Two New Families and a New Species of Myxosporeans (Myxozoa, Myxosporea) from the Mediterranean and Black Sea Fishes
Descriptions of two new families – Polysporoplasmidae fam. n. (Syn. Sphaerosporidae Davis, 1917 in Sitja-Bobadilla, Alvarez-Pellitero, 1995) and Gadimyxidae fam. n. (syn. Parvicapsulidae Schulman, 1953 in Køie et al., 2007), and one new species of myxosporeans – Gadimyxa ovale sp. n. – parasites of...
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Інститут зоології ім. І.І. Шмальгаузена НАН України
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irk-123456789-658422014-07-04T03:01:46Z Two New Families and a New Species of Myxosporeans (Myxozoa, Myxosporea) from the Mediterranean and Black Sea Fishes Yurakhno, V.M. Краткие сообщения Descriptions of two new families – Polysporoplasmidae fam. n. (Syn. Sphaerosporidae Davis, 1917 in Sitja-Bobadilla, Alvarez-Pellitero, 1995) and Gadimyxidae fam. n. (syn. Parvicapsulidae Schulman, 1953 in Køie et al., 2007), and one new species of myxosporeans – Gadimyxa ovale sp. n. – parasites of fishes of the Mediterranean and the Black Seas are presented. Species of the genus Gadimyxa is found for the first time in the Black Sea. Описаны два новых семейства – Polysporoplasmidae fam. n. (син. Sphaerosporidae Davis, 1917 по Sitja-Bobadilla, Alvarez-Pellitero, 1995) и Gadimyxidae fam. n. (син. Parvicapsulidae Schulman, 1953 по Kшie et al., 2007), а также нового вида миксоспоридий – Gadimyxa ovale sp. n. – паразитов рыб Средиземного и Чёрного морей. Вид из рода Gadimyxa впервые заре-гистрирован в Чёрном море. 2011 Article Two New Families and a New Species of Myxosporeans (Myxozoa, Myxosporea) from the Mediterranean and Black Sea Fishes / V.M. Yurakhno // Вестник зоологии. — 2011. — Т. 45, № 6. — С. 555–559. — Бібліогр.: 16 назв. — англ. 0084-5604 http://dspace.nbuv.gov.ua/handle/123456789/65842 593.194:597.2/.5(262+262.5) en Вестник зоологии Інститут зоології ім. І.І. Шмальгаузена НАН України |
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Краткие сообщения Краткие сообщения Yurakhno, V.M. Two New Families and a New Species of Myxosporeans (Myxozoa, Myxosporea) from the Mediterranean and Black Sea Fishes Вестник зоологии |
| description |
Descriptions of two new families – Polysporoplasmidae fam. n. (Syn. Sphaerosporidae Davis, 1917 in Sitja-Bobadilla, Alvarez-Pellitero, 1995) and Gadimyxidae fam. n. (syn. Parvicapsulidae Schulman, 1953 in Køie et al., 2007), and one new species of myxosporeans – Gadimyxa ovale sp. n. – parasites of fishes of the Mediterranean and the Black Seas are presented. Species of the genus Gadimyxa is found for the first time in the Black Sea. |
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Yurakhno, V.M. |
| author_facet |
Yurakhno, V.M. |
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Yurakhno, V.M. |
| title |
Two New Families and a New Species of Myxosporeans (Myxozoa, Myxosporea) from the Mediterranean and Black Sea Fishes |
| title_short |
Two New Families and a New Species of Myxosporeans (Myxozoa, Myxosporea) from the Mediterranean and Black Sea Fishes |
| title_full |
Two New Families and a New Species of Myxosporeans (Myxozoa, Myxosporea) from the Mediterranean and Black Sea Fishes |
| title_fullStr |
Two New Families and a New Species of Myxosporeans (Myxozoa, Myxosporea) from the Mediterranean and Black Sea Fishes |
