Scorpions of Greece

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Scorpions of Greece and adjacent lands current
advances in systematics Victor Fet Marshall
University Huntington, West Virginia, USA
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GARY A. POLIS (1946 - 2000) a friend, a
teacher, a researcher
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• Ancient origin
• known in fossil record since Silurian (420 Ma)
  fossil forms may constitute 3 orders many
  aquatic taxa.
• modern Order Scorpiones from Carboniferous (250
  Ma), all terrestrial.
• Tertiary fossils (amber 60-30 Ma) already have
  extant families and genera.

Scorpions are a small (2000 species worldwide)
but diverse group of arthropods (14 families,
200 genera)
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Scorpion systematics is undergoing an exciting
time. Our understanding of traditional taxa at
all levels is changing rapidly and considerably
due to introduction of modern methods of analysis
(both morphological and molecular), discovery of
new important characters, intensive new
collection, and study of old museum material. A
picture that emerges of scorpions in Greece and
adjacent lands is that of a diverse and complex
group.
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...adjacent lands ? ?
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From Aristotle one jumps to Brullé (1832) to
first discover a few Greek scorpions with proper
Linnean names
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• Species of the genus Euscorpius Thorell, 1876 are
  ecologically diverse and they occupy a variety of
  habitats from xeric to mesic, from the
  Mediterranean shoreline to the high altitudes of
  the Alps, Balkans, and the Cilician Taurus.

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Already C. L. Koch (1837) ingeniously outlined
local species such as Euscorpius tergestinus, ?.
sicanus and E. naupliensis however later, the
lumping trend prevailed.
Carl Ludwig Koch (1778-1857)
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Alexei Birula (1864-1937)
Karl Kraepelin (1848-1915)
Franz Werner (1867-1939)
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Old museum labels and archives in French,
German, Italian, Latin, Bulgarian, Russian
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From Kraepelin (1899) to Kinzelbach (1975), the
Balkans and Anatolia do not enjoy very diverse
scorpion fauna a toxic buthid Mesobuthus
gibbosus, two relict monotypic iurid genera Iurus
(Greece and Turkey) and Calchas (Turkey), and a
few widely ranging Euscorpius (E. italicus, E.
carpathicus, E. germanus) were identified from
the region, known for its amazing diversity in
other animal groups. Are scorpions special? Why
are not they diverse? But they are.
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Hadi, 1929, 1930
Jovan Hadi (1884-1972)
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Lodovico di Caporiacco (1900-1951)
As indicated early by a record number of
subspecies in the Mediterranean Euscorpius
(Hadzi, 1930 Caporiacco, 1950), this genus
indeed contained a not-so-hidden diversity.
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Max Vachon (1908-1991)
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Vachon, 1981
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Vachon, 1971
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Important papers of 1970-1980s by Curcic,
Kritschner, Crucitti, and others on the Balkan
scorpions
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Scorpion fauna of Turkey 1950-1990s new data,
Vachon, Kritschner, Crucitti, Lacroix, and others
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Kinzelbach, 1975
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Fet Braunwalder, 2000 The fauna
and zoogeography of the scorpions in the Aegean
area are not well researched and based upon
specimens randomly collected and described by
various authors in the past 150 years. A first
revision ot the scorpions in this region was
provided by Kinzelbach in 1975. However, the
taxonomical validity of some species and most of
the subspecies, first of all those in the genus
Euscorpius (Euscorpiidae) is still unclear and
their geographic ranges remain rather uncertain.

Kaltsas et al., 2008
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INTENSIVE STUDY OF RICH, OLD COLLECTIONS
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VIENNA
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LONDON
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FRANKFURT
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SOFIA
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INTENSIVE NEW COLLECTING(nice places
preferred!)
Anamur
Mt Ossa
Thasos
bbbb
Mt. Ossa
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OLD METHODS, NEW CHARACTERS
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• Traditional diagnostic morphological
  characters
• in scorpions
• appendages (pedipalps claws femur, patella,
  chela)
• carapace (cephalothorax)
• metasoma (tail),
• pectinal organs, etc.

