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Figure 8 in Five new species of Enchytraeidae (Annelida: Clitellata) from Mediterranean woodlands of Italy and reaffirmed validity of Achaeta etrusca, Fridericia bulbosa and F. miraflores

Figure 8. Fridericia bulbosa (Rosa, 1887) sensu stricto. (A) Anterior body segments (dorsal view); (B) lateral view of segments I–V. (C and D) Clitellum in dorsal (C) and ventrolateral views (D); the elliptical contour in (D) shows the midventral granular field behind the male pores; (E) chylus cells in segment XIV; (F) shallow dorsolateral view of segments IV–V, showing the conspicuous epidermal gland cells; (G) midventral close-up of clitellum revealing the I-shaped male slits and the granular field behind them (contour); (H) dorsal view of segments IV–VI, showing the bent tail of a peptonephridium, the spermathecae and the pharyngeal glands. I, coelomocytes. (A) to (D) from permanent whole-mounted specimens, (E) to (I) from live specimens. Anterior to the right, except in (G) and (H) where anterior is to the top.

Figure 1 in Five new species of Enchytraeidae (Annelida: Clitellata) from Mediterranean woodlands of Italy and reaffirmed validity of Achaeta etrusca, Fridericia bulbosa and F. miraflores

Figure 1. Achaeta borbonica sp. nov. (A) Anterior body half in a dorsal view. Note the intestinal loop in IX; (B) lateral view of segments IV–V, showing the right spermatheca; asterisks indicate the oesophageal dorsal ridge; (C) ventral view of segment V, revealing the closely spaced spermathecal pores; (D) sperm funnel in segment X; (E) lateral view of nephridium in segment VIII; (F) coelomic cavity of caudal segments (lateral view), showing the coarse granulation of the chloragogenous cells and the small size of coelomocytes as compared with the flask-shaped glands. (G and H) Lateral views of clitellum, in vivo (G) and after fixation (H). All except (H) from live specimens. In (A), (D) and (E) anterior to the top; in all others, anterior to the right.

Figure 3 in Five new species of Enchytraeidae (Annelida: Clitellata) from Mediterranean woodlands of Italy and reaffirmed validity of Achaeta etrusca, Fridericia bulbosa and F. miraflores

Figure 3. Achaeta giustii sp. nov. (A) Cephalic region (lateral view); (B) dorsolateral view of segments I–VIII, showing the well-developed pharyngeal glands; (C) dorsal view of clitellum. Note the granular cells bordering the middorsal interruption (dg); (D) shallow lateral view of body wall (segment IX), revealing the lozenge pattern of the longitudinal muscle fibres; (E) lateral view of clitellum, showing the reticulate pattern of the gland cells; (F) lateral view of same clitellum in a deeper optical section, documenting its midventral continuity and middorsal gap; (G) ectal portion of the spermatheca; (H) sperm funnel. All from permanent wholemounted specimens, anterior to the left.

Figure 7 in Five new species of Enchytraeidae (Annelida: Clitellata) from Mediterranean woodlands of Italy and reaffirmed validity of Achaeta etrusca, Fridericia bulbosa and F. miraflores

Figure 7. Fridericia meridiana sp. nov. (A) Anterior body segments (dorsal view). The white arrow points to the small spur of the last pharyngeal glands into VII; (B) clitellum (dorsal view); (C) male pores and midventral interruption of clitellum between and before them; (D) lateral view of clitellum and male opening after fixation; (E) coelomic cavity of segments XI–XII, showing the minute sperm funnel; (F) dorsolateral view of segments IV–V, showing first two pairs of pharyngeal glands and the right spermatheca; (G) coelomocytes; (H) peptonephridium. (A) and (D) from permanent whole-mounted specimens; all others from live specimens. In all, anterior to the right.