| title_full_unstemmed |
Two New Families and a New Species of Myxosporeans (Myxozoa, Myxosporea) from the Mediterranean and Black Sea Fishes |
| title_sort |
two new families and a new species of myxosporeans (myxozoa, myxosporea) from the mediterranean and black sea fishes |
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Інститут зоології ім. І.І. Шмальгаузена НАН України |
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2011 |
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http://dspace.nbuv.gov.ua/handle/123456789/65842 |
| citation_txt |
Two New Families and a New Species of Myxosporeans (Myxozoa, Myxosporea) from the Mediterranean and Black Sea Fishes / V.M. Yurakhno // Вестник зоологии. — 2011. — Т. 45, № 6. — С. 555–559. — Бібліогр.: 16 назв. — англ. |
| series |
Вестник зоологии |
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AT yurakhnovm twonewfamiliesandanewspeciesofmyxosporeansmyxozoamyxosporeafromthemediterraneanandblackseafishes |
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2025-07-05T16:13:24Z |
| last_indexed |
2025-07-05T16:13:24Z |
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1836824138913677312 |
| fulltext |
UDC 593.194:597.2/.5(262+262.5)
TWO NEW FAMILIES AND A NEW SPECIES
OF MYXOSPOREANS (MYXOZOA, MYXOSPOREA)
OF THE MEDITERRANEAN AND BLACK SEA FISHES
V. M. Yurakhno
Institute of Biology of the Southern Seas of NAS of Ukraine,
Nakhimov Av., 2, Sevastopol, 99011 Ukraine
E-mail: viola_taurica@mail.ru
Received 18 August 2010
Accepted 10 November 2011
Two New Families and a New Species of Myxosporeans (Myxozoa, Myxosporea) from the Mediter rane -
an and Black Sea Fishes. Yurakhno V. M. – Descriptions of two new families – Polysporoplasmidae
fam. n. (Syn. Sphaerosporidae Davis, 1917 in Sitja-Bobadilla, Alvarez-Pellitero, 1995) and
Gadimyxidae fam. n. (syn. Parvicapsulidae Schulman, 1953 in Køie et al., 2007), and one new species
of myxosporeans – Gadimyxa ovale sp. n. – parasites of fishes of the Mediterranean and the Black
Seas are presented. Species of the genus Gadimyxa is found for the first time in the Black Sea.
Ke y wo r d s: Myxosporea, Polysporoplasmidae fam. n., Gadimyxidae fam. n., Gadimyxa ovale sp. n.,
fishes, Mediterranean, Black Sea.
Äâà íîâûõ ñåìåéñòâà è íîâûé âèä ìèêñîñïîðèäèé (Myxozoa, Myxosporea) èç ðûá Ñðåäèçåìíîãî è
×åðíîãî ìîðåé. Þðàõíî Â. Ì. – Îïèñàíû äâà íîâûõ ñåìåéñòâà – Polysporoplasmidae fam. n.
(ñèí. Sphaerosporidae Davis, 1917 ïî Sitja-Bobadilla, Alvarez-Pellitero, 1995) è Gadimyxidae fam. n.
(ñèí. Parvicapsulidae Schulman, 1953 ïî Køie et al., 2007), à òàêæå íîâîãî âèäà ìèêñîñïîðèäèé –
Gadimyxa ovale sp. n. – ïàðàçèòîâ ðûá Ñðåäèçåìíîãî è ׸ðíîãî ìîðåé. Âèä èç ðîäà Gadimyxa
âïåðâûå çàðå-ãèñòðèðîâàí â ׸ðíîì ìîðå.
Êëþ÷åâûå ñ ëîâ à: ìèêñîñïîðèäèè, Polysporoplasmidae fam. n., Gadi-myxidae fam. n.,
Gadimyxa ovale sp. n., ðûáû, Ñðåäèçåìíîå ìîðå, ׸ðíîå ìîðå.
Our investigations of myxosporean fauna of mullets (family Mugilidae) in the Black and Mediterranean
seas showed that attribution of Polysporoplasma mugilis Sitja-Bobadilla et Alvarez-Pellitero, 1995 (Sitja-
Bobadilla, Alvarez-Pellitero, 1995) to the family Sphaerosporidae Davis, 1917 was a mistake. The genus
Polysporoplasma Sitja-Bobadilla et Alvarez-Pellitero, 1995 presented now by two species forms a new family
Polysporoplasmidae fam. n.
