
meadow, leg. S. Taiti, 30 April 2004 (MZUF 9501).
Distribution
Porcellionides sexfasciatus (Budde-Lund, 1885) is widely distributed in the western
Mediterranean region, Atlantic coasts of Europe and northern Africa, including
Atlantic islands. It has been also introduced to many other parts of the world. At
present P. sexfasciatus includes several subspecies. The specimens from the Rif
examined by us belong to P. sexfasciatus lusitanus distributed in Portugal and
Morocco (Vandel 1946). New record for the Rif region.
Porcellionides sp.
(Figures 16 and 17)
Material examined
1♀, St. 4, dune under stones, leg. S. Taiti and C. Rossano, 1 May 2004 (MZUF
9502); 1 ♂, St. 6, near well and wheat field, leg. S. Taiti and C. Rossano, 26 April
2004 (MZUF 9503).
Remarks
These specimens certainly belong to the genus and subgenus Porcellionides, but they
do not seem to belong to any of the species of Porcellionides recorded from Morocco
or the Iberian Peninsula. This species is morphologically close to P. pruinosus from
which it differs in the more granulated dorsum and shape of the male pleopod 2 with
exopod having a triangular posterior lobe and endopod tip slightly bent inwards.
Since only two specimens were examined, we illustrate the main characters of the
species (Figures 16 and 17) but we prefer not to identify the species until a more
abundant material can be analysed.
Genus Soteriscus gen. nov.
Type species: Soteriscus gaditanus Vandel, 1956b by present designation
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Figure 16. Porcellionides sp., ♀from St. 4: (A) adult specimen, dorsal view; (B) dorsal scale-
seta; (C) cephalon, dorsal view; (D) cephalon, frontal view; (E) pleonites 4, 5, telson and
uropods; (F) antenna. ♂from St. 6: (G) pereopod 1.
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Figure 17. Porcellionides sp., ♂from St. 6: (A) pereopod 7; (B) genital papilla and pleopod 1;
(C) pleopod 2; (D) pleopod 3 exopod; (E) pleopod 4 exopod; (F) pleopod 5 exopod.
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Remarks
Soteriscus was erected by Vandel (1956b) as a subgenus of Metoponorthus Budde-
Lund, 1879 (junior synonym of Porcellionides) to accommodate four species:
Metoponorthus (Soteriscus) stricticauda Dollfus, 1893 from the Canary Islands;
Metoponorthus (Soteriscus) wollastoni Paulian de Félice, 1939 from Madera Island;
Metoponorthus (Soteriscus) fuscovariegatus (Lucas, 1849) from Algeria; and the new
species Metoponorthus (Soteriscus) gaditanus from south-western Spain and north-
western Morocco. No type species for the subgenus Soteriscus was selected neither by
Vandel (1956b), nor by Vandel and Matsakis (1959) when the subgenus was elevated
to genus rank, nor in any other subsequent publication dealing with this genus. Thus,
according to article 13.3 of ICZN (1999), the name is unavailable (see also Schmidt
and Leistikow 2004). In order to revalidate the genus we designate here
Metoponorthus (Soteriscus) gaditanus as type species of the genus Soteriscus. For
the diagnosis of the genus see Vandel (1960b).
Soteriscus gibbosus sp. nov.
(Figures 18–20)
Material examined
Holotype: ♂, St. 14, Phillyrea wood, leg. S. Taiti, 27 April 2004 (MZUF 9504).
Paratypes: 3 ♂♂,11♀♀, same data as holotype (MZUF 9504); 1 ♂,6♀♀, same
locality, leg. S. Taiti and C. Rossano, 27 September 2005 (MZUF 9505); 1 ♀, St. 10,
leg. S. Taiti and C. Rossano, 26 April 2004 (MZUF 9506); 1♀, St. 12, leg. C.
Rossano, 26 April 2004 (MZUF 9507); 1 ♀, St. 20, under stones along path to
waterfall, leg. S. Taiti and C. Rossano, 2 May 2004 (MZUF 9508).
Description
Maximum length: ♂, 11 mm; ♀, 15 mm. Body outline as in Figure 18A. Brown
colour with numerous yellowish muscle spots; a round pale spot at the base of pereon
epimera in the frontal half of the segment; an elongated pale spot in the middle of
pereonites and second or third to fifth pleonite; males darker than females. Back
smooth with some scattered pointed scale-setae (Figure 18B); a distinct sulcus mar-
ginalis at the sides of pereon epimera with numerous gland pores along its whole
length (Figure 18G); numerous gland pores scattered on the whole dorsal surface of
the body; noduli laterales clearly visible, more or less at the same distance from the
lateral margin of the pereonites, b/c and d/c co-ordinates as in Figure 18C. Cephalon
(Figure 18D–F)with no suprantennal line, frontal line straight; very small lateral
lobes bent downwards and not protruding frontwards; eye with about 25 ommatidia.
Pereonites 1–3 with posterior margin regularly convex; pereonite 4 with posterior
margin straight; pereonites 5–7 with posterior corners pointing backwards. Pleonites
3–5 with distinct but short posterior points (Figure 18H). Telson triangular with
distinctly concave sides (Figure 18H). Antennule (Figure 18I) with first article longer
than second and third; third article with a tuft of elongated aesthetascs at apex.
Antenna (Figure 19A) reaching back the posterior margin of pereonite 3; fifth article
of peduncle slightly curved, as long as flagellum; first flagellar article about 1.6 as
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Figure 18. Soteriscus gibbosus sp. nov. from St. 14, paratype ♀: (A) adult specimen, dorsal
view; (B) dorsal scale-seta; (C) co-ordinates of noduli laterals; (D) cephalon, dorsal view; (E)
cephalon, frontal view; (F) cephalon and pereonite 1, lateral view; (G) pereonite 7, right side;
(H) pleonites 4, 5, telson and uropods; (I) antennule.
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Figure 19. Soteriscus gibbosus sp. nov. from St. 14, paratype ♂: (A) antenna. Paratype ♀: (B)
left mandible; (C) right mandible; (D) maxillule; (E) maxilla; (F) maxilliped; (G) uropod,
lateral view.
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Figure 20. Soteriscus gibbosus sp. nov. from St. 14, paratype ♂: (A) pereopod 1; (B) pereopod
7; (C) genital papilla and pleopod 1; (D) pleopod 2; (E) pleopod 3 exopod; (F) pleopod 4
exopod; (G) pleopod 5 exopod.
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long as second. Mandibles (Figure 19B,C) with molar penicil dichotomized and a
line of several free penicils. Maxillule outer branch with 4 + 6 teeth (3 slightly cleft);
inner branch with a distinct posterior point and two long and thin penicils
(Figure 19D). Maxilla (Figure 19E) bilobate with setose apex, inner lobe quadran-
gular, much smaller than outer one; two long setae on the margin between the two
lobes. Maxilliped (Figure 19F) endite with two small triangular setae on distal margin
and no penicil; first article of palp with two strong setae. Pleopod 1 and 2 exopods
with monospiracular covered lungs. Uropod (Figure 18H) with a triangular depres-
sion on protopodal outer margin; exopod almost twice as long as endopod; endopod
proximally inserted.