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Figs 7–9 in Achaeta Antefolliculata Sp. N., A New Enchytraeid Species (Oligochaeta, Enchytraeidae) From The Rock Grassland Of The Sas-Hegy In Hungary

Figs 7–9. Achaeta antefolliculata sp. n. (photographs of stained whole mounts): 7 = segments 0–VII of the holotype epidermal follicles marked with arrows, 8 = anterior segments, dorsal view, follicles marked with arrow, b = brain, 9 = segments IV–VI, dorso-lateral view; arrow: spermatheca, pg = pri-

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Figs 1–6 in Achaeta Antefolliculata Sp. N., A New Enchytraeid Species (Oligochaeta, Enchytraeidae) From The Rock Grassland Of The Sas-Hegy In Hungary

Figs 1–6. Achaeta antefolliculata sp. n.: 1 = anterior segments (0–VIII) dorsal view (a = oesophageal appendages, b = brain, c = chloragogen cells, dv = dorsal vessel, f = epidermal follicles, n = nephridia, oe = oesophagus, pg = primary pharyngeal glands, ph = pharynx, s = spermatheca, spg = secondary pharyngeal gland), 2 = brain, 3 = spermatheca, 4 = coelomocytes (oblique and upper view), 5 =

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Figs 10–13 in Achaeta Antefolliculata Sp. N., A New Enchytraeid Species (Oligochaeta, Enchytraeidae) From The Rock Grassland Of The Sas-Hegy In Hungary

Figs 10–13. Achaeta antefolliculata sp. n. (photographs of stained whole mounts): 10 = male copulatory organ (penial bulb and glands), lateral view (marked with arrow), 11 = male copulatory organ, ventro-lateral view (marked with arrow), and the granular gland cells of clitellum in rows, 12 = sperm

Table 2 in Five new species of Enchytraeidae (Annelida: Clitellata) from Mediterranean woodlands of Italy and reaffirmed validity of Achaeta etrusca, Fridericia bulbosa and F. miraflores

<p>Table 2. Key morphological characters distinguishing <i>Fridericia rara</i> sp. nov. from <i>F. bulbosa</i> sensu stricto, the only known quadrisetose congener possessing adiverticulate spermathecae, hyaline coelomocytes and short, simple peptonephridia. Measurements in this table refer to live specimens, except where specified by (f), meaning fixed material.</p><table><tbody><tr><th>Species</th><th><i>F. rara</i> sp. n.</th><th><i>F. bulbosa</i> sensu Rota, 1995</th><th><i>F. bulbosa</i> – urban SI</th><th><i>F. bulbosa</i> – urban NA</th></tr></tbody><tbody><tr><th>Length</th><td>5 mm</td><td>7–9 mm</td><td>6 mm</td><td>5–7 mm</td></tr><tr><th>Width (at clitellum)</th><td>0.25 mm</td><td>0.23–0.28 mm</td><td>0.23 mm</td><td>0.22–0.26 mm</td></tr><tr><th>Segments</th><td>27–32</td><td>39–41</td><td>27–37</td><td>24–36</td></tr><tr><th>Chaetal formula</th><td>0,2 – 2: 2,3,4 – 2</td><td>2,3,4– 4,3,2: 2,3,4–4,3,2</td><td>2,3,4 – 2: 2,3,4 – (4,3),2</td><td>2,3,4 – 2: 2,3,4 – 4,3,2</td></tr><tr><th>Chaetal length precl.