Our study of myxosporeans from Mediterranean Sea burbot Gaidropsarus mediterraneus in the Black
Sea revealed a new species – Gadimyxa ovale sp. n. Representative of this genus was found for the first time
in the Black Sea.
Material and methods
Mullets were purchased in June 2004 and May 2005 at local fish markets from two localities in the
Spanish Mediterranean, namely, from the Ebro River delta and Santa Pola Bay and in June 2004 from
Yuzhnaya Bay of Sevastopol (the Black Sea, Ukraine; caught with a fishing rod). Myxosporeans of mullets
were taken from alive or freshly frozen fish. Mediterranean Sea burbot Gaidropsarus mediterraneus (Linnaeus)
was received from fishermen of the fishing brigade of the Institute of Biology of the Southern Seas of NASU,
from Sevastopol bays of the Black Sea during October, November, and December of 2002—2004.
Myxosporeans of the Mediterranean Sea burbot were taken only from alive fish. A total of 246 specimens
belonging to three species of Mediterranean mullets were examined: thin-lip mullet Liza ramada (Risso,
1826) (207 specimens), golden grey mullet Liza aurata (Risso, 1810) (36 specimens) and thick-lip grey mul-
let Chelon labrosus (Risso, 1827) (3 specimens). Totally 9 specimens of L. aurata and 40 specimens of
G. mediterraneus were examined from the Black Sea. All tissues and organs of fish were examined for myx-
osporean infections. The studies of spores and plasmodia were conducted either directly on alive material or
Vestnik zoologii, 45(6): 555—559, 2011 Êðàòêèå ñîîáùåíèÿ
on gelatine-glycerine slides using the phase-contrast device (Donets, Shulman, 1973). Spores were pho-
tographed and measured on digital images. Descriptions of the spores were based on the guidelines of Lom
and Dykova (1992) and Schulman et al. (1997) taking into account modern classification of Myxozoan par-
asites (Lom, Dykova, 2006).
POLYSPOROPLASMIDAE Yurakhno, fam. n.
Type genus: Polysporoplasma Sitja-Bobadilla et Alvarez-Pellitero, 1995
Myxosporeans of the order Bivalvulida Schulman, 1959, the suborder Variisporina
Lom et Noble, 1984 with very big spores and polar capsules of spherical or subspheri-
cal shape. There are two polar capsules, equal in size, located on the anterior pole of
the spore on the sides of sutural line. Sutural line is straight. The spores are character-
ized by very great thickness of valves, thickest at the anterior and posterior poles of the
spore. There are two distinct bulges or horns (one on each valve) at the posterior pole
of the spore in front view. Presence of several (from 4 to 12) uninucleate sporoplasms
in a spore is the most prominent peculiarity of the family. Vegetative forms are two-
spore and polyspore plasmodia of round and oval form. One genus, Polysporoplasma,
and two species, P. sparis Sitja-Bobadilla, Alvarez-Pellitero, 1995 and P. mugilis Sitja-
Bobadilla, Alvarez-Pellitero, 1995 are known. Parasites on the sea fish kidneys.
Polysporoplasma mugilis Sitja-Bobadilla et Alvarez-Pellitero, 1995 (fig. 1, 2)
Ho s t s. In the Mediterranean Sea: thinlip grey mullet Liza ramada (Risso), gold-
en grey mullet L. aurata (Risso), thicklip grey mullet Chelon labrosus Risso; in the
Black Sea: L. aurata (Risso) (fam. Mugilidae).
S i t e: kidneys.
Lo c a l i t y a nd t ime o f r e v e a l i n g. Mediterranean Sea: Spain, river Ebro
Delta, June 2004: in 2 of 37 (prevalence – 5 %) of studied L. ramada with general
length (L) 41—42.5 ñm; in 1 of 36 (3 %) of studied L. aurata with length 32.5 ñm; May
2005: in 2 of 109 (2 %) of studied L. ramada with length 39.2—39.8 ñm, and in 1 of 3
C. labrosus; Santa Pola – in 3 of 61 (prevalence – 8 %) L. ramada with length 34—35
ñm; Black Sea: Sevastopol, Yuzhnaya bay, June 2004: in 1 of 9 of studied L. aurata
with length 21.7 ñm.