Male: Carpus of pereopod 1 (Figure 20A) to 3 with a brush of pointed setae
increasing in length distally. Pereopod 7 (Figure 20B) ischium with straight sternal
margin and a longitudinal depression in the middle of the rostral surface; merus with
a distinct hump on the posterior half of tergal margin. Pleopod 1 (Figure 20C)
exopod with large medial lobe about twice as long as wide, with largely rounded
apex bearing a line of short setae; endopod with distal part with almost parallel sides
and a tuft of short setae at apex. Pleopod 2 (Figure 20D) endopod slightly longer than
exopod. Pleopod 3–5 exopods as in Figure 20E–G.
Etymology
From the Latin ‘gibbosus’= having a hump. The name refers to the male pereopod 7
merus which shows a distinct hump on the posterior half of tergal margin.
Remarks
At present the genus Soteriscus includes 15 species from Atlantic islands (Madeira
Archipelago, Canary Islands and Cape Verde), northern Morocco, northern Algeria
and southern Spain (Schmalfuss 2003). Three species have been recorded in north-
eastern Africa (Vandel 1956b,1958a,1960b): S. gaditanus, S. virescens (Budde-Lund,
1885) and S. fuscovariegatus (Lucas, 1849). Of these species, only S. gaditanus was
recorded from the Rif region (Vandel 1956b,1958a; Achouri et al. 2008a,2008c), but
this species has not been collected by us. Re-examination of the material identified as
S. gaditanus from the Rif region is necessary to confirm its occurrence.
Soteriscus gibbosus differs from all the other species in the genus in having a
distinct hump on the male pereopod 7 merus. In having the male pleopod 1 exopod
with a large medial lobe, the news species shows affinities with S. gaditanus and S.
fuscovariegatus, but in both these two species the medial lobe is distinctly more
slender. It also differs from the former in having a broadly rounded instead of
triangular apical part of the medial lobe of the male pleopod 1 exopod (see
Figure 2B in Vandel 1956b), and in the thicker and shorter uropodal exopods (see
Figure 2A in Vandel 1956b); from the latter in having shorter frontal lateral lobes and
longer and more slender uropodal exopods (see Figure 3A, B in Vandel 1956b).
Soteriscus laouensis sp. nov.
(Figures 21 and 22)
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Figure 21. Soteriscus laouensis sp. nov. from St. 16, paratype ♀: (A) adult specimen, dorsal
view; (B) dorsal scale-seta; (C) co-ordinates of noduli laterals; (D) cephalon, dorsal view; (E)
cephalon, frontal view; (F) cephalon and pereonite 1, lateral view; (G) pereonite 7, right side;
(H) pleonites 4, 5, telson and uropods; (I) antennule; (J) antenna; (K) uropod, lateral view.
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Figure 22. Soteriscus laouensis sp. nov. from St. 16, paratype ♂: (A) pereopod 1; (B) pereopod
7; (C) genital papilla and pleopod 1; (D) pleopod 2; (E) pleopod 3 exopod; (F) pleopod 4
exopod; (G) pleopod 5 exopod.
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Material examined
Holotype: ♂, St. 16, along road margin under stones, leg. S. Taiti, 28 April 2004
(MZUF 9509). Paratypes: 1 ♂♂,35♀♀, same data as holotype (MZUF 9509).
Description
Maximum length: ♂,13mm;♀, 16 mm. Body enlarged, outline as in Figure 21A.
Colour: male brown-grey with the usual yellowish muscle spots; female light
brown with a marbled pattern, two darker spots per side on the anterior part of
pereonites; antennae uniformly grey; pereopods pale with numerous dark spots;
pleopodal exopods dark. Back smooth with some scattered short triangular scale-
setae (Figure 21B); a distinct sulcus marginalis on lateral margins of pereon
epimera with numerous gland pores along its whole length (Figure 21G); numer-
ous gland pores scattered on the whole dorsal surface of the body; noduli laterales
clearly visible, inserted more or less at the same distance from the lateral margin
of the pereonites, b/c and d/c co-ordinates as in Figure 21C. Cephalon
(Figure 21D–F) with no suprantennal line, frontal line straight; very small lateral
lobes bent downwards and not protruding frontwards; eye with about 26 omma-
tidia. Pereonites 1–3 with posterior margin regularly convex; pereonite 4 with
posterior margin straight; pereonites 5–6 with posterior corners pointing back-
wards, pereonite 7 with acute posterior corners and slightly sinuous posterior
margin at sides. Pleonites 3–5 with well-developed falciform posterior points
(Figure 21H). Telson triangular with distinctly concave sides (Figure 21H).
Antennule (Figure 21I) with first article longer than second and third; third article
with a short triangular point and a tuft of elongated aesthetascs at apex. Antenna
(Figure 21J) reaching back posterior margin of pereonite 3; fifth article of ped-
uncle almost as long as flagellum; first flagellar article about 1.5 longer than
second. Buccal pieces as in the preceding species. Pleopodal exopods 1 and 2
with monospiracular covered lungs. Uropod (Figure 21K) with a triangular
depression on protopodal outer margin; exopod about twice as long as endopod;
endopod proximally inserted.
Male: Carpus and distal part of merus of pereopod 1 (Figure 22A), pereopod 2 and,
to a lesser extent, pereopod 3 with a brush of pointed setae. Pereopod 7 (Figure 22B)
ischium with slightly convex sternal margin and a longitudinal depression and a
setose area on rostral surface; merus elongated, without peculiar structures.
Pleopod 1 (Figure 22C) exopod with long medial lobe almost three times as long as
wide, with some short setae along its margin and a broadly rounded apex; endopod
with distal part with almost parallel sides and a tuft of short setae at apex. Pleopod 2
(Figure 22D) endopod distinctly longer than exopod. Pleopod 3–5 exopods as in
Figure 22E–G.
Etymology
The species is named after the Oued Laou basin, where the specimens were
collected.
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Remarks
In having the male pleopod 1 exopod with a long medial lobe S. laouensis is similar to
S. gaditanus, S. fuscovariegatus and S. gibbosus sp. nov. It is readily distinguishable from
S. gaditanus in having the male pleopod 1 exopod with broadly rounded instead of
triangular apical part and shorter uropodal exopods; from S. fuscovariegatus in the less
protruding lateral lobes of cephalon (see Figure 3A in Vandel 1956b) and comparatively
longer and thinner uropods; and from S. gibbosus in lacking the hump on the male
pereopod 7 merus and distinctly thinner medial lobe of the male pleopod 1 exopod.