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Pectinal organs(paired modified mesosomal
appendages mechano- and chemosensory arrays)
number of teeth is a popular meristic characted
sine 18th century each tooth carries dozens to
hundreds sensilla
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NEW or well-forgotten old CHARACTERS

• Trichobothriotaxy numbers, patterns, variation
  pedipalps - detailed analysis
• Chaetotaxy legs, pedipalps, chelicerae,
  metasoma spines versus setae (Soleglad Fet
  2003, etc.)
• Other sensory organs eyes, pectines, mechano-
  and chemoreceptory setae, etc.
• Constellation array a new sensory organ! (Fet
  et al. 2006)
• Specialized non-sensory structures denticles
  fingers, chelicerae, carinae, leg spurs
• Serrula a neotenic structure? (Graham Fet 2006)

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Trichobothria (sensory bristles on pedipalps)
the most elaborate data set in scorpion
morphology
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Trichobothria number variation
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Trichobothria positional patterns variation
(patella)
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Constellation array (putative thermoreceptors)
(Calchas gruberi Fet et al., 2009) 15 sensilla
the highest known number in scorpions
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Tibial spur unique leg feature of the Anatolian
genus Calchas (Iuridae) and Old World Buthidae
and Pseudochactidae, as well as some fossil
scorpions
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Tarsal setation spination
Left, Calchas diagnostic combination of
socketed setae and non-socketed spines and
spinules on vetral surface of leg tarsus
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Tarsal setation spinationunique tufts of
Iurus (Iuridae) (Fet et al. 2004)
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• CHELICERA
• Two fingers, movable and fixed
• Dentition is diagnostic at high level
  (family/genus) (Vachon 1963)
• Setation is NOT studied

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• SERRULA
• on ventral surface of movable cheliceral finger
• discovered in scorpions only in 1944 (Mulaik
  Higgins (in Superstitionia) Vachon) (in
  Belisarius)
• a grooming device?
• Its tines (denticles) are very often broken
  could it be a juvenile device, retained but
  suboptimal in adults?

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Other novel structures discoveredDetailed SEM
survey reveals previously unknown microstructures
of scorpion cuticleleft, basitarsus of Calchas
a rosette of denticles, size of a an animal cell
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NEW METHODS

• A number of DNA phylogenies in genus Euscorpius
  has been published, based on mitochondrial rRNA
  markers, first introduced by Gantenbein et al.
  (1999)
• Based on same mt rRNA markers, the first DNA
  phylogenies are published for the region both for
  Mesobuthus and Iurus (Parmakelis et al., 2006a,
  2006b).

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16S rRNA Mitochondrial DNA molecules are
maternally inherited in most animals. Comparative
analyses of mitochondrial ribosomal RNA gene
sequences (12S and 16S) are now routinely used in
molecular systematics for resolving species-level
phylogenies of different animal groups.
Mitochondrial ribosomal RNA sequences contain
areas of high mutation rate known as
non-conserved regions as well as regions with low
mutation rates known as conserved regions (Avise,
1994). By using the mitochondrial ribosomal RNA
gene sequences for the 16S gene, we take
advantage of all these features.
16S rRNA molecule, secondary structure
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DNA sequencing