The size of P. mugilis spores from fresh smears of L. ramada kidney appeared to
be a bit differing from that in the first description. The length of spores was 18.0*—22.0,
width of spores 16.0—19.5, thickness of spores 16.5, diameter of polar capsules 5.5—6.0.
Polar thread formed 4 spires.
556 V. M. Yurakhno
Fig. 1—2: 1 – Plasmodium of Polysporoplasma mugilis with spores. 2 – Spores of Polysporoplasma mugilis.
Ðèñ. 1—2: 1 – Ïëàçìîäèé Polysporoplasma mugilis ñî ñïîðàìè. 2 – Ñïîðû Polysporoplasma mugilis.
* All measurements in micrometers (μm) unless otherwise indicated.
Liza ramada and C. labrosus are found to be new hosts for P. mugilis in the
Mediterranean Sea. In the Black sea, P. mugilis is a new species for the local fauna
(Ovcharenko, Jurakhno, 2006; Ovcharenko et al., 2007; Yurakhno, 2008; Yurakhno,
Ovcharenko, 2008).
GADIMYXIDAE Yurakhno, fam. n.
Type genus: Gadimyxa Køie, Karlsbakk et Nylund, 2007
Myxosporeans of the order Bivalvilida, the suborder Variisporina, with spores of
semicircular or oval, transversally elongated shape, sometimes (sub) spherical. Anterior
spore pole is often flattened. Valves are smooth, without protrusions, non-engraved.
The sutural line is thin, strait or weakly curved. It is characteristic, that when seen from
the front the line goes over the anterior and posterior spore poles horizontally, but not
vertically, as in the representatives of Leptotheca Thélohan, 1895 (Thélohan, 1895) and
Ceratomyxa Thélohan, 1892 (Thélohan, 1892), in which the spore shape is similar to
that in Gadimyxa (Køie et al, 2007). Presence of the spindle-shaped bulge surrounding
polar capsules and located along the sutural line is another peculiarity of the family rep-
resentatives. This bulge is not always seen on the spores from the fresh smears. Two
polar capsules of rounded shape and not of big size are located at the anterior spore
pole and open near the sutural line. There is one sporoplasm, it has two nuclei and dis-
tributed all over the spore plane under the polar capsules. Vegetative forms are round-
ed, oval or elongated two-spore, coarse-grained plasmodia, sometimes with one end
sharpened, with well seen pseudopodia. Parasites of the sea fish urinary bladder. The
family contains one genus Gadimyxa and four species – G. atlantica Køie, Karlsbakk
et Nylund, 2007, G. sphaerica Køie, Karlsbakk et Nylund, 2007, G. arctica Køie,
Karlsbakk et Nylund, 2007 (Køie et al., 2007) and G. ovale sp. n. Diagnosis of the
genus Gadimyxa corresponds to the diagnosis of the family.
Gadimyxa ovale Yurakhno, sp. n. (figs 3, 4)
Ho s t: Mediterranean Sea burbot Gaidropsarus mediterraneus (Linnaeus) (fam.
Gadidae).
S i t e: urinary bladder.
Lo c a l i t y a nd t ime o f r e v e a l i n g: Black sea, Sevastopol (Karantinnaya
Bay – November 2002: in 2 of 6 studied fishes; October 2004: in 3 of 11, November
2004: in 2 of 5, December 2004: in 1 of 2. Martynova Bay – December 2002: in 3 of
3. Alexandrovskaya Bay – October 2003: in 4 of 13 studied fishes).
Syntypes: slides N 1, 9, 30, 32, BAS 6, BAS 5.
Syntypes of Gadimyxa ovale sp. n. are deposited in the collection of the
Department of Ecological Parasitology of IBSS, NAS of Ukraine (Sevastopol).
Vegetative forms: rounded, oval and wide—oval bisporous plasmodia, sometimes
with one sharpened end, with coarse-grained endoplasma, with many well seen pseu-
dopodia, 0.5— 2.0 long. Size of plasmodia 10.5 × 12.5, 9.7 × 17.0, 27.0 × 34.0.
Spores are elongated, oval, smooth. Polar capsules are rounded, small, situated at
front spore pole and opening near the sutural line. Sutural line is thin, slightly curved.