Genus Porcellio Latreille, 1804
Porcellio pseudornatus sp. nov.
(Figures 23–25)
Material examined
Holotype: ♂, St. 5, under stones near beach, leg. S. Taiti and C. Rossano, 1 May
2004 (MZUF 9510). Paratypes: 7 ♂♂,10♀♀, same data as holotype (MZUF 9510);
1♂,1♀, same data as holotype (SMNS 15674); 1 ♂,3♀♀, St. 2, under stones at base
of cliff, leg. S. Taiti and C. Rossano, 1 May 2004 (MZUF 9511); 1 ♀, St. 7, leg. S.
Taiti, 29 April 2004 (MZUF 9512); 2 ♂♂,2♀♀, St. 9, under stones in meadow, leg.
S. Taiti, 28 April 2004 (MZUF 9513); 1 ♂,1♀, St. 14, Phillyrea wood, leg. S. Taiti,
27 April 2004 (MZUF 9514); 1 ♂, same locality, leg. S. Taiti and C. Rossano, 27
September 2005 (MZUF 9515); 3 ♂♂,1♀, St. 16, along road margin under stones,
leg. S. Taiti, 28 April 2004 (MZUF 9516); 2 ♀♀, St. 17, leg. S. Taiti, 29 April 2004
(MZUF 9517).
Other material examined
1♂,1♀, Al Hoceima, near road under stones, leg. C. Rossano, 4 May 2004 (MZUF
9518).
Description
Maximum length: ♂, 22 mm; ♀, 23 mm. Body outline as in Figure 23A. Colour
brown-grey with pale marginal parts of frontal lateral lobes and epimera of pereon
and pleon; sometimes with two paramedian yellowish round spots on pereonites 5–7
and pleonites 2–4; antennae uniformly grey, pereopods and pleopod 1–2 exopods
pale, pleopod 3–5exopods dark. Dorsal surface of cephalon, pereon and pleon
distinctly granulated with many scattered short triangular scale-setae (Figure 22B);
numerous gland pores in rounded fields disposed near the anterior corner of pereonite
1 and more or less in the middle of pereonites 2–7 close to the lateral margins of the
segments (Figure 22A,E); noduli laterales small, the ones on pereonites 1–4 inserted
about twice more distant from the lateral margin than those on pereonites 5–7
(Figure 23A). Cephalon (Figure 22C,D) with no suprantennal line; large rounded
lateral lobes obliquely bent downwards and distinctly protruding frontwards, median
lobe widely rounded, often slightly incised at the apex; eye with about 30 ommatidia.
Pereonites 1–3 with hind margin slightly concave at sides, more distinct on pereonite
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Figure 23. Porcellio pseudornatus sp. nov. from St. 5, paratype ♀: (A) adult specimen, dorsal
view; (B) dorsal scale-seta; (C) cephalon, dorsal view; (D) cephalon, frontal view; (E) pereonite
3, right side. Paratype ♂: (F) pleon, telson and uropods; (G) antennule; (H) antenna.
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1; pereonites 3–7 with posterior corners bent backwards and progressively more
acute. Pleonites 3–5 with falciform epimera (Figure 23A,F); tips of pleonite 5
epimera reaching 2/3 of telson length. Telson (Figure 23F) about twice as wide as
Figure 24. Porcellio pseudornatus sp. nov. from St. 5, paratype ♀: (A) left mandible; (B) right
mandible; (C) maxillule; (D) maxilla; (E) maxilliped.
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Figure 25. Porcellio pseudornatus sp. nov. from St. 5, paratype ♂: (A) pereopod 1; (B)
pereopod 7; (C) genital papilla and pleopod 1; (D) pleopod 2; (E) pleopod 3 exopod; (F)
pleopod 4 exopod; (G) pleopod 5 exopod.
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long with a triangular distal part. Antennule (Figure 23G) with first article distinctly
longer than second and third; third article with a tuft of short aesthetascs near the
apex. Antenna (Figure 23H) reaching back or slightly surpassing the posterior margin
of pereonite 3; fifth article of peduncle shorter than flagellum; first flagellar article
about twice as long as second. Mandibles (Figure 24A,B) with molar penicil
dichotomized and a line of eight free penicils. Maxillule (Figure 24C) outer branch
with 4 + 6 teeth, all simple; inner branch with a distinct posterior point and two long
and thin penicils. Maxilla (Figure 24D) bilobate with setose apex, inner lobe quad-
rangular, much smaller than outer one; two long setae on the margin between the two
lobes. Maxilliped (Figure 24E) endite with two small triangular setae on distal margin
and no penicil; first article of palp with two strong setae. Pleopodal exopods 1 and 2
with monospiracular covered lungs (Figure 25C,D). Uropod (Figure 23A,F)witha
triangular depression on protopodal outer margin, not visible in dorsal view.
Male: Uropodal exopods (Figure 23F) flattened, elongated and much longer than in
females (4 times as long as wide in males, less than 3 times in females). Merus and
carpus of pereopod 1 (Figure 25A) to 4 and, to a lesser extent, 5 with a brush of
pointed setae. Pereopod 7 (Figure 25B) ischium distally enlarged with a transversal
depression and a setose area on rostral surface; sternal margin slightly convex; carpus
with a large rounded lobe on distal half of tergal margin. Pleopod 1 (Figure 25C)
exopod with long medial lobe with parallel sides, oblique and sinuous distal margin
with three strong setae at apex, and some short setae along the medial margin;
endopod with distal part straight, rounded and setose apex. Pleopod 2
(Figure 25D) exopod triangular and slightly shorter than endopod. Pleopod 3–5
exopods as in Figure 25E–G.
Etymology
From the stem of the Greek ‘Pseudes’= false, erroneous + ornatus. The species name
refers to the similarity with Porcellio ornatus Milne-Edwards, from south-eastern
Spain.
Remarks
According to Vandel (1958a), four species of the Rif-Betic group of Porcellio occur
in the Rif region: P. ornatus Milne-Edwards; P. wagneri Brandt, 1841;
P. hoffmannseggii Brandt, 1833; and P. echinatus Lucas, 1849. The specimens here
examined do not seem to belong to any of these four species, nor to any other species
in this group, even if they are strictly related to P. ornatus and P. wagneri. Specimens
of P. ornatus collected from the type locality in south-eastern Spain (many ♂♂ and
♀♀, Cartagena, Murcia, leg. M. Rizzotti Vlach, 4 August 1981, MZUF 9594) have
been examined and are here illustrated (Figures 26 and 27) for comparison with our
specimens from the Oued Laou basin. Porcellio pseudornatus differs from P. ornatus
in the dorsal body surface more granulated, wider dorsal scale-setae, posterior margin
of pereonites 1–3 more concave at the sides, thinner antennae, much longer
male uropodal exopods and more pronounced rounded lobe on the male pereopod
7 carpus. According to the figures provided by Lucas (1849, plate 6, Figures 6A–C);
for P. wagneri originally described from Algeria, the new species differs in having the
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Figure 26. Porcellio ornatus from Cartagena, ♀: (A) adult specimen, dorsal view; (B) dorsal
scale-seta; (C) cephalon, dorsal view; (D) cephalon, frontal view; (E) pereonite 4, right side. ♂:
(F) pleon, telson and uropods; (G) antennule; (H) antenna.