</th><td>24–27 μm (lat.); 27-35 μm (v.)</td><td>up to 45 μm</td><td>up to 42 μm</td><td>up to 40 μm</td></tr><tr><th>length postcl.</th><td>26–35 μm (lat. & v.)</td><td>50 μm</td><td>up to 42 μm</td><td>40 μm</td></tr><tr><th>Clitellar pattern</th><td>regular rows, chequered, interr. ventrally in XII; cells mostly hyaline, large</td><td>regular rows, ventr. absent between &male; pores, granular behind them, mosaic in XIII; hyaline cells large</td><td>regular rows, absent between &male; pores, granular behind them, mosaic in XIII; hyaline cells large</td><td>regular rows, absent between &male; pores, granular behind them, mosaic in XIII; hyaline cells large</td></tr><tr><th>Male openings</th><td>longitudinal, glandular duct end</td><td>longitudinal</td><td>longitudinal</td><td>longitudinal</td></tr><tr><th>Peptonephridia</th><td>Type a</td><td>Type a</td><td>Type a</td><td>Type a</td></tr><tr><th>Coelomocytes:</th></tr><tr><th>nucleated</th><td>Type a,.24–30 μm long(f)</td><td>Type a, up to 30 μm long</td><td>Type a</td><td>Type a, 32–40 μm long</td></tr><tr><th>anucleate</th><td>small (2.5–6 μm)(f)</td><td>small (4–10 μm)</td><td>small</td><td>small (4–6 μm)</td></tr><tr><th>Chylus cells</th><td>XII–XIV</td><td>XIII–XIV</td><td>XIII–XIV</td><td>XIII–XIV</td></tr><tr><th>Dorsal vessel from</th><td>XV</td><td>XVI–XVII</td><td>XVI</td><td>XV–XVII</td></tr><tr><th>Preclitellar nephridia</th><td>3 pairs (6/7–8/9) midventral</td><td>5 pairs (6/7–10/11)</td><td>5 pairs (6/7–10/11) subterminal</td><td>5 pairs (6/7–10/11)</td></tr><tr><th>Sperm funnels l: w</th><td>88: 50 μm</td><td>152–184: 65 μm 112: 50(f)</td><td>80: 32 μm (f)</td><td>75–100: 42–50 μm</td></tr><tr><th>Penial bulb length</th><td>50 μm(f)</td><td>120 μm, soft, 80(f)</td><td>50–58 μm(f)</td><td>60–64 μm(f)</td></tr><tr><th>Spermathecal ampulla w;</th><td>45 μm; 32(f)</td><td>30 μm; 26(f)</td><td>30 μm; 26–28(f)</td><td>32 μm; 24–29(f)</td></tr><tr><th>ental ducts;</th><td>adjacent;</td><td>adjacent;</td><td>adjacent, on one side;</td><td>merging;</td></tr><tr><th>ectal duct l: w;</th><td>112: 10 μm(f);</td><td>245: 12 μm;</td><td>225: 15 μm;</td><td>175: 15 μm;</td></tr><tr><th>glands at pore</th><td>one, small, 8 μm long(f)</td><td>one, 28 μm long</td><td>one, small, 13 μm long</td><td>one, small, 10 μm long(f)</td></tr></tbody></table>

Table 1 in Five new species of Enchytraeidae (Annelida: Clitellata) from Mediterranean woodlands of Italy and reaffirmed validity of Achaeta etrusca, Fridericia bulbosa and F. miraflores

<p>Table 1. Key morphological characters distinguishing <i>Fridericia meridiana</i> sp. nov. from the known quadrisetose congeners possessing adiverticulate spermathecae, peripherally granular coelomocytes and short, simple peptonephridia. Measurements in this table refer to live specimens, except where specified by (f), meaning fixed material.</p><table><tbody><tr><th>Species</th><th><i>F. meridiana</i> sp. nov.</th><th><i>F. bretscheri</i> Southern, 1907</th><th><i>F. schmelzi</i> Cech & Dózsa-Farkas, 2005</th><th><i>F. composti</i> Schmelz, 2003</th><th><i>F. semisetosa</i> Dózsa-Farkas, 1970</th></tr></tbody><tbody><tr><th>Length</th><td>5–6 mm</td><td>5–10 mm</td><td>5–7.7 mm</td><td>6–8 mm</td><td>5–7 mm</td></tr><tr><th>Width (at clitellum)</th><td>0.19–0.20 mm</td><td>0.25–0.32 μm</td><td>0.22–0.29 mm</td><td>0.30 mm</td><td>0.15–0.24 μm</td></tr><tr><th>Segments</th><td>28–36</td><td>30–42</td><td>34–37</td><td>33–36</td><td>24–34</td></tr><tr><th>Chaetal formula</th><td>2,3 – 2: 2,3,4 – (3),2</td><td>(2,3),4 – 4,3,2: 3,4 – 4,3,2</td><td>(2),3,4 – (3),2: 3,4 – (4),2</td><td>0,1,2,(3,4)</td><td>1,0 – 0: 2,3,4 – (3),2,1</td></tr><tr><th>Chaetal length precl.</th><td>25–40 μm</td><td>up to 40 μm</td><td>up to 29 μm</td><td>up to 40 μm</td><td>20–30 μm</td></tr><tr><th>Max length postcl.</th><td>50 μm</td><td>65 μm</td><td>?</td><td>40 μm</td><td>up to 30 μm</td></tr><tr><th>Clitellar pattern</th><td>regular rows, interrupted ventrally in XII; cells mostly hyaline, large</td><td>irregular; complete; rows of granular cells between male pores</td><td>regular rows, often double, interrupted ventr. in XII; hyaline cells large</td><td>irregular; complete; rows of granular cells between male pores</td><td>regular rows, interrupted dorsally and ventrally; hyaline cells larger</td></tr><tr><th>Male openings</th><td>T-shaped</td><td>longitudinal (staple)</td><td>transversal</td><td>L or T-shaped</td><td>T-shaped</td></tr><tr><th>Peptonephridia</th><td>Type a</td><td>Type a</td><td>Type a</td><td>Type a</td><td>Type a</td></tr><tr><th>Coelomocytes:</th></tr><tr><th>nucleated</th><td>Type b, 18–30 μm long</td><td>Type b, 30–36 μm long</td><td>Type b, 24–34 μm</td><td>Type b, 22–25 μm</td><td>Type b, 20 μm(f)</td></tr><tr><th>anucleate</th><td>small (5–10 μm)</td><td>small (6–10 μm)</td><td>large (8–12 μm)</td><td>large (8–12 μm)</td><td>large (9–12 μm)(f)</td></tr><tr><th>Chylus cells</th><td>IX–X or X–XI</td><td>XII–XV</td><td>IX–X, X–XI or XI–XII</td><td>X–XII</td><td>XII–XIII or XI–XII</td></tr><tr><th>Dorsal vessel from</th><td>XIII–XIV</td><td>XV–XVIII</td><td>XIV–XVI</td><td>XIV</td><td>XIV–XVI</td></tr><tr><th>Preclitellar nephridia</th><td>4 pairs (6/7–9/10) midventral</td><td>4 pairs (6/7–9/10) midventral</td><td>5 pairs (6/7–10/11) midventral</td><td>5 pairs (6/7–10/11) subterminal</td><td>5 pairs (6/7–10/11) midventral</td></tr><tr><th>Sperm funnels l: w</th><td>75: 38 μm</td><td>90–200: 50–100 μm</td><td>48–62: 29–34 μm</td><td>80: 35 μm</td><td>70–110: 37–65 μm</td></tr><tr><th>Penial bulb length</th><td>55 μm</td><td>80–120 μm</td><td>110 μm</td><td>62 μm</td><td>70–90 μm(f)</td></tr><tr><th>Spermathecal ampulla w; ental ducts; ectal</th><td>20 μm(f); separate or adjacent; 175 μm</td><td>43–49 μm; adjacent or merging;</td><td>22–25 μm; merging dorsally;</td><td>25–28 μm(f); merging dorsally;</td><td>22 μm; merging dorsally;</td></tr><tr><th>duct l: w</th><td></td><td>250: 15 μm</td><td>130–160: 10 μm</td><td>100–120: 10 μm(f)</td><td>100: 10 μm(f)</td></tr><tr><th>glands at pore</th><td>1, small, 8–10 μm long(f)</td><td>1or 2, large, 60 μm long</td><td>1, 20–30 μm</td><td>1, 20 μm long(f)</td><td>1, 20–22 μm long</td></tr></tbody></table>

FIGURE 6 in Mata Atlântica enchytraeids (Paraná, Brazil): The genus Achaeta (Oligochaeta, Enchytraeidae)