The line observed from the front is situated horizontally, over the anterior and poste-
rior spore poles. Sporoplasm is distributed all over the spore plane under the polar cap-
sules. Measurements of the spores are given in table 1.
The general length (L) of infected fish was 20.0—27.5 ñm. Single specimens were
observed in the smears.
D i f f e r e n t i a l d i a g no s i s. Gadimyxa ovale sp. n. is similar to the three other
species of the genus Gadimyxa (Køie et al., 2007) from Gadiidae fish – Atlantic cod
Gadus morhua L. (Gadidae) from the Øresund, Denmark, western Norway (G. atlanti-
557Two New Families and a New Species of Myxosporeans (Myxozoa, Myxosporea)...
ca and G. sphaerica) and Arctogadus glacialis (Peters) from Greenlandic seas (G. arcti-
ca). G. ovale sp. n. vegetative stages have rounded or oval and wide—oval shape, where-
as other species of this genus have elongated or spherical shape. The spores of G. ovale
sp. n. differ by their shape and size from those of G. atlantica, G. sphaerica and G. arc-
tica. Semicircular shape of the spores occurs rarely, spherical shape was not found.
Length of the spores of G. ovale sp. n. is larger than in G. atlantica and smaller than in
G. sphaerica and G. arctica. Spores of Gadimyxa ovale sp. n. are wider and with thick-
er valves than in G. sphae rica, with considerably smaller polar capsule diameter than in
other Gadimyxa species.
Discussion
Very large size of Polysporoplasma mugilis spores and polar capsules, very great
thickness of its valves, thickest at the anterior and posterior poles of the spore, two well
seen bulges or horns at the posterior pole of the spore in front view, several (from 4 to
12) uninucleate sporoplasms in the spore, very big size and unusual form of vegetative
stages are the evidence of this genus uniqueness, in contrast to other representatives of
Sphaerosporidae; this set of features supports the existence of a separate myxosporean
family Polysporoplasmidae.
558 V. M. Yurakhno
Fig. 3—4. 3 – Plasmodia and spores of Gadimyxa ovale sp. n.: a, b, c – plasmodia with different forms and
sizes; d, e, f – spores. 4 – Spore of Gadimyxa ovale sp. n.
Ðèñ. 3—4. 3 – Ïëàçìîäèè è ñïîðû Gadimyxa ovale sp. n.: a, b, c – ïëàçìîäèè ðàçíûõ ôîðì è ðàçìå-
ðîâ; d, e, f – ñïîðû. 4 – Ñïîðà Gadimyxa ovale sp. n.
Ta b l e 1. The size of Gadimyxa ovale sp. n. spores on the fresh and fixed preparations
Ò à á ëèö à 1. Ðàçìåðû ñïîð Gadimyxa ovale sp. n. íà ñâåæèõ è ôèêñèðîâàííûõ ìàçêàõ
Size of spores and polar capsules Fresh smears Gelatine-glycerine preparations
Length of spores 5.3—6.5 5.4—6.6
Width of spores 8.7—9.2 6.5—8.4
Thickness of spores 6.0
Diameter of polar capsules 1.2 1.2—1.3 (1.5)
Differences in Gadimyxa ovale sp. n. size and the shape of the spores in compari-
son with other species of this genus prove the validity of this new species. Attribution
of G. ovale sp. n. and other Gadimyxa spp. to the family Parvicapsulidae we consider
to be erroneous, as the shape of typical representatives of this family differs complete-
ly from that characteristic for Gadimyxa genus. We presume it was the molecular stud-
ies that demonstrated the affinities between Parvicapula and Gadimyxa inhabiting the
same organ – the urinary bladder. Similar situation occurred in the case of
Pseudokudoa trifolia Holzer, Blasco-Costa, Sarabeev, Ovcharenko et Balbuena, 2006
(Holzer et al., 2006): the molecular-biological data have shown very high similarity of
this species to Kudoa unicapsula Yurakhno, Ovcharenko, Holzer, Sarabeev et Balbuena,
2007 (Yurakhno et al., 2007), which is completely different morphologically, but occurs
in the same organs – intestine mesenterium and pyloric appendage of the same hosts,
Liza ramada and L. aurata (Yurakhno, 2007).