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Figure 27. Porcellio ornatus from Cartagena, ♂: (A) pereopod 1; (B) pereopod 7; (C)
pleopod 1; (D) pleopod 2; (E) pleopod 3 exopod; (F) pleopod 4 exopod; (G) pleopod 5
exopod.
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male uropodal exopods much wider (4 times as long as wide vs. 7.5 times in P.
wagneri). The material of P. ornatus and P. wagneri from the Rif studied by
Vandel (1958a) needs to be re-examined to confirm that also these two species are
present in the area.
Porcellio hoffmannseggii hoffmannseggii Brandt, 1833
Porcellio Hoffmannseggi; Budde-Lund, 1879, p. 3; Budde-Lund, 1885, p. 108;
Dollfus, 1896, p. 534; Dollfus, 1898,p.134
Porcellio (Porcellio) Hoffmannseggü; (sic) Arcangeli, 1935,p.14
Porcellio hoffmannseggi tamaricis; Verhoeff, 1937, p. 306, Figures 8,9
Porcellio (Rogopus) Hoffmannseggii tamaricis; Paulian de Félice, 1939, p. 197,
Figure 34
Porcellio hoffmannseggi; Vandel, 1946, p. 327; Schmölzer, 1971,p.46
Porcellio hoffmannseggi hoffmannseggi; Vandel, 1958a, p. 129; Schmalfuss, 1987,p.
285, Figures 1–25
Porcellio hoffmannseggii; Schmalfuss, 2003, p. 226; Achouri et al., 2008c, pp. 76–78.
Material examined
1♂,5♀♀, St. 3, under stones, leg. S. Taiti and C. Rossano, 1 October 2005 (MZUF
9519); 1 ♀, St. 4, dune under stones, leg. S. Taiti and C. Rossano, 1 May 2004
(MZUF 9520); 1 ♂, St. 14, Phillyrea wood, leg. S. Taiti and C. Rossano, 27
September 2005 (MZUF 9521); 1 ♂, same data (SMNS 15675); 4 ♀♀, St. 23,
under stones, leg. S. Taiti, 30 April 2004 (MZUF 9522); 1 ♀, St. 25, under stones
in meadow, leg. S. Taiti, 30 April 2004 (MZUF 9523).
Previous Rif records
Taxdirt, west of Melilla; Djebel Kerker, south of Melilla; Tizi Ifri, south of Targuist;
Beni-Bufrah, Tétouan; Benitez, near Ceuta (Vandel 1958a); Chechauen; Cap Spartel;
15 km south of Ceuta (Schmalfuss 1987); Oued Laou region (Achouri et al. 2008c).
Distribution
Portugal, southern Spain and northern Morocco.
Remarks
For diagnostic characters of P. hoffmannseggii hoffmanseggii see Schmalfuss (1987,p.
285, Figures 1–25).
Porcellio flavocinctus Budde-Lund, 1885
(Figures 28 and 29)
Porcellio (Euporcellio) flavocinctus; Paulian de Félice, 1939, p. 199
Porcellio flavocinctus; Dollfus, 1896, p. 534; Vandel, 1958a, p. 129
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Figure 28. Porcellio flavocinctus from St. 10, ♀: (A) adult specimen, dorsal view; (B) dorsal
scale-seta; (C) cephalon, dorsal view; (D) cephalon, frontal view; (E) pereonite 4, left side. ♂:
(F) pleon, telson and uropods; (G) antennule; (H) antenna.
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Figure 29. Porcellio flavocinctus from St. 10, ♂: (A) pereopod 1; (B) pereopod 7; (C) genital
papilla and pleopod 1; (D) pleopod 2; (E) pleopod 3 exopod; (F) pleopod 4 exopod; (G)
pleopod 5 exopod.
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Material examined
3♂♂,6♀♀, St. 10, leg. S. Taiti and C. Rossano, 26 April 2004 (MZUF 9524).
Remarks
These specimens are identified as P. flavocinctus after comparison with the redescrip-
tion of this species provided by Arcangeli (1936, p. 190, Figures 1–7). The main
characters of the specimens from the Oued Laou valley are here illustrated (Figures
28 and 29) and they fit well those of P. flavocinctus provided by Arcangeli. The record
of Budde-Lund (1885) from El Araisch (= Larache) in northern Morocco was
considered doubtful by Arcangeli (1936). Our record from the Oued Laou valley
together with those by Vandel (1958a) from the Rif confirms that this species is also
present in this part of north-western Africa.
Previous Rif records
Melilla; Mt Gurugu and Ixmoart, near Melilla; Sidi Yacoub el Basi, near Peñón de
Velez, Tétouan; Benitez, near Ceuta (Vandel 1958a).
Distribution
Southern Spain, southern Portugal and northern Morocco.
Porcellio laevis (Latreille, 1804)
Porcellio laevis; Verhoeff, 1938, p. 67; Vandel, 1958a, p. 129; Achouri et al., 2008c,
pp. 76–78; Achouri et al., 2008a, p. 129
Porcellio (Mesoporcellio) laevis; Arcangeli, 1932, p. 225; Paulian de Félice, 1939,
p. 210
Material examined
1♂,1♀, St. 3, under stones, leg. S. Taiti and C. Rossano, 1 October 2005 (MZUF
9525); 1 ♂, St. 4, dune under stones, leg. S. Taiti and C. Rossano, 1 May 2004
(MZUF 9526); 2 ♂♂,1♀, St. 5, beach under stones, leg. S. Taiti and C. Rossano, 1
May 2004 (MZUF 9527); 1 ♂,2♀♀, St. 6, near well and wheat field, leg. S. Taiti and
C. Rossano, 26 April 2004 (MZUF 9528).
Previous Rif records
Granja de la Muluya; Benitez, near Ceuta (Vandel 1958a); Oued Laou region
(Achouri et al. 2008a,2008c).
Distribution
Europe and northern Africa, introduced to many parts of the world.
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Remarks
For diagnostic characters of P. laevis see Vandel (1962, p. 684, Figures 331, 332).