• Sequences are obtained in text format
• gtEgOB (AJ249553 GenBank submission number),
  Euscorpius germanus, Austria, Carinthia,
  Oberdrauenburg (Scherabon et al. 2000), 16S mt
  rDNA
• 1 tcgccagact tccttatttt attattgcat
  gaaataggta tcttaatcca acatcgaggt
• 61 cacaaacttt cttgatgata agaactcttg
  aagaaaatta tgctgttatc cctatagtaa
• 121 cttattcttt tttaaaaact ttttgattgt
  tcaagagtta ctcccacgtt attgaaataa
• 181 aattttattt atttactgcc ccagtagaat
  aatttttagt ttatgttgat attaaattga
• 241 gattataaag cttaataggg tcttcttgtc
  tttaaaatgg attttagctt ttttactaaa
• 301 atataaaatt tgaagtataa taataagaca
  tgattattta gttaaaccat tcattccagt
• 361 cctaaattac aagacta
• gtEalOL (AJ286754), Euscorpius alpha, Olmo al
  Brembo, Valle Brembana, Lombardia, Italy
  (Gantenbein et al. 2000), 16S mt rDNA
• 1 cttccttatt ttattattgc atgaaatagg
  tatcttaatc caacatcgag gtcacaaact
• 61 ttcttgatga taagaactct ttaagaaaat
  tatgctgtta tccctatagt aacttgttct
• 121 tttttaaaaa ctttttggtt tttcaagagt
  tattctcata ttattaaaat aaaattttat
• 181 ttatttactg ccccagtata ataatttgtg
  atttatattg atgttaaatt ggaattataa

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• Sequence alignment (ClustalX PAUP format)

NEXUS BEGIN DATA gene mitochondrial 16S rRNA,
amplified by PCR dimensions ntax8 eight
species nchar385 385 DNA base pairs format
missing? symbols"ABCDEFGHIKLMNPQRSTUVWXYZ" inter
leave datatypeDNA gap - matrix EcRO1
TCGAACAGA-CTTCCTTAATTTTATTATTGCATGAAAAAGGTCTCTTAAT
EcRO2 TCGAACAGNACTTCCTTAATTTTATTNTTGCATGAAA
AAGGTCTCTTAAT EtMD2 TCGAACAGA-CTTCCTTAATTTTA
TTATTGCATGAAAAAGGTTTCTTAAT EtMA1
TGGCACAGA-CTTCCTTAATTTTATTATTGCATGAAAAAGGTTTCTTAAT
EcKA2 TCGAACAGA-CTTCCTTAATTTTGTTATTGCACAAAA
AAGGTATCTTAAT EhDU2 TCGAACAGA-CTTCCTTAATTTTA
TTATTGCATGAAAAAGGTATCTTAAT EbBA1
TCTAACAGA-CTTCCTTAATTT-ATTATTGCATGAAAAAGGTA-CTTAAT
EfLA TCGAACAGA--CTCCCTTATTTTATTATTGCATGAAA
TAGGGCTCTTAAT EcRO1 CCAACATCGAGGTCACAAACTTT
CTTGATGATAAGAACTCTTTAAGAAAA EcRO2
CCAACATCGAGGTCACAAACTTTCTTGATGATAAGAACTCTTTAAGAAAA
EtMD2 CCAACATCGAGGTCACAAACTTTCTTGATGATAAGAA
CTCTTTAAGAAAA EtMA1 CCAACATCGAGGTCACAAACTTTC
TTNACGATGAGAACTCTTTAAGAAAA EcKA2
CCAACATCGAGGTCACAAACTTTCTTGATGATAAGAACTCTTTAAGAAAA
EhDU2 CCAACATCGAGGTCACAAACTTTCTTGATGATAAGAA
CTCTTTAAGAAAA EbBA1 CCAACATCGAGGTCACAAACTTTC
TTGATGATAAGAACTCTTTAAGAAAA EfLA
CCAACATCGAGGTCACAAACTTTCTTGATGATAAGGACTCTTAAAGAAAA
EcRO1 TAATGCTGTTATCCCTATAGTAACTTATTCCTTTTT
AAAAATTTTTTGAT EcRO2 TAATGCTGTTATCCCTATAGTAA
CTTATTCCTTTTTAAAAATTTTTTGAT EtMD2
TAATGCTGTTATCCCTATAGTAACTTATTCCTTTTTAAAAATTTTTTGGT
EtMA1 TAATGCTGTTATCCCTATAGTAACTTATTCTTTTTTA
AAAATTTTTTGAT EcKA2 TTATGCTGTTATCCCTATAGTAAC
TTATTTCTTTTTAAAAATTTTTTGAT EhDU2
TAATGCTGTTATCCCTATAGTAACTTATTCATTTTTAAAAATTTTTTGAT
EbBA1 TTATGCTGTTATCCCTATAGTAACTTATTCCTTTTTA
AAAACTTTTTGAT EfLA TTATGCTGTTATCCCTA-AGTAAC
TTGTTCCTTCTTAAGAAATTTTTGAT end
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BUTHIDAE
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Mesobuthus gibbosus (Brullé, 1832) (Buthidae)
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• Gantenbein Largiadèr (2002)
• Allozyme data suggest that the Rhodes population
  of Mesobuthus gibbosus is a hybrid population of
  recent origin. Namely, it is a mixture between an
  autochthonous population and an artificially
  introduced population probably from the Greek
  mainland. All samples were mainly composed of F1
  hybrid genotypes and genotypes either fixed for
  autochthonous or introduced alleles. MtDNA
  analysis, in contrast, revealed only one group of
  closely related haplotypes that are unique for
  the Rhodes populations, thus suggesting
  asymmetric introgression of the two marker
  classes.