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P. 743—755.
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559Two New Families and a New Species of Myxosporeans (Myxozoa, Myxosporea)...
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/ConvertImagesToIndexed true
/PassThroughJPEGImages true
/CreateJDFFile false
/CreateJobTicket false
/DefaultRenderingIntent /Default
/DetectBlends true
/DetectCurves 0.0000
/ColorConversionStrategy /CMYK
/DoThumbnails false
/EmbedAllFonts true
/EmbedOpenType false
/ParseICCProfilesInComments true
/EmbedJobOptions true
/DSCReportingLevel 0
/EmitDSCWarnings false
/EndPage -1
/ImageMemory 1048576
/LockDistillerParams false
/MaxSubsetPct 100
/Optimize true
/OPM 1
/ParseDSCComments true
/ParseDSCCommentsForDocInfo true
/PreserveCopyPage true
/PreserveDICMYKValues true
/PreserveEPSInfo true
/PreserveFlatness true
/PreserveHalftoneInfo false
/PreserveOPIComments false
/PreserveOverprintSettings true
/StartPage 1
/SubsetFonts true
/TransferFunctionInfo /Apply
/UCRandBGInfo /Preserve
/UsePrologue false
/ColorSettingsFile ()
/AlwaysEmbed [ true
]
/NeverEmbed [ true
]
/AntiAliasColorImages false
/CropColorImages true
/ColorImageMinResolution 300
/ColorImageMinResolutionPolicy /OK
/DownsampleColorImages true
/ColorImageDownsampleType /Bicubic
/ColorImageResolution 300
/ColorImageDepth -1
/ColorImageMinDownsampleDepth 1
/ColorImageDownsampleThreshold 1.50000
/EncodeColorImages true
/ColorImageFilter /DCTEncode
/AutoFilterColorImages true
/ColorImageAutoFilterStrategy /JPEG
/ColorACSImageDict <<
/QFactor 0.15
/HSamples [1 1 1 1] /VSamples [1 1 1 1]
>>
/ColorImageDict <<
/QFactor 0.15
/HSamples [1 1 1 1] /VSamples [1 1 1 1]
>>
/JPEG2000ColorACSImageDict <<
/TileWidth 256
/TileHeight 256
/Quality 30
>>
/JPEG2000ColorImageDict <<
/TileWidth 256
/TileHeight 256
/Quality 30
>>
/AntiAliasGrayImages false
/CropGrayImages true
/GrayImageMinResolution 300
/GrayImageMinResolutionPolicy /OK
/DownsampleGrayImages true
/GrayImageDownsampleType /Bicubic
/GrayImageResolution 300
/GrayImageDepth -1
/GrayImageMinDownsampleDepth 2
/GrayImageDownsampleThreshold 1.50000
/EncodeGrayImages true
/GrayImageFilter /DCTEncode
/AutoFilterGrayImages true
/GrayImageAutoFilterStrategy /JPEG
/GrayACSImageDict <<
/QFactor 0.15
/HSamples [1 1 1 1] /VSamples [1 1 1 1]
>>
/GrayImageDict <<
/QFactor 0.15
/HSamples [1 1 1 1] /VSamples [1 1 1 1]
>>
/JPEG2000GrayACSImageDict <<
/TileWidth 256
/TileHeight 256
/Quality 30
>>
/JPEG2000GrayImageDict <<
/TileWidth 256
/TileHeight 256
/Quality 30
>>
/AntiAliasMonoImages false
/CropMonoImages true
/MonoImageMinResolution 1200
/MonoImageMinResolutionPolicy /OK
/DownsampleMonoImages true
/MonoImageDownsampleType /Bicubic
/MonoImageResolution 1200
/MonoImageDepth -1
/MonoImageDownsampleThreshold 1.50000
/EncodeMonoImages true
/MonoImageFilter /CCITTFaxEncode
/MonoImageDict <<
/K -1
>>
/AllowPSXObjects false
/CheckCompliance [
/None
]
/PDFX1aCheck false
/PDFX3Check false
/PDFXCompliantPDFOnly false
/PDFXNoTrimBoxError true
/PDFXTrimBoxToMediaBoxOffset [
0.00000
0.00000
0.00000
0.00000
]
/PDFXSetBleedBoxToMediaBox true
/PDFXBleedBoxToTrimBoxOffset [
0.00000
0.00000
0.00000
0.00000
]
/PDFXOutputIntentProfile ()
/PDFXOutputConditionIdentifier ()
/PDFXOutputCondition ()
/PDFXRegistryName ()
/PDFXTrapped /False
/Description <<
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/NLD (Gebruik deze instellingen om Adobe PDF-documenten te maken die zijn geoptimaliseerd voor prepress-afdrukken van hoge kwaliteit. De gemaakte PDF-documenten kunnen worden geopend met Acrobat en Adobe Reader 5.0 en hoger.)