Porcellio riffensis Caruso and Di Maio, 1990
Porcellio hoffmannseggi nemethi; Vandel, 1958a, p. 129
Porcellio riffensis Caruso and Di Maio, 1990, p. 204, Figures 6–10; Schmalfuss, 2003,
p. 234
Material examined
1♂,4♀♀, 3 juvs, St. 7, leg. S. Taiti and C. Rossano, 29 September 2005 (MZUF
9529); 1 ♂,2♀♀, St. 10, leg. S. Taiti and C. Rossano, 26 April 2004 (MZUF 9530); 6
♂♂,2♀♀, St. 12, leg. C. Rossano, 26 April 2004 (MZUF 9531); 1 ♂,7♀♀, St. 14,
Phillyrea wood, leg. S. Taiti and C. Rossano, 27 September 2005 (MZUF 9532); 6
♂♂,7♀♀, St. 16, along road margin under stones, leg. S. Taiti, 28 April 2004
(MZUF 9533); 1 ♀, St. 18, cork-oak wood, leg. S. Taiti, 29 April 2004 (MZUF 9534);
9♂♂,12♀♀, St. 19, leg. S. Taiti, 29 April 2004 (MZUF 9535).
Previous Rif records
Einzoren, south of Villa Sanjurjo; Issaguen (Vandel 1958a; Caruso and Di Maio
1990); Bab Taza; Cedreta, 17 km from Ketama; Bab Berred (Caruso and Di Maio
1990).
Distribution
Northern Morocco.
Remarks
For diagnostic characters of P. riffensis see Caruso and Di Maio (1990, p. 204,
Figures 6–10).
Porcellio echinatus Lucas, 1849
Porcellio (Euporcellio) echinatus; Paulian de Félice, 1939, p. 198
Porcellio echinatus; Dollfus, 1896, p. 532; Vandel, 1958a, p. 129; Achouri et al.,
2008c, pp. 76–78
Material examined
2♂♂,2♀♀, St. 6, near well and wheat field, leg. S. Taiti and C. Rossano, 26 April
2004 (MZUF 9536); 3 ♂♂,3♀♀,St. 7, leg. S. Taiti, 29 April 2004 (MZUF 9537); 1
♂,1♀, same locality, leg. S. Taiti and C. Rossano, 29 September 2005 (MZUF
9538); 2 ♀♀, St. 9, under stones in meadow, leg. S. Taiti, 28 April 2004 (MZUF
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9539); 1 ♀, St. 10, leg. S. Taiti and C. Rossano, 26 April 2004 (MZUF 9540); 1 ♂,4
♀♀, St. 14, Phillyrea wood, leg. S. Taiti, 27 April 2004 (MZUF 9541); 1 ♂,2♀♀,
same locality, leg. S. Taiti and C. Rossano, 27 September 2005 (MZUF 9542); 1 ♂,3
♀♀, St. 19, leg. S. Taiti, 29 April 2004 (MZUF 9543); 2 ♂♂,3♀♀, St. 20, under
stones along path to waterfall, leg. S. Taiti and C. Rossano, 2 May 2004 (MZUF
9544); 1 ♂,2♀♀, St. 26, cork-oak wood, leg. S. Taiti, 30 April 2004 (MZUF 9545).
Previous Rif records
Taxdirt, west of Melilla (Vandel 1958a); Oued Laou region (Achouri et al. 2008c).
Distribution
Southern Portugal, southern Spain, northern Morocco, and north-western Algeria.
Remarks
For diagnostic characters of P. echinatus see Vandel (1946, p. 304, Figures 125–129).
Porcellio humberti Paulian de Félice, 1939
(Figures 30 and 31)
Porcellio Humberti Paulian de Félice, 1939, p. 210, Figures 70–74
Porcellio humberti; Vandel, 1958a, p. 129; Vandel, 1958c, p. 445, Figures 1–3
Material examined
1♀, St. 4, leg. S. Taiti and C. Rossano, 1 May 2004 (MZUF 9546); 1 ♀, St. 7, leg. S.
Taiti, 29 April 2004 (MZUF 9547); 5 ♀♀, 1 juv., same locality, leg. S. Taiti and C.
Rossano, 29 September 2005 (MZUF 9548); 1 ♂,3♀♀, St. 12, leg. C. Rossano, 26
April 2004 (MZUF 9549); 2 ♀♀, 3 juvs, St. 14, Phillyrea wood, leg. S. Taiti and C.
Rossano, 27 September 2005 (MZUF 9550); 5 ♂♂,6♀♀, St. 15, sieved ground near
stream, leg. S. Taiti, 27 April 2004 (MZUF 9551); 2 ♀♀, St. 16, along road margin
under stones, leg. S. Taiti, 28 April 2004 (MZUF 9552); 1 ♀, St. 18, cork-oak wood,
leg. S. Taiti, 29 April 2004 (MZUF 9553); 5 ♂♂,11♀♀, St. 19, leg. S. Taiti, 29 April
2004 (MZUF 9554); 1 ♀,St. 20, under stones along path to waterfall, leg. S. Taiti and
C. Rossano, 2 May 2004 (MZUF 9555); 4 ♂♂,6♀♀, St. 21, leg. S. Taiti and C.
Rossano, 30 September 2005 (MZUF 9556); 2 ♀♀, St. 24, cork-oak wood, leg. S.
Taiti, 29 April 2004 (MZUF 9557); 1 ♂,2♀♀, St. 25, under stones in meadow, leg. S.
Taiti, 30 April 2004 (MZUF 9558); 3 ♂♂,7♀♀, St. 26, cork-oak wood, leg. S. Taiti,
30 April 2004 (MZUF 9559).
Previous Rif records
Mt Gurugu, near Melilla; Tétouan; Benitez, near Ceuta (Vandel 1958a,1958c).
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Distribution
Southern Spain and Morocco.
Remarks
Porcellio humberti was described by Paulian de Félice (1939) on specimens from Jebel
Ayachi, in the High Atlas Range. Vandel (1958c) redescribed and partially illustrated
Figure 30. Porcellio humberti from St. 19, ♂: (A) adult specimen, dorsal view; (B) dorsal scale-
seta; (C) cephalon, dorsal view; (D) cephalon, frontal view; (E) pereonite 4, right side; (F)
pleonites 4, 5, telson and uropods; (G) antennule; (H) antenna.
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Figure 31. Porcellio humberti from St. 19, ♂: (A) pereopod 1; (B) pereopod 7; (C) genital
papilla and pleopod 1; (D) pleopod 2; (E) pleopod 3 exopod; (F) pleopod 4 exopod; (G)
pleopod 5 exopod.
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the species, and added several localities in Morocco and southern Spain. A full set of
figures of this species that integrate Vandel’s redescription is here provided on speci-
mens from St. 19.