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• Attention to relatively uniform Mesobuthus yields
  an endemic species on Cyprus (Gantenbein et al.
  2000) the Anatolian fauna of Mesobuthus is
  addressed in detail, including recently restored
  M. nigrocinctus (Karatas Karatas 2001, 2003,
  Karatas 2007 etc.). Other buthids (Hottentotta,
  Compsobuthus) are recorded for SE Anatolia.

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EUSCORPIIDAE
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For almost a century, only four species were
listed in Euscorpius E. carpathicus, E.
germanus, E. flavicaudis and E. italicus
(Kraepelin, 1899 Birula, 1900, 1917
Caporiacco, 1950). Bonacina (1980)
reestablished E. mingrelicus (Kessler,
1874). Further revisional work (1999-2007)
resulted in 17 formally recognized species E.
alpha Caporiacco, 1950 E. balearicus Caporiacco,
1950 E. beroni Fet, 2000 E. carpathicus
(Linnaeus, 1767) E. concinnus (C. L. Koch,
1837) E. flavicaudis (DeGeer, 1778) E. gamma
Caporiacco, 1950 E. germanus (C. L. Koch,
1837) E. hadzii Caporiacco, 1950 E. italicus
(Herbst, 1800) E. koschewnikowi Birula, 1900 E.
mingrelicus (Kessler, 1874) E. naupliensis (C.L.
Koch, 1837) E. oglasae Caporiacco, 1950 E.
sicanus (C.L. Koch, 1837) E. tauricus (C.L.
Koch, 1837) E. tergestinus (C.L. Koch, 1837)
(Fet 2000, 2003 Gantenbein et al. 2000, 2001,
2002 Scherabon et al. 2000 Fet Soleglad 2002
Fet et al. 2003a, 2003b, 2006 Vignoli et al.
2006 Salomone Vignoli 2007).
Ongoing splitting in Euscorpius a trend
started by Alberto Bonacina (1980) who separated
E. mingrelicus from E. germanus.
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Euscorpius systematics

• We outlined related species E. hadzii and E.
  sicanus as separate from the main carpathicus
  complex (Fet Soleglad 2002 Fet et al. 2003),
  which also likely includes several species, some
  undescribed.
• We rediscovered the forgotten Peloponnese endemic
  E. naupliensis, a sister group to a possibly
  glacial bottleneck survivor E. italicus
  (Gantenbein et al. 2003 Fet et al. 2006), both
  in subgenus Polytrichobothrius.
• At least five species of Euscorpius inhabit
  Greece alone, often sympatrically the
  collaborative international work is under way to
  obtain a detailed molecular and morphological
  phylogeny of these species and their populations,
  including many island groups.

