Fridericia connatiformis, Dozsa-Farkas, 2015

Dózsa-Farkas, Klára, Ortmann-Ajkai, Adrienne & Horváth, Ferenc, 2015, New Enchytraeid Species (Oligochaeta: Enchytraeidae) From The Danube-Dráva National Park, Acta Zoologica Academiae Scientiarum Hungaricae 61 (4), pp. 305-327 : 307-312

Fridericia connatiformis Dózsa- Farkas, sp. n.

( Figs 1A–D, 2–4)

Type material – Holotype: F.22. slide No. 721. Type locality: Páprád , Bükkhát Forest Reserve, Danube –Dráva National Park, 45°52’19.4”N, 18°00’47.4”E, 109 m a.s.l., riverine oak- elm- ash Woodlands, leg. Ortmann- Ajkai, A., 08. 11. 2012.

Paratypes (intotal 15 specimens): P . 99.1 slideNo. 1142, onespecimenfromtypelocal- ity leg. Dózsa- Farkas, K ., Ortmann- Ajkai , A . & Farkas , J., 28. 03. 2011; P . 99.2–99.8. slide No. 715, 720, 747, 748, 749, 750, 755, seven specimens, from type locality, leg. Ortmann- Ajkai, A., 08. 11. 2012; P . 99.9. slide No. 746, from type locality, leg. Dózsa- Farkas, K ., Farkas , J., 13. 11. 2013; P . 99.10. slide No. 740. One specimen, Bükkhát Forest Reserve, 45°52’35.2”N, 18°00’41.3”E, 127 ma.s.l., loWlandpedunculateoak- hornbeamWoodlandsunder Ruscusaculeatus shrubs, leg. Dózsa- Farkas, K ., