Contributions to Zoology, 78 (3) - 2009Valentina Delogu; Marco Curini-Galletti: The Parotoplana jondelii species-group (Platyhelminthes: Proseriata): a microturbellarian radiation in the Mediterranean

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Appendix

Family OTOPLANIDAE Hallez, 1892

Subfamily PAROTOPLANINAE Ax, 1956

Genus Parotoplana Meixner, 1938

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Parotoplana jondelii Delogu and Curini Galletti, 2007

(Fig. 3A)

Material examined: two whole mounts (SMNH 6666 - holotype; CZM 55) and one specimen sagittally sectioned (CZM 56) from the type locality: Santa Maria di Leuca (Lecce, Apulia, Italy), La Principessa cave, about 5 m deep in coarse sand, May 2005.

Description: The sclerotised apparatus of this recently described species (Delogu and Curini-Galletti 2007), is here re-drawn and re-described, using the terminology of the spine pairs presented above. The species has 18 spines, arranged as follows:

- Dorsal pairs: D1 : straight, needle like, thin spines, 75-79 μm long; D2 : much stouter, straight spines, 70-80 μm long.

- The girdle typically consists of 6 pairs of spines: G1 : similar in size and shape to D2 , but with blunt apices; G2 : two broad spines (62-65 μm long), stout, straight, with falcate apices and small submarginal apophyses; G3 : large straight spines, 75-83 μm long, with straight distal tips, and small subterminal apophyses; G4 -G6 : three pairs of comparatively small spines (53-55 μm long), with apices progressively longer from G4 to G6 ; G5 distinctly slender and straighter than the other pairs.

- V: similar in size to G3 but with longer, more recurved distal tips, with distinctly blunt apices and a more developed, downwards pointing, submarginal apophyses.

In whole mounts, the presence of numerous bursal spines, arranged into two blocks, could be detected. These spines are sharply triangular in shape (up to 4-5 μm long), and weakly sclerotised (Delogu and Curini-Galletti, 2008, Fig. 7B, p. 29).

Karyotype. Chromosome number: n = 6; FN = 12. All chromosomes are isobrachial; the first three pairs are metacentric and distinctly larger than the remaining pairs, at the border between meta- and submetacentric chromosomes. Full karyometrical data are available in Delogu and Curini-Galletti, 2007.

Remarks. The species is only known from the type locality, a marine cave in southern Apulia (Italy), about 5 m deep in medium-coarse sand (Delogu and Curini-Galletti, 2007)

Parotoplana axi sp. n.

(Figs 3B; 4A)

FIG2

Fig. 4. Male sclerotised apparatus of Parotoplana axi (A) (holotype); P. carthagoensis (B) (holotype); P. gracilispina (C) (holotype); P. crassispina (holotype, from Castiglione della Pescaia (D); from Le Bombarde (E); from Canet (G)); P. ichnusae (G) (holotype); P. obtusispina (H) (holotype); P. pulchrispina (I)(holotype). Scale bars: 10 µm.

Holotype: one karyological slide made permanent with lactophenol (SMNH 7573).

Type locality: Catania, Agnone (Sicily, Italy) (lat. 37°18’38.03”N, long. 15°6’24.63”E); lower intertidal, medium-coarse sand, October 2008.

Etymology: the species is dedicated to dr. prof. Peter Ax (Göttingen, Germany), for his long-lasting contribution to the study of the Otoplanidae.

Description: Sclerotised apparatus consisting of 16 spines:

- D1 -D2 : dorsal spines are similar: both pairs are slender, 50-52 µm long, tapering into an acute, outwardly curved point, particularly evident in D2 . As in all the species of the group, D1 is thinner (maximum width: 1,5 µm) than D2 (max. width: about 3 µm).

- G1 -G5 : the girdle consists of five pairs of slender spines, with maximum width of stem about 2 µm, and length ranging from 39 to 46 µm. All have falcate distal tips, conical at least in G4 and G5 , and small, triangular apophyses.

- V: distinctive spines, about 49-51 µm um long, provided with an apex 17-21 µm long, tapering into a narrowly elongate distal tip, and a small (1.5 µm long) apophysis. Stem about 4 µm broad at its basis, narrowing to 1.5 µm just below the apophysis.

Parotoplana carthagoensis sp. n.

(Figs 3C; 4B; 5B)

Holotype: One specimen prepared as a whole mount (SMNH 7566).

Type locality: Sidi Bou Said (Tunis, Tunisia), (lat. 36°51’59.62”N, long. 10°20’52.74”E), about 20 cm deep in medium sand, March 2008.

Paratype: one specimen sagittally sectioned (SMNH 7567).

Additional material: nine specimens sagittally sectioned (CZM 168-176) and two karyological slides (CZM 177-178), all from the type locality.

Etymology: the name refers to the type locality, adjacent to the ruins of the ancient city of Carthago.

Description: The sclerotised apparatus consists of 14 spines (13 in specimen CZM 177, 15 in specimen CZM 178) (Fig. 1B):

- Dorsal pairs: D1 : very slender, straight spines (only one in CZM 177), 40-45 µm long in the holotype, ranging 38-45 µm, with a maximum width of 1 µm; D2 : stout and broader spines, 43-50 µm long (holotype 46-50 µm), 4-5 µm wide at their bases, tapering distally into a sharp point.

- Girdle: G1 : two spines, 39-40 µm long in the holotype (range 37-45 µm), 2-3 µm thick, with falcate apices and pointed distal tip. They are provided with small apophyses, located at about 8 µm from the tip; G2 : two distinctive, straight spines, 40-47 µm long (holotype 43-45 µm), with long, straight apices, with obtuse distal tips, and sharp, upturned apophyses placed at about 14 µm from the tip; G3 -G4 : two pairs of spines (five spines in specimen CZM 176), 37-39 µm in the holotype, ranging 37-41 µm in length and 3-4 µm wide, with falcate apices and obtuse distal tips, and short, broad apophyses placed at about 9 µm from the distal tip.

- V: two spines, ranging 40-45 µm in length (holotype about 40 µm), and 3 µm broad, with long falcate apices and blunt distal tip; apophysis flat, rather well developed (3-5 μm long) placed at about 12 µm from the tip.

The species is provided with numerous, broadly triangular bursal spines, 2 to 3 µm in length, 3 to 4 μm wide at their bases, arranged in concentric girdles, that, in a few specimens, are fused into a single tube, about 25 µm long and 20 µm wide (Fig. 4 B).

Karyotype. Chromosome number: n = 6; FN = 11. The first three pairs of chromosomes are distinctly larger than the three remaining pairs. Karyometrical data (based on measurements of three plates): Chrom. I = r.l.: 26.09 ± 1.44 ; c.i.: 44.95 ± 0.96 (m); Chrom. II = r.l.: 25.48 ± 1.11; c.i.: 33.92 ± 0.7 (sm); Chrom. III = r.l.: 23.6 ± 1.68; c.i.: 42.41 ± 1.48 (m); Chrom. IV = r.l.: 9.02 ± 1.41; c.i.: 30.74 ± 6.42 (sm); Chrom. V = r.l.: 8.32 ± 0.86; c.i.: 45.27 ± 2.54 (m); Chrom. VI = r.l.: 7.42 ± 1.28; c.i.: 10.41 ± 2.97 (t); haploid genome length: 7.4 ± 0.5 µm.

Remarks. A common species in shallow water in the Sidi Bou Said harbour area, but not found elsewhere during extensive samplings performed along the Tunisian eastern coast during BIOIMPA campaign (2008).

Parotoplana crassispina sp. n.

(Figs 1; 2; 3E-G; 4D-F; 5A, C-F)

FIG2

Fig. 5. A-B. Bursal sclerotised apparatus (“nozzles”). A: Parotoplana crassispina (from Castiglione della Pescaia); B: P. carthagoensis. C-F. Genital area of P. crassispina. C-D: from living, slightly squeezed specimens (C: Le Bombarde; D: Castiglione della Pescaia). E-F: from sagittal sections of specimens from Castiglione della Pescaia. Scale bars: A, B: 10 µm; C-F: 25 µm.

Holotype: one whole mount (SMNH 7568).

Type locality: Castiglione della Pescaia, Capo Capezzolo (Tuscany, Italy) (42°45’55.29”N, long. 10°51’46.11”E); lower intertidal, medium sand, March 2007.

Paratype: one specimen from the type locality sagittally sectioned (SMNH 7569).

Additional material: Tuscany: Castiglione della Pescaia,CapoCapezzolo. Six specimens sagittally sectioned (CZM 121-126) and four karyological slides made permanent with lactophenol (CZM 127-130), March 2007. Punta Ala, Torre Civette (lat. 42°50’41.70”N, long. 10°46’30.34”E), lower intertidal, medium to coarse sand. 14 specimens sagittally sectioned (CZM 131-144) and eight karyological slides made permanent with lactophenol (CZM 145-152), March 2007. Sardinia: Le Bombarde beach, Alghero (lat. 40°35’2.59”N, long. 8°15’37.36”E), in medium to coarse sand, from the lower intertidal to about 2 m deep. One specimen sagittally sectioned (CZM 153) and five karyological slides made permanent with lactophenol (CZM 154-158). Several specimens observed alive, July 2006. Archi Cave, Capo Caccia (lat. 40°34’10.71”N, long. 8°13’43.93”E), medium sand, about 10 m deep. One specimen made as whole mounth (CZM 159), July 2006. Girin beach, Carloforte Is. (lat. 39°6’56.68”N, long. 8°18’33.33”E), about 50 cm deep, medium sand. Three specimens made as whole mount (CZM 160-162), June 2008. France: Canet, city beach (lat. 42°41’41.36”N, long. 3°2’14.98”E), about 2 m deep, medium sand. Five specimens made as whole mounts (CZM 163-167), August 2008.

Etymology: from latin crassus, fat – with reference to the peculiar, broad spines of the new species.

Description: With 14 copulatory spines, arranged as follows:

- Dorsal pairs: D1 : thin, straight, needle-shaped spines, ranging 45-58 μm long in specimens from Tuscany (holotype: 56-58 μm) with maximum width at the basis of about 2.5 μm; and 42-48 μm and 51-53 μm long in specimens from Sardinia (Le Bombarde) and Canet, respectively; D2 : broad, straight spines, ranging 47-64 μm long (holotype: 59 µm) and 6-7 μm broad at their basis (Tuscany); 42-50 μm long, 4.5 μm broad (Sardinia); 53-56 μm long, 6 μm broad (Canet).

- Girdle: G1 : short, straigth spines with slightly falcate, conical apices, provided with very small apophyses, ranging 37-44 μm long, 5-6 μm wide (Tuscany) (Holotype: 41-44 µm; distance between apophysis and tip about 8.5 µm; stem 5.5 µm wide); 37-43 μm long, 4.5 μm wide (Sardinia); 41-46 μm long, 4.5 μm wide (Canet). Sardinian specimens have a comparatively longer (about 1/3 of the whole length of the spine) distal tip; G2 -G4 : three pairs of broad spines, with characteristic flat and broad apophyses, falcate distal tips and blunt apices. Stems become progressively longer from G2 to G4 , and spines gradually assume a more slender shape. Lengths range 32-52 μm long (Holotype: 42-50 µm), stem 3.5-6.5 μm broad, length of apex 11-13 μm (Tuscany); 37-46 μm long, 3.5-5 μm wide, length of apex 9-14.5 μm (Sardinia); 42-47 μm long, 3.5-5 um wide, length of apex 11-12 μm (Canet).

- Ventral spines differ in morphology among populations. In Tuscan specimens, V spines are 47-65 μm long (holotype: 58-59 µm), 7.5 μm broad at their bases, with straight, very long (18-24 μm) distal tips, and long (11-20 μm) and flat apophyses. Sardinian and French specimens have more recurved and comparatively shorter distal tips, and shorter, ovoid apophyses. Ranges are 47-55 μm long, 7 μm broad, distal tip 11-19 μm long, apophysis 9-12 μm long (Sardinia) and 51-53 μm long, 5-6 μm broad, distal tip 12-17 μm long; apophysis 6-8 μm long (Canet).

With numerous bursal spines, arranged in girdles. Each girdle consists of up to 30 broadly triangular spines, with acute tips, increasing in length from proximal (about 1 μm) to distal (about 5 μm) girdles (Fig. 5A). The arrangement of spines is somewhat variable. In a few specimens they are arranged into a single structure, consisting of about six girdles of spines. In other specimens girdles are less regular, while in others two distinct series of girdles are apparent. However, the level of the variability (either individual or interpopulational) has yet to be ascertained.

Karyotype. Chromosome number: n = 6; FN = 10. The first three pairs of chromosomes are distinctly larger than the three remaining pairs. Karyometrical data were obtained from two populations. Castiglione della Pescaia (Tuscany): Chrom. I = r.l.: 25.46 ± 2.86 ; c.i.: 35.4 ± 1.79 (sm); Chrom. II = r.l.: 24.03 ± 0.84; c.i.: 41.52 ± 3.28 (m); Chrom. III = r.l.: 23.35 ± 0.61; c.i.: 32.39 ± 2.53 (m); Chrom. IV = r.l.: 9.85 ± 1.13; c.i.: 36.95 ± 4.59 (sm); Chrom. V = r.l.: 8.72 ± 0.26; c.i.: 15.93 ± 4.39 (st); Chrom. VI = r.l.: 8.61 ± 1.57; c.i.: 10.12 ± 1.43 (t); haploid genome length: 9.6 ± 0.8 µm (based on measurements of four plates). Le Bombarde (Sardinia): Chrom. I = r.l.: 25.47 ± 2.43 ; c.i.: 44.08 ± 2.09 (m); Chrom. II = r.l.: 25.32 ± 2.63; c.i.: 40.06 ± 0.84 (m); Chrom. III = r.l.: 24.67 ± 0.36; c.i.: 33.15 ± 2.57 (m); Chrom. IV = r.l.: 8.66 ± 0.97; c.i.: 20.8 ± 4.08 (st); Chrom. V = r.l.: 8.43 ± 1.41; c.i.: 39.15 ± 2.73 (m); Chrom. VI = r.l.: 7.45 ± 0.55; c.i.: 8.97 ± 0.95 (t); haploid genome length: 10.1 ± 0.9 µm (based on measurements of five plates).

Remarks.A shallow water species, often common in medium-grained sediments up to the low water mark. In this habitat, it may be the numerically dominating proseriate, as in the Tuscan sites.

P. crassispina is the only species of the group known for a comparatively broad range, extending across the north-western Mediterranean region. Populations in this range appear to differ noticeably, particularly in the morphology of V and G1 spines. The suspicion that different species have been lumped into a single taxon, may thus arise. However, the range of variation of the spines, and thus the reliability as diagnostic markers of the differences observed, should be assessed on more adequate geographical sampling than that presently available. At the moment, given the overall similarity of most spine pairs across the range, and the overlap in measures among populations, differences in morphology are assumed to reflect a degree of interpopulational (clinal?) divergence.

Parotoplana gracilispina sp. n.

(Figs 3D; 4C)

Holotype: one karyological slide made permanent with lactophenol (SMNH 7599).

Type locality: Atlit (Israel), beach near the Castle (lat. 32°42’21.01”N, long. 34°56’6.90”E), lower intertidal, medium sand, February 1986.

Additional material: one karyological slide made permanent with lactophenol (CZM 184)

Etymology: the name refers to the slender (latin gracilis) appearance of the copulatory spines.

Description: The sclerotised apparatus consists of 15 spines in the holotype, and 14 in CZM 184:

- Dorsal pairs: D1 : slender, straight, spines ranging 43-44 µm long, with a maximum width at their bases of 1 µm; D2 : stouter spines, 47-51 µm long, about 3 µm wide at their bases, tapering distally.

- Girdle: G1 : two spines, 37-39 µm long, 2-3 µm wide in their middle, tapering towards their bases. With slender, slightly falcate apices and obtuse distal tips. They are provided with small apophyses, located at about 7 µm from the tip; G2 : two straight spines, 46-48 µm long, about 4.5 µm wide in their middle, and tapering basally to 1 µm wide. With very long, slender, straight apices with obtuse distal tips, and with very small apophyses, placed at about 20 µm from the apex; G3 -G4 : ranging in length from 41-44 µm (G3 ) to 48-51 µm (G4 ); stems 2-3 µm wide in their middle. Both pairs have long, falcate apices, with obtuse distal tips and short apophyses placed at about 10-16 µm from the distal tip.

- V: two spines, ranging 44-48 µm in length, and 2.5 µm broad, with very long, slender, falcate apices and blunt distal tip; apophysis flat, 3-3.5 µm long, placed at 17-20 µm from the tip.

No bursal spines were retrieved in the specimens studied.

Karyotype. Chromosome number: n = 6; FN = 10. The first three pairs of chromosomes are distinctly larger than the three remaining pairs. Karyometrical data (based on measurements of nine plates): Chrom. I = r.l.: 25.92 ± 2.13 ; c.i.: 37.13 ± 0.91 (sm); Chrom. II = r.l.: 24.6 ± 1.42; c.i.: 29.03 ± 3.76 (sm); Chrom. III = r.l.: 23.85 ± 2.56; c.i.: 45.24 ± 1.27 (m); Chrom. IV = r.l.: 8.93 ± 1.21; c.i.: 18.07 ± 5.91 (st); Chrom. V = r.l.: 8.84 ± 1.15; c.i.: 28.48 ± 5.81 (sm); Chrom. VI = r.l.: 7.76 ± 2.05; c.i.: 12.18 ± 5.81 (t); haploid genome length: 11.4 ± 0.9 µm.

Parotoplana ichnusae sp. n.

(Figs 3H; 4G)

Holotype: one karyological slide made permanent with lactophenol (SMNH 7570).

Type locality: Le Bombarde beach, Alghero (Sardinia, Italy) (lat. 40°35’1.50”N, long. 8°15’38.05”E), about 2 m deep, medium-fine sand, May 2008

Additional material: Porto Alabe (Sardinia) (lat. 40°14’46.87”N, long. 8°28’28.49”E), about 1.5 m deep, medium sand.One karyological slide made permanent with lactophenol (CZM 177), June 2008.

Etymology: the species epithet refers to the ancient greek name of Sardinia, Hyknusa (latinised as Ichnusa).

Description: With a sclerotised apparatus consisting of 16 spines:

- Dorsal pairs: D1 : two very thin, needle shaped, straight spines, 62 µm long in the holotype (range 61-65 µm), ending into a sharp distal tip. Stem about 1.5 µm broad; D2 : two broad, straight spines, 62-64 µm long in the holotype (range 60-65 µm) slightly tapering distally into a rather obtuse tip. Stem about 4.5 µm thick.

- Girdle: G1 : similar to D2 , but with a blunter apex, and with a very small apophysis 13-15 µm from the slightly curved distal tip. 64 µm long in the holotype, ranging 60-65 µm; stem 5.5 µm broad; G2 : transient between the morphology of G1 and the rest of the spines of the girdle. 63-65 µm long in the holotype (range 61-65 µm) with a comparatively very long slightly curved apex, with an obtuse distal tip, and with a small apophysis at about 23 µm from the apex. Stem 7.5 µm broad at its maximum, about 5 µm at its basis; G3 -G5 : three pairs of rather similar spines, 76-80 µm long in the holotype (range 72-81 µm) with falcate apices progressively shorter and with small, obtuse subterminal apophyses. Stems ranging 6-8 µm in width.

- V: slender spines (79-81 µm long in the holotype, ranging 70-81 µm), with small falcate apices, and obtuse distal tips. They are provided with a narrow, upward turned apophysis, about 3 µm long. The distance between apices and apophysis is about 12 µm. Stem about 4 µm broad, narrowing to 1.5 µm at its basis.

Without bursal spines.

Karyotype. Chromosome number: n = 6; FN = 11. The first three pairs of chromosomes are distinctly larger than the three remaining pairs. The only plate suitable for karyometric analysis from the specimen from Le Bombarde yielded the following data: Chrom. I = r.l.: 24.84; c.i.: 44.72 (m); Chrom. II = r.l.: 24.36; c.i.: 30.71 (sm); Chrom. III = r.l.: 24.31; c.i.: 40.67 (m); Chrom. IV = r.l.: 9.32; c.i.: 31.83 (sm); Chrom. V = r.l.: 9.32; c.i.: 18.18 (st); Chrom. VI = r.l.: 8.75; c.i.: 46.37 (m); haploid genome length: 9.4 µm.

Parotoplana obtusispina sp. n.

(Figs 3I; 4H)

Holotype: one karyological slide made permanent with lactophenol (SMNH 7572).

Type locality: St. Cyprien, France (lat. 42°37’ 45.97”N, long. 3°2’11.55”E), about 2 m deep, medium sand, August 2008.

Additional material: three karyological slides made permanent with lactophenol (CZM 181-183).

Etymology: the name refers to the blunt (latin obtusus) apices of the spines.

Description: The sclerotised apparatus consists of 14 spines (15 in specimen CZM 181) arranged as follows:

- Dorsal pairs: D1 : straight spines, with obtuse distal tips, 70-72 µm long in the holotype, ranging 69-73 µm in the sample; stem about 3 µm broad; D2 : straight spines, distinctly broader than D1 , with a recurved, obtuse distal tip. 71-73 µm long in the holotype, ranging 70-73 µm; stem about 7.5 µm broad; narrowing basally to 2 µm.