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• E. carpathicus does not live here any more we
  restrict it to the type populations in Romania.
• At least three Euscorpius s. str. species live in
  Bulgaria, where northern and southern
  carpathicus complex members are not closely
  related (Fet Soleglad 2007)
• We project more than one Euscorpius species
  inhabiting Greek islands (e.g. Thasos) and
  isolated peninsulas (Chalkidiki, Peloponnese).
• Human introduction likely plays role for many
  Euscorpius (Fet et al. 2006) and Mesobuthus
  (Gantenbein Largiadèr 2003).
• The Western Balkans and the Anatolian Peninsula
  have a relict and diverse populations of subgenus
  Alpiscorpius, absent from the mainland Greece
  (exept maybe northern Epiros) and Aegean islands
  (except Samos).

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Four Euscorpius species from carpathicus
complex in southern Europe (Fet Soleglad,
2002) cooperation with researchers from France,
Italy, Greece, Bulgaria, Slovenia, Austria,
Spain, Switzerland
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Euscorpius sicanus in Italy, Greece Malta (Fet
et al. 2003) in cooperation with University of
Siena, Italy University of Crete, Greece
University of Malta, Malta.
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Among the denizens of Mt. Olympus, Greece TWO
sympatric Euscorpius species (first discovered
by Ragnar Kinzelbach, 1975)
S???p???
(what do you know! ?? ??da ?t? ??d?? ??da)
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DNA-based phylogenetic tree Euscorpius sicanus
(possibly a species complex)
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At least four species of scorpions are recorded
for Bulgaria (up to 1850 m a.s.l.). The Balkan
species Euscorpius hadzii inhabits the southwest
of Bulgaria. Other Bulgarian populations of
Euscorpius, widely ranging from north to south
belong to Euscorpius carpathicus complex. A
clearly separated cluster of Rhodope populations
could represent a different taxon from the Stara
Planina ones and those from the southwest of
Bulgaria.
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A new (or rather well-forgotten old), relict
species of Euscorpius (subgenus
Polytrichobothrius) from Peloponnesos
(Gantenbein et al., 2002)
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Trichobothria positional patterns variation
(fixed finger of chela)
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Vachon, 1981
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DNA-based phylogenetic tree Euscorpius
italicus and E. naupliensis
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Little or no genetic divergence is detected using
mitochondrial 16S rDNA sequence comparisons
across the entire geographic range of the
scorpion Euscorpius italicus (Herbst 1800) from
Switzerland, Italy, Slovenia, Greece and Turkey.
This is consistent with known absence of patterns
of allozymes and morphological variation.
Euscorpius italicus is found almost exclusively
in human habitations. Its sister species, E.
naupliensis, exhibits much higher genetic
diversity within southern Greece. We suggest that
the natural populations of the thermophilic E.
italicus underwent a bottleneck during the
glaciations, and that its modern range could be a
result of dispersal with humans.
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IURIDAE
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• Two monotypic, relict genera of Iuridae (Greece
  and Turkey), often collected but ignored for
  decades, yield new insights
• unexpected and diverse neobothriotaxy of Iurus is
  discovered (Soleglad et al., 2009)
• three allopatric species of Calchas are
  discovered (Fet et al., 2009)
• more surprises are in store for these fascinating
  iurid relicts (Soleglad, Fet Kovarík, in
  progress).

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• Neobothriotaxy in Iurus (discovered for the
  first time Soleglad et al. 2009)
• Based on the examination of 101 specimens of
  Iurus, from mainland Greece, Crete, Rhodes, and
  Turkey, no less than nine distinct types of
  neobothriotaxy have been detected
• CHELA
• basal aspect of the internal surface(1)
  suprabasal petite accessory trichobothrium (t)
  (one instance) (2) basal petite accessory t (one
  instance)
• (3) distal accessory t (11 instances)
• external surface(4) one or two accessory t are
  present in the Et series (47 instances) (5) one
  petite accessory t in the Est series (four
  instances)
• ventral surface
• (6) a petite t in the V series (one instance)

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Neobothriotaxy in Iurus PATELLA three types
of neobothriotaxy on the patella external
surface (7) one to two accessory t on the
dorsal half of the surface (seven instances)
(8) one accessory t on the ventral surface in
the em series (one instance) (9) one petite
accessory t on the ventral surface in the et
series (four instances).