FIGURE 6. Achaeta singularis sp. nov. A. Juvenile specimen with primordial testis (2) and ovary (3), dorsal view. Dotted areas mark perikarya in brain and ventral nerve cord. The perikarya of the ganglia in II and III are concentrated on the lateral sides. Note nephridia in VI and VII, the oesophageal appendage in V, the dorsal blood vessel in VI, and the winding intestine. From XII on the ganglia are fused among segments; 4: posterior growth zone. The outer pair of postpharyngeal bulbs was not distinguished in this specimen (comp. B). B. Anterior end, lateral view. Perikarya of ventral nerve cord, brain, and prostomial epithelial thickenings are drawn in. Ganglia of II and III are not completely separate, but there is no uniform suboesophageal ganglion over II–IV as in other Achaeta species (comp. Fig. 3B). 1: Prostomial papilla. 2: Oesophageal appendage. 3: Dorsal blood vessel origin. C. Prostomium and first segment, dorsal view. Note the brain, deeply incised anteriorly and posteriorly, and the conspicuous prostomial papillae. Here dotted areas mark the perikarya-free areas. Laterally and posteriad, the prostomial epithelial protrusions gradually merge into the lateral line. D. The brain as seen in living specimens. E. Anterior nephridia, lateral view. - F. Posterior nephridia, ventral view. G. Coelomocytes and piles of rod-shaped structures. D and G are freehand sketches from living specimens, the rest is from stained whole mounts.

FIGURE 5 in Mata Atlântica enchytraeids (Paraná, Brazil): The genus Achaeta (Oligochaeta, Enchytraeidae)

FIGURE 5. Achaeta paranensis sp. nov. A. Anterior body region, lateral view, mature specimen. Spermatheca (1) and nephridium (2) are shaded grey here. The prostomial epithelium is drawn as seen in sagittal optical section; the left prostomial papilla as seen in parasagittal optical section is indicated by a dotted line (3). The region above the dotted line in the brain (4) is filled with perikarya; beneath is the neuropile. Dorsal (5) and ventral (6) epidermal glands are different; lateral glands are not shown here. The circumferal intestinal diverticulum (7) is enclosed by mesodermal tissue and blood lacunae (8); 9: oesophageal appendage; 10: dorsal blood vessel. B. Prostomium with brain and prostomial papillae (1) of a juvenile specimen. C. Coelomocyte, ca. 27 µm long. D, E. Segments IV–VII, lateral view (D) and dorsal view (E), with circumferal intestinal diverticulum and spermathecae. F. Posterior body region with nephridium and pars tumida of midgut epithelium (1); 2: intestinal lumen; 3: ventral nerve cord; 4: chloragocytes. G, H. Hindmost segments with pygidium, in sagittal (G) and parasagittal (H) optical section; sketches from same whole-mounted specimen. I. Clitellar region, lateral view. C–E freehand sketches from living specimens, A, B, F–I from stained whole mounts. Bars =100 µm, bar in A valid also for B, F, I; one bar for G, H. C-E without bar.

FIGURE 1 in Mata Atlântica enchytraeids (Paraná, Brazil): The genus Achaeta (Oligochaeta, Enchytraeidae)

FIGURE 1. Location of the 'Reserva Natural Rio Cachoeira', study area and type locality of Achaeta hanagarthi, A. paranensis and A. singularis spp. nov.

FIGURE 2. Achaeta piti Bittencourt, 1974, specimens from the Cachoeira Natural Reserve. B in Mata Atlântica enchytraeids (Paraná, Brazil): The genus Achaeta (Oligochaeta, Enchytraeidae)

FIGURE 2. Achaeta piti Bittencourt, 1974, specimens from the Cachoeira Natural Reserve. B is a freehand sketch from living specimens, the rest is from stained whole mounts. Bar = 100 µm, valid for A, C, D. A. Submature specimen, side view, anterior 9 segments, with thick body wall and dorsally thick cuticle (1). There are dorsal, lateral and ventral pyriform glands (2,3,4, respectively; only glands of left body side shown). The ventral glands insert more posteriorly than dorsal and lateral glands. The dorsal blood vessel (5) begins at 1/3 VIII in this specimen. The thickened septa in 4/5–7/8 are a constant feature of this species. 6: Oesophageal appendage; 7: pharyngeal gland; 8: postpharyngeal bulb; 9: brain; 10: suboesophageal ganglion; 11: ganglion of segment V. Perikarya are omitted here, they are as in Figs 3B, 4A. 12: nephridia; 13: intestine with chloragocytes; 14: spermatheca with ectal asymmetry; 15: recesses in frontal prostomial epithelium. B. Oesophageal appendage and pharyngeal glands, dorsal view, as seen in a living specimen. The two separately winding canals in IV, one for each glomerulus in V, were already mentioned in the original description. C. Postclitellar nephridia, side view. The organs lie in consecutive segments, one to the left, one to the right. D. Clitellum and male reproductive system, side view, specimen with thick body wall and cuticle. The dorsal pyriform gland of XI is drawn in (1). The clitellar border cells (2) extend farther ventrally than the granulocytes.