- Girdle: G1 : straight spines, 63 µm long in the holotype (range: 61-65 µm), without apophysis, with very narrow bases (about 1 µm broad), widening progressively to 4 µm towards the subapical region. Apex provided with an obtuse distal tip. G2 : similar to G1 , 67 µm long in the holotype, ranging 67-70 µm, with narrow bases, about 1.5 µm wide, widening distally to a maximum width of 6 µm at about 20 µm from the apex, which is nearly straight, and provided with an obtuse distal tip. Without distinct apophysis. G3 -G4 : two pairs of similar spines, about 60 µm long, with a maximum width of the stem of 4-5 µm, and narrowing towards a basis about 2 µm wide. Both pairs have very short (about 5 µm long) obtuse apices. With small apophyses, more developed in G4 .

- V: 55 µm long in the holotype, ranging 54-59 µm. Straight, rod-shaped spines, with a slightly recurved, obtuse distal tip. With a flat subterminal apophysis, inserted at about 13 µm from the apex.

Parotoplana pulchrispina sp. n.

(Figs 3L; 4I)

Holotype: one karyological slide made permanent with lactophenol (SMNH 7571).

Type locality: Le Bombarde beach, Alghero (Sardinia, Italy) (lat. 40°35’1.50”N, long. 8°15’38.05”E), in medium-fine sand, about 2 m deep, May, December 2008.

Additional material: two karyological slides made permanent with lactophenol (CZM 178-179) and one specimen sagittally sectioned (CZM 180), all from the type locality.

Etymology: the name refers to the graceful, elegant morphology of the spines (latin pulcher = fine, beautiful).

Description: The sclerotised apparatus consists of numerous spines (21 in the holotype, 20 in CZM 178 and 22 in CZM 179).

- dorsal pairs D1 : two thin, needle shaped spines, 52-53 µm long in the holotype (range: 46-53 µm); stem 2-3 µm wide; D2 : two broad, straight spines, tapering into an acute point, 55-56 µm long in the holotype (range: 51-56 µm); stem 4-5.5 µm wide in the middle, and about 3 µm at its basis.

- Girdle: G1 -G6 : A somewhat variable number of similar, straight spines (11 in the holotype, 12 in CZM 179, 10 in CZM 178), ranging 42-54 µm in length in the sample, with slightly falcate apex, ending into an obtuse distal tip, and narrow, upturned subterminal apophyses, 2-5 µm long. Stem width ranging 1.5-2.5 µm; G7 : a pair of spines with straight, pointed apex, and very small apophysis inserted at about 14 µm from the distal tip. 49-50 µm long in the holotype (range: 50-53 µm); stem 3-3.5 µm wide in the middle, about 1.5 µm at its basis; G8 : very similar to G7 , but with more acute and slender, comparatively shorter (about 10 µm long) apex. 44-46 µm long in the holotype, ranging 44-51 µm in the whole sample; stem 3-3.5 µm broad.

- V: medially to pair G8 , all specimens had a pair of smaller spines, similar in morphology to G2 -G7 spines, but with straighter distal tip, and with very small, obtuse apophyses. In the holotype, these spines are 42-46 µm long, with a thin and feebly sclerotised stem. In the rest of the sample, they are about 46 µm long, with a stem slightly widening towards its base, which is 3 µm across.

Without bursal spines.

Karyotype. Chromosome number: n = 6; FN = 10. The first three pairs of chromosomes are distinctly larger than the three remaining pairs. Karyometrical data (based on measurements of four plates): Chrom. I = r.l.: 25.23 ± 1.19 ; c.i.: 42.45 ± 1.33 (m); Chrom. II = r.l.: 24.89 ± 0.06; c.i.: 38.01 ± 1.46 (m); Chrom. III = r.l.: 24.02 ± 1.24; c.i.: 32.5 ± 1.77 (sm); Chrom. IV = r.l.: 8.39 ± 0.65; c.i.: 15 ± 4.36 (st); Chrom. V = r.l.: 8.13 ± 0.98; c.i.: 36.43 ± 2.88 (sm); Chrom. VI = r.l.: 7.68 ± 0.74; c.i.: 11.3 ± 4.41 (t); haploid genome length: 10 ± 1 µm.