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Genus Calchas (Iuridae) - three allopatric
species!
Artvin
Schmetterlingenbuche
Schmetterlingenbuche
Schmetterlingenbuche
Schmetterlingenbuche
Schmetterlingenbuche
Mardin
Anamur
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PEDIPALP CHELA Diagnostic combinations of
trichobothria on the fingertip constellation
array of sensilla (Calchas birulai Fet et al.,
2009)
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Prepectinal plate in Calchas sexual
dimorphism, function unknown
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Unique and virtually unknown structure!
Sternopectinal area in Calchas the genital
operculum, prepectinal plate (in female), and
genital papillae (in male). In male, the large
long subtriagular genital operculum with
separated sclerites exposing significant genital
papillae female has a narrow operculum with
fused sclerites. Top left, C. nordmanni, female,
Tortum, Turkey.right. C. birulai, sp. nov.,
femaleC. nordmanni, Mardin, Turkey. Bottom left.
C. gruberi, sp. nov., female, 12 km S. Akseki,
Turkey right, male, Turkey. (Fet et al. 2009)
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BACK TO ARISTOTLE?
And now to something entirely different.
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ARISTOTLE (384-322 B.C.)History of Animals,
iv.7 ."..Some insects have the sting inside, as
the bee and the wasp, others outside, as the
scorpion and, by the way, this is the only
insect furnished with a long tail. And, further,
the scorpion is furnished with claws..."
History of Animals, viii.29 locality is an
important element in regard to the bite of an
animal. Thus, in Pharos and other places, the
bite of the scorpion is not dangerous elsewhere
in Caria, for instances where scorpions are
venomous as well as plentiful and of large size,
the sting is fatal to man or beast

• Aristotle tutoring Alexander,
• modified from J L G Ferris 1895

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Caria, part of classical Greece in Southwestern
Anatolia (Mugla Province, modern Turkey), is
inhabited by several scorpion species, among them
a relatively toxic Mesobuthus gibbosus

• But where is Pharos?
• The most famous ancient Pharos was a small island
  off coast Alexandria, Egypt, a natural harbor
  mentioned in the Odyssey.
• When Alexander marched there in the autumn of 332
  BC, the 1500-m mole connecting Pharos to a
  fishing village, Rhacotis, was built, and the
  great city of Alexandria begun.
• the famous, at least 100 m tall, Lighthouse of
  Alexandria, the last of the Seven Wonders of the
  World, was built on Pharos between 285 and 247
  BC
• the site of Septuagint Project, the translation
  of Torah (Pentateuch) into Koine Greek,
  commissioned for the Library of Alexandria by
  Ptolemy II Philadelphus (285246 BC)

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But why should the scorpions of Pharos be
non-toxic? Scorpions of Egypt (mainly Buthidae)
always were, and still are, widely and justly
known for their toxicity since the child Horus
was stung in the heel We see no reason why the
small island of Pharos would be singled out by
Aristotle in 340s BC for having scorpions that
are not dangerous.
Our attention turns now to another Pharos the
modern Hvar Island, a popular tourist resort off
the Croatian coast in the Adriatic Sea. This
Pharos was a Greek colony, founded by Ionian
settlers from Paros in 385 BC
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Euscorpius tergestinus. Photo by Ryszard
Wiejski-Wolschendorf, made on Hvar Island,
Croatia (4309.881 N 1626.667 E), October 3,
2006.

• We think it quite plausible that Aristotle had in
  mind non-dangerous Euscorpius spp. from the
  Illyrian Pharos (as well as other places!) as
  compared with toxic Mesobuthus spp. from
  Anatolia (and most of modern Greece).