Family ARMADILLIDIIDAE (Brandt, 1833)
Genus Eluma Budde-Lund, 1885
Eluma caelata (Miers, 1877)
(Figures 32 and 33)
Material examined
1♂,1♀, St. 7, leg. S. Taiti, 29 April 2004 (MZUF 9560); 2 ♂♂, St. 8, leg. S. Taiti
and C. Rossano, 28 April 2004 (MZUF 9561); 1 ♀, same locality and collectors, 28
September 2005 (MZUF 9562); 3 ♀♀, St. 10, leg. S. Taiti and C. Rossano, 26 April
2004 (MZUF 9563); 1 ♂,1♀, St. 11, leg. S. Taiti, 27 April 2004 (MZUF 9564); 4
♂♂,29♀♀, St. 18, cork-oak wood, leg. S. Taiti, 29 April 2004 (MZUF 9565); 7 ♂♂,
2♀♀, St. 20, under stones along path to waterfall, leg. S. Taiti and C. Rossano, 2
May 2004 (MZUF 9566); 1 ♀, St. 22, leg. S. Taiti and C. Rossano, 30.IX.2005
(MZUF 9567).
Distribution
Southern British Isles, western France, Portugal, Spain, north-western Africa, Canary
Islands, Madeira, and Azores. Introduced to Tasmania and French Guiana
(Schmalfuss 2003). New record for the Rif region.
Remarks
The synonymy of Eluma purpurascens Budde-Lund, 1885 with Armadillidium caela-
tum Miers, 1877 was definitely established by Schmalfuss (2003) after examination of
the type material of the latter species. The main characters of this species are
illustrated here (Figures 32 and 33) on specimens from St. 18 (Fahsa) for comparison
with the new species Eluma praticola described below.
Eluma praticola sp. nov.
(Figures 34–36)
Material examined
Holotype: ♂, St. 25, under stones in meadow, leg. S. Taiti, 30 April 2004 (MZUF
9568). Paratypes: 9 ♂♂,19♀♀, same data as holotype (MZUF 9568).
Description
Maximum length: ♂, 5.5 mm; ovigerous ♀,7 mm. Body very convex, able to roll up
into a ball, euspheric type (Figure 34A). Colour brown with paler epimera of
pereonites 2–7. Dorsal surface without ornamentation and with numerous round
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Figure 32. Eluma caelata from St. 18, ♀: (A) cephalon and pereonites 1, 2, lateral view; (B)
cephalon and pereonite 1, frontal view; (C) pleonite 5, telson and uropods; (D) pereonite 7 with
noduli laterales. ♂: (E) pereopod 7, caudal surface; (F) pereopod 7 ischium and merus, caudal
surface; (G) pereopod 7 ischium and merus, rostral surface.
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Figure 33. Eluma caelata from St. 18, ♂: (A) dorsal scale-seta; (B) pereopod 1; (C) genital
papilla and pleopod 1; (D) pleopod 2; (E) pleopod 3 exopod; (F) pleopod 4 exopod; (G)
pleopod 5 exopod.
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Figure 34. Eluma praticola sp. nov. from St. 25, paratype ♀: (A) adult specimen, lateral view;
(B) cephalon and pereonites 1, 2, lateral view; (C) cephalon and pereonite 1, frontal view; (D)
pereonite 7 with noduli laterales; (E) pleonite 5, telson and uropods. Paratype ♂: (F) pereopod
7, caudal surface; (G) pereopod 7 ischium and merus, caudal surface.
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Figure 35. Eluma praticola sp. nov. from St. 25, paratype ♂: (A) dorsal scale-seta; (B) right side
of pleonites 1 and 2, ventral view; (C) antennule; (D) antenna. Paratype ♀: (E) left mandible;
(F) right mandible; (G) maxillule; (H) maxilla; (I) maxilliped.
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Figure 36. Eluma praticola sp. nov. from St. 25, paratype ♂: (A) uropod; (B) pereopod 1; (C)
genital papilla and pleopod 1; (D) pleopod 2; (E) pleopod 3 exopod; (F) pleopod 4 exopod; (G)
pleopod 5 exopod.
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pits and tiny pointed scale-setae (Figure 35A); one line of noduli laterales per side far
from the lateral margin of pereonites 1–6, two noduli laterales per side on pereonite 7
(Figure 34D); no visible gland pores. Cephalon (Figure 34B,C) with a wide trian-
gular scutellum distinctly separated from and not bent over vertex; no postscutellar
line; antennary lobes quadrangular, directed frontwards; eye consisting of a single
large ocellus. Pereonite 1 (Figures 34B,35B) with a flattened lateral margin and a
schisma on posterior corners; inner lobe of schisma rounded, more protruding back-
wards than the outer one; a small triangular lobe on ventral surface; posterior margin
slightly sinuous at sides. Pereonite 2 (Figure 35B) with rounded epimera, a small
triangular ventral lobe and straight posterior margin. Pereonite 3 with rounded
epimera and straight posterior margin; pereonites 4–7 with quadrangular epimera
and straight posterior margins. Pleonites 3–5 with rectangular epimera, slightly
divergent. Telson (Figure 34E) triangular, almost 1.5 as wide as long, with straight
sides and broadly rounded apex. Antennule (Figure 35C) of three articles, second
article much shorter than first and third, third article with a tuft of superimposed
aesthetascs subapically. Antenna (Figure 35D) short and stout with flagellum slightly
shorter than fifth article of peduncle, second flagellar article almost three times longer
than first and bearing two rows of aesthetascs. Mandibles (Figure 35E,F) with
semidichotomized molar penicil and four free penicils; right mandible with one
penicil and left mandible with two penicils on the hairy lobe. Maxillule
(Figure 35G) outer lobe with 4 + 6 (5 cleft) teeth; inner lobe with two subequal
penicils and a small triangular distal point. Maxilla (Figure 35H) apically setose, with
quadrangular inner lobe, much smaller than rounded outer lobe. Maxilliped
(Figure 35I) with quadrangular endite bearing three short triangular setae on distal
margin and a longer subapical seta near the inner corner; basal article of palp with
two long setae. Uropod (Figures 34E,36A) flattened; exopod about twice as wide as
long, with concave distal margin; endopod distinctly more protruding backwards
compared with exopod.
Male: Pereopod 1 (Figure 36B) with a line of pointed setae on sternal margin of
carpus and, to a lesser extent, merus. Pereopod 7 (Figure 34F,G) ischium distally
with a ridge on caudal surface, sternal margin slightly concave with numerous long
setae; merus with no ridges or lobes, sternal margin slightly convex with some long
setae. Pleopod 1 (Figure 36C) exopod with a short, rounded medial lobe; endopod
with distal part pointed and bent outwards. Pleopod 2 (Figure 36D) exopod trian-
gular and slightly shorter than endopod. Pleopod 3–5 exopods as in Figure 36E–G.
Etymology
From the Latin pratum = meadow + stem of colere = to live. The name refers to the
habitat where the specimens have been collected.