/NOR <FEFF004200720075006b00200064006900730073006500200069006e006e007300740069006c006c0069006e00670065006e0065002000740069006c002000e50020006f0070007000720065007400740065002000410064006f006200650020005000440046002d0064006f006b0075006d0065006e00740065007200200073006f006d00200065007200200062006500730074002000650067006e0065007400200066006f00720020006600f80072007400720079006b006b0073007500740073006b00720069006600740020006100760020006800f800790020006b00760061006c0069007400650074002e0020005000440046002d0064006f006b0075006d0065006e00740065006e00650020006b0061006e002000e50070006e00650073002000690020004100630072006f00620061007400200065006c006c00650072002000410064006f00620065002000520065006100640065007200200035002e003000200065006c006c00650072002000730065006e006500720065002e>
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/SUO <FEFF004b00e40079007400e40020006e00e40069007400e4002000610073006500740075006b007300690061002c0020006b0075006e0020006c0075006f00740020006c00e400680069006e006e00e4002000760061006100740069007600610061006e0020007000610069006e006100740075006b00730065006e002000760061006c006d0069007300740065006c00750074007900f6006800f6006e00200073006f00700069007600690061002000410064006f0062006500200050004400460020002d0064006f006b0075006d0065006e007400740065006a0061002e0020004c0075006f0064007500740020005000440046002d0064006f006b0075006d0065006e00740069007400200076006f0069006400610061006e0020006100760061007400610020004100630072006f0062006100740069006c006c00610020006a0061002000410064006f00620065002000520065006100640065007200200035002e0030003a006c006c00610020006a006100200075007500640065006d006d0069006c006c0061002e>
/SVE <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>
/ENU (Use these settings to create Adobe PDF documents best suited for high-quality prepress printing. Created PDF documents can be opened with Acrobat and Adobe Reader 5.0 and later.)
>>
/Namespace [
(Adobe)
(Common)
(1.0)
]
/OtherNamespaces [
<<
/AsReaderSpreads false
/CropImagesToFrames true
/ErrorControl /WarnAndContinue
/FlattenerIgnoreSpreadOverrides false
/IncludeGuidesGrids false
/IncludeNonPrinting false
/IncludeSlug false
/Namespace [
(Adobe)
(InDesign)
(4.0)
]
/OmitPlacedBitmaps false
/OmitPlacedEPS false
/OmitPlacedPDF false
/SimulateOverprint /Legacy
>>
<<
/AddBleedMarks false
/AddColorBars false
/AddCropMarks false
/AddPageInfo false
/AddRegMarks false
/ConvertColors /ConvertToCMYK
/DestinationProfileName ()
/DestinationProfileSelector /DocumentCMYK
/Downsample16BitImages true
/FlattenerPreset <<
/PresetSelector /MediumResolution
>>
/FormElements false
/GenerateStructure false
/IncludeBookmarks false
/IncludeHyperlinks false
/IncludeInteractive false
/IncludeLayers false
/IncludeProfiles false
/MultimediaHandling /UseObjectSettings
/Namespace [
(Adobe)
(CreativeSuite)
(2.0)
]
/PDFXOutputIntentProfileSelector /DocumentCMYK
/PreserveEditing true
/UntaggedCMYKHandling /LeaveUntagged
/UntaggedRGBHandling /UseDocumentProfile
/UseDocumentBleed false
>>
]
>> setdistillerparams
<<
/HWResolution [2400 2400]
/PageSize [612.000 792.000]
>> setpagedevice
|