FIGURE 3 in Two new species of Achaeta (Enchytraeidae, Oligochaeta) from afforested post-mining and post-fire sites in Poland

FIGURE 3. Achaeta gemmata sp. nov., drawings from whole mounts. A. Anterior end, segments 0–VIII, latero-dorsal view. In this specimen, ventral papillae begin at IV. Pharyngeal glands shaded dark-grey. B. Clitellar region, ventral view.

FIGURE 1 in Two new species of Achaeta (Enchytraeidae, Oligochaeta) from afforested post-mining and post-fire sites in Poland

FIGURE 1. Achaeta florens sp. nov., drawing from whole mounted holotype (A–C) and living non-type specimen (D). A. Head region, lateral view, with epidermal papillae (dark-grey), prostomial ganglion (medium-grey), and head pore. B. Clitellar region, lateral view. C. Segments IV–VII, lateral view. In the original, the ental part of the spermatheca lies between pharyngeal glands and the oesophagus. D. Spermatheca, free-hand drawing.

FIGURE 4 in Two new species of Achaeta (Enchytraeidae, Oligochaeta) from afforested post-mining and post-fire sites in Poland

FIGURE 4. Achaeta gemmata sp. nov., photographs from living specimens. A. Head region, dorsal view, showing prostomial papillae (arrows) antero-laterally of brain (asterisk). B. Nephridium at 7/8; the funnel is covered by the ental reservoir (er) of the spermatheca. C. Spermathecae between pharyngeal glands of IV and V, dorsal view, both ectal pores focused. D. Coelomocytes. E. Clitellum, dorso-lateral view, as seen in a subadult specimen. F. Sperm funnels on both sides of the intestine. G. Lateral papillae (arrows) of segments III–V. H. Clitellum, lateral view. White asterisk: First lateral row with alternating hyalocytes and granulocytes. Dark asterisk: incipient double row of granulocytes. I. Clitellum, dorso-lateral view. Note the absence of a mid-dorsal interruption.

FIGURE 2 in Two new species of Achaeta (Enchytraeidae, Oligochaeta) from afforested post-mining and post-fire sites in Poland

FIGURE 2. Achaeta florens sp. nov., photographs from living specimens. A. Nephridium at 6/7, ventro-lateral view. Asterisks: Dorsal pyriform glands (ventral pyriform glands out of focu). B. Sperm funnels on both sides of the intestine. Asterisks: collar. C. Segments IV, V, ventral view, with ectal part of spermathecae. D. Segment IV, ventro-lateral view. The free-ending ental part of the spermatheca (ct, er) is pushed forwards. E. Segment XII, ventro-lateral view, showing ventral borders of clitellum and male glandular bulbs with central male pores. Ventral nerve cord slightly out of focus. F. Coelomocytes, extruded from the body.

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FIGURE 2. Pseudopolydora achaeta adult morphology. A, B in Cryptic invasions of Pseudopolydora (Annelida: Spionidae), with description of a new species from Queensland, Australia

FIGURE 2. Pseudopolydora achaeta adult morphology. A, B, anterior ends, dorsal view. C, D, anterior end, left lateral view. Scale bars: A, B, D = 200 µm; C = 500 µm. A, Nha Trang Bay, Vietnam (MIMB 40946). B, Paranaguá Bay, Paraná, Brazil (MIMB 40945). C, D—Florida, USA (VIR 24579, photos by Michael Scott Jones).

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FIGURE 4. Pseudopolydora achaeta larval morphology. A–D, 20 in Cryptic invasions of Pseudopolydora (Annelida: Spionidae), with description of a new species from Queensland, Australia

FIGURE 4. Pseudopolydora achaeta larval morphology. A–D, 20-chaetiger larva from plankton: A, dorsal view, in transmitted light; B, same, ventral view; C, dorsal view, in reflected light; D, same, ventral view. Scale bars: A–D = 100 µm. Nha Trang Bay, Vietnam (in life, VIR 25357).