Remarks
Up to date the genus Eluma included only two species, the widespread E. caelata and
E. tuberculata Cruz, 1991 from Fatima, Portugal. The new species is similar to E.
caelata from which it differs in smaller size (maximum length in ovigerous females
7 mm vs 16 mm), the dorsal surface with shorter and less numerous scale-setae
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(compare Figure 34A–Eand Figure 32A–D), shorter uropodal exopod (compare
Figures 34E and Figure 32C), male pereopod 7 without a basal triangular lobe on
merus and less sharp ridge on the caudal surface of the ischium (compare Figure 34F,
Gand Figure 32E–G), and pleopod 1 exopod with a shorter and round medial lobe
(compare Figure 36C and Figure 33C).
Eluma praticola is readily distinguishable from E. tuberculata in the dorsal surface
without ornamentation and lateral margin of pereonite 1 not grooved. No compar-
ison is possible with the male characters since E. tubercolata was described on female
specimens (Cruz 1991).
Genus Armadillidium Brandt, 1833
Armadillidium album Dollfus, 1887
Armadillidium album; Colombini et al., 2008,p.86
Material examined
2♂♂,1♀, St. 4, under logs, leg. S. Taiti and C. Rossano, 1 May 2004 (MZUF 9569).
Previous Rif records
Oued Laou (Colombini et al. 2008).
Distribution
Atlantic coast of Europe, western and central Mediterranean as far East as Greece.
Remarks
For diagnostic characters of A. album see Vandel (1962, p. 841, Figure 403) and
Schmalfuss (2008, p. 154, Figures 1–12).
Armadillidium granulatum Brandt, 1833
Armadillidium granulatum; Dollfus, 1896, p. 528; Paulian de Félice, 1939, p. 212;
Vandel, 1958a, p. 130; Achouri et al., 2008c, pp. 76–78; Achouri et al., 2008a,
p. 129
Material examined
Many ♂♂ and ♀♀, St. 2, under stones at base of cliff, leg. S. Taiti and C. Rossano, 1
May 2004 (MZUF 9570).
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Previous Rif records
Melilla; Taxdirt and Ixmoart, near Melilla; Sidi Yacoub el Basi, near Peñón de Velez;
Tétouan; Benitez, near Ceuta (Vandel 1958a); Oued Laou region (Achouri et al.
2008a,2008c).
Distribution
Littoral species distributed along the coasts of the Mediterranean, Atlantic coasts of
Morocco, Portugal, France, Madeira and the Azores.
Remarks
For diagnostic characters of A. granulatum see Vandel (1962, p. 796, Figure 383).
Armadillidium vulgare (Latreille, 1804)
Armadillidium vulgare; Dollfus, 1896, p. 530; Dollfus, 1898, p. 134; Verhoeff, 1938,p.
67; Paulian de Félice, 1939, p. 212; Vandel, 1958a, p. 130; Achouri et al., 2008c,
pp. 76–78; Colombini et al., 2008,p.86
Armadillidium cinereum; Arcangeli, 1932, p. 225
Material examined
1♀, St. 3, under stones, leg. C. Rossano, 28 April 2004 (MZUF 9571); 3 ♀♀, St. 4,
dune under stones, leg. S. Taiti and C. Rossano, 1 May 2004 (MZUF 9572); 1 ♂,3
♀♀, St. 5, beach under stones, leg. S. Taiti and C. Rossano, 1 May 2004 (MZUF
9573); 3 ♂♂, same locality, leg. S. Taiti and C. Rossano, 1 October 2005 (MZUF
9574); 1 ♂,9♀♀, St. 6, near well and wheat field, leg. S. Taiti and C. Rossano, 26
April 2004 (MZUF 9575); 1 ♂,1♀, St. 7, leg. S. Taiti and C. Rossano, 29 September
2005 (MZUF 9576); 1 ♀, St. 14, Phillyrea wood, leg. S. Taiti, 27 April 2004 (MZUF
9577); 5 ♂♂,8♀♀, St. 15, sieved ground near stream, leg. S. Taiti, 27 April 2004
(MZUF 9578); 1 ♀, St. 23, under stones, leg. S. Taiti, 30 April 2004 (MZUF 9579).
Previous Rif records
Benitez, near Ceuta (Vandel 1958a); Oued Laou region (Achouri et al. 2008c); Oued
Laou (Colombini et al. 2008).
Distribution
Cosmopolitan species of Mediterranean origin.
Remarks
For diagnostic characters of A. vulgare see Vandel (1962, p. 826, Figures 397, 398)
and Gruner (1966,p. 320, Figures 251, 252).
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Family ARMADILLIDAE Brandt
Genus Armadillo Latreille, 1802
Armadillo officinalis Duméril, 1816
Armadillo officinalis; Dollfus, 1896, p. 527; Paulian de Félice, 1939, p. 212; Vandel,
1958a, p. 130; Achouri et al., 2008c, pp. 76–78
Material examined
6♂♂,5♀♀, St. 2, under stones at base of cliff, leg. S. Taiti and C. Rossano, 1 May
2004 (MZUF 9580); 1 ♀, St. 7, leg. S. Taiti, 29 April 2004 (MZUF 9581); 6 ♂♂,5
♀♀, same locality, leg. S. Taiti and C. Rossano, 29 September 2005 (MZUF 9582); 5
♂♂,11♀♀, 2 juvs, St. 10, leg. S. Taiti and C. Rossano, 26 April 2004 (MZUF 9583);
1♂,5♀♀, St. 11, leg. S. Taiti, 27 April 2004 (MZUF 9584); 1 ♂,4♀♀, St. 12, leg. C.
Rossano, 26 April 2004; (MZUF 9585) 1 ♂,3♀♀, St. 14, Phillyrea wood, leg. S.
Taiti, 27 April 2004 (MZUF 9586); 3 ♂♂,3♀♀, 1 juv., same locality, leg. S. Taiti
and C. Rossano, 27 September 2005 (MZUF 9587); 9 ♂♂,5♀♀, 6 juvs, St. 15, sieved
ground near stream, leg. S. Taiti, 27 April 2004 (MZUF 9588); 6 ♂♂,9♀♀, St. 16,
along road margin under stones, leg. S. Taiti, 28 April 2004 (MZUF 9589); 2 ♂♂,9
♀♀, St. 19, leg. S. Taiti, 29 April 2004 (MZUF 9590); 4 ♀♀, St. 20, under stones
along path to waterfall, leg. S. Taiti and C. Rossano, 2 May 2004 (MZUF 9591); 2
♂♂,2♀♀, St. 21, leg. S. Taiti and C. Rossano, 30 September 2005 (MZUF 9592); 1
♀, St. 23, under stones, leg. S. Taiti, 30 April 2004 (MZUF 9593).
Previous Rif records
Ixmoart and Mt Gurugu, near Melilla; Isla de Alhucemas; Tétouan; Benitez, near
Ceuta (Vandel 1958a); Oued Laou region (Achouri et al. 2008c).