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• The first observation on scorpion biogeography
  by Aristotle
• Victor Fet,
• Hisham El-Hennawy,
• Matt E. Braunwalder
• John L. Cloudsley-Thompson
• Boletin S.E.A., 2009 44

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CONCLUSIONS
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• A picture that emerges of scorpions in Greece
  and adjacent lands is that of a diverse and
  complex group.
• They are a promising group for speciation and
  other evolutionary studies, employing character
  sets from new morphology (e.g. constellation
  sensory array Fet et al. 2006, in progress) to
  karyotypes (first steps now attempted by F.
  Stahlavsky in Prague) to detailed, multiple DNA
  markers, both mitochondrial and nuclear etc.
  (allozymes, microsatellites).
• Scorpion biology and ecology in the region,
  studied mainly by the Italian and Greek authors
  since 1980s, still remains in its infancy.

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ACKNOWLEDGMENTS I thank all colleagues,
co-authors and friends from many countries for
their enthusiastic collaboration, for sharing
specimens, literature, time and expertise over
all the years of my scorpion studies... Their
list includes (but is not limited to) Luis
Acosta, Luis de Armas, Mark Barker, Janet
Beccaloni, Günther Bechly, Petar Beron,
Alexander Blinov, Alberto Bonacina, Nikos
Botsaris, Philip Brownell, Matt Braunwalder,
Michael Brewer, H. D. Cameron, Terry Christenson,
John Cloudsley-Thompson, Jonathan Coddington,
James Cokendolpher, Marco Colombo, Pat Craig,
Pierangelo Crucitti, Hieronymus Dastych, Christo
Deltshev, Dobrin Dobrev, Jason Dunlop, Gérard
Dupré, Marshall Hedin, Hisham El-Hennawy, Roger
Farley, Günther and Gerta Fleissner, Douglas
Gaffin, Benjamin Gantenbein, Arnold Gegechkori,
Matt Graham, Manfred Grasshoff, Charles Griswold,
Joshua Greenwood, Alexander Gromov, Jürgen
Gruber, Paul Hillyard, Christoph Hörweg, Dietmar
Huber, Peter Jäger, Dimitris Kaltsas, Ahmet
Karatas, Aysegul Karatas, Ragnar Kinzelbach,
Christian Komposch, Frantiek Kovarík, Jirí Král,
Alex and Elena Kreuzberg, Viktor Krivochatsky,
Christian Kropf, Oleg Kryzhanovsky, Kadir Bogac
Kunt, Matjaz Kuntner, Adolf Kurka, Jean-Bernard
Lacroix, Scott Larcher, Laura Leibensperger,
Herbert Levi, Gershom Levy, Wilson Lourenço,
Graeme Lowe, Yuri Marusik, Kirill Mikhailov,
Moysis Mylonas, Shakhrokh Navidpour, David Neff,
Vladimir Ovtsharenko, Ivan Pandourski, Aris
Parmakelis,Rusty Parrett, Vera Pkhakadze, Gary
Polis, Guillem Pons, Alexi Popov
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Valentin Popa, Vlado Sakalian, Nicola Salomone,
Jorge Santiago-Blay, Patrick Schembri, Bernhard
Scherabon, Adolph Scholl, Paul Selden, Petra
Sierwald, David Sissom, Boris Sket, Michael
Soleglad, Verena Stagl, Frantiek Stahlavsky,
Iasmi Stathi, Alex Stewart, Roland Stockmann,
Rolando Teruel, Konrad Thaler, Ian Towler,
Ryszard Wiejski-Wolschendorf, Sarah Whitman,
Stanley Williams, Alex Winkler, Max Vachon,
Marco Valle, Milen Vasilev, Nikolay Vasilyev,
Mark Volkovich, Valerio Vignoli, Ersen Yagmur,
T. Yamashita, Eric Ythier, and Jirí Zidek. My
special thanks to my family Galina, Elizabeth
and Simon, for their enthusiastic help in
studying scorpions of Greece and adjacent
countries!
Ivan Pandourski, Simon Fet, and Christo Deltshev
catching Euscorpius sp. in NW Bulgaria, 2005
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Great thanks to 25th ECA organizers!