Distribution
Mediterranean basin, Morocco and Portugal.
Remarks
For diagnostic characters of A. officinalis see Vandel (1962, p. 855, Figures 408, 409).
Discussion
In the present study 34 species of terrestrial isopods are recorded from the Oued Laou
basin and summarized in Figure 37.Two genera (Paractenoscia and Soteriscus) and
seven species (Trichoniscus microphthalmus, Paractenoscia cavernicola, Bathytropa
rifensis, Soteriscus gibbosus, S. laouensis, Porcellio pseudornatus and Eluma praticola)
are described as new. Nine species (Ligia italica, Graeconiscus thermophilus, Ctenoscia
minima, Stenoniscus carinatus, Platyarthrus caudatus, P. parisii, Lucasius pallidus,
Porcellionides sexfasciatus lusitanus and Eluma caelata) represent new records for
the Rif region.
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Figure 37. Map of Oued Laou basin with distribution records of Oniscidean species. The
stations (St.) refer to those listed in Table 1. 1=Ligia italica;2=Tylos europaeus;
3=Trichoniscus microphthalmus sp. nov.; 4 = Graeconiscus thermophilus;5=Halophiloscia
couchii;6=Stenophiloscia glarearum;7=Ctenoscia minima;8=Paractenoscia cavernicola sp.
nov.; 9 = Bathytropa rifensis sp. nov.; 10 = Stenoniscus carinatus;11=Platyarthrus caudatus;
12 = Platyarthrus parisii;13=Leptotrichus panzerii;14=Agabiformius lentus;
15 = Agabiformius obtusus;16=Lucasius pallidus;17=Porcellionides pruinosus;
18 = Porcellionides sexfasciatus lusitanus;19=Porcellionides sp.; 20 = Soteriscus gibbosus sp.
nov.; 21 = Soteriscus laouensis sp. nov.; 22 = Porcellio pseudornatus sp. nov.; 23 = Porcellio
hoffmannseggii hoffmannseggii;24=Porcellio flavocinctus;25=Porcellio laevis;26=Porcellio
riffensis;27=Porcellio echinatus;28=Porcellio humberti;29=Eluma caelata;30=Eluma
praticola sp. nov.; 31 = Armadillidium album;32=Armadillidium granulatum;
33 = Armadillidium vulgare;34=Armadillo officinalis.
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Vandel (1958a) recorded 17 species from the Rif area, many of which have also
been encountered in the Oued Laou basin. Six of these species (Lucasius myrmeco-
philus, Soteriscus gaditanus, Porcellio ornatus, P. wagneri, Armadillidium djebalensis
Vandel, 1958b and A. pardoi Vandel, 1956a) are not present in the collection exam-
ined by us, but we must point out the taxonomic uncertainties concerning L. myrme-
cophilus (see remarks under L. pallidus) and the possible misidentification of P.
ornatus which probably has to refer to P. pseudornatus sp. nov.
In a more recent publication on the diversity of terrestrial isopods from the same
area considered in this study, Achouri et al. (2008c) listed 19 species, six of which
identified only at genus level (Porcellio sp. 1, Porcellio sp. 2, Porcellionides sp.,
Acaeroplastes sp. Philoscia sp. and Chaetophiloscia sp.). The material studied by
those authors should be re-examined to complete the identifications and confirm
the presence of the genera Acaeroplastes Verhoeff, 1918,Philoscia Latreille, 1804
and Chaetophiloscia Verhoeff, 1908 in the Rif area. Only three species (Lucasius
myrmecophilus, Soteriscus gaditanus and Porcellio lamellatus Budde-Lund, 1885)
recorded by the authors for the Oued Laou region were not encountered by us.
According to their present distributions, the 39 species identified with certainty
from the Rif region belong to the following biogeographical categories:
1. Widespread species (4)
Agabiformius lentus, Porcellionides pruinosus, Porcellio laevis and Armadillidium
vulgare: these species, all of Mediterranean origin, have been introduced into many
parts of the world and can be considered as cosmopolitan.
2. Mediterranean-Atlantic species (8)
Ligia italica, Tylos europaeus, Halophiloscia couchii, Stenophiloscia glarearum,
Leptotrichus panzerii, Porcellio lamellatus, Armadillidium album and A. granulatum
3. Mediterranean species (4)
Graeconiscus thermophilus, Leptotrichus panzerii, Agabiformius obtusus and
Armadillo officinalis
4. West-Mediterranean-Atlantic species (6)
Stenoniscus carinatus, Ctenoscia minima, Porcellionides sexfasciatus lusitanus,
Lucasius pallidus,Platyarthrus parisii and Eluma caelata
5. West-Mediterranean species (1)
Platyarthrus caudatus
6. Rif-Betic species (6)
Soteriscus gaditanus, Porcellio hoffmannseggii hoffmannseggii, P. flavocinctus, P.
echinatus, P. wagneri and P. humberti
7. Rif endemics (10)
Trichoniscus microphthalmus sp. nov., Paractenoscia cavernicola sp. nov.,
Bathytropa rifensis sp. nov., Soteriscus gibbosus sp. nov., S. laouensis sp. nov.,
Porcellio pseudornatus sp. nov., P. riffensis, Eluma praticola sp. nov.,
Armadillidium djebalensis and A. pardoi.
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About 59% of the species have wide distributions (Groups 1–5), all of
Mediterranean origin. Seven of these species (Ligia italica, Tylos europaeus,
Halophiloscia couchii, Stenophiloscia glarearum, Stenoniscus carinatus, Porcellio
lamellatus and Armadillidium album) are halophilic and widely distributed along the
sandy and rocky shores of the Mediterranean Sea and some also on the Atlantic
coasts of Africa and Europe.
The number of species with more limited distributions and more significant from
a zoogeographical point of view is quite high (Groups 6 and 7, 16 spp., 41%). Most of
these species are also distributed in the southern part of the Iberian Peninsula (Group
6) or are endemic to the Rif but showing affinities with Iberian species: Soteriscus
gibbosus, S. laouensis which are morphologically close to S. gaditanus;Porcellio
pseudornatus and P. riffensis, belonging to the Rif-Betic group of Porcellio;Eluma
praticola belonging to an Atlantic genus; and Armadillidium djebalensis and A.
pardoi, belonging to the serratum group of Armadillidium. All these examples seem
to show a common origin of the fauna of the Rif with that of the Betic cordillera,
which were in connection until the middle Miocene (Rosenbaum et al. 2002).
Acknowledgements
We wish to thank Dr C. Giordano (CeME-CNR, Florence) for her help with SEM analysis of
some species.
Funding
This work was funded by the MEDCORE project of the European Community INCO-DC 5th
Framework Programme Contract no. ICA3-2002-10003 (2003–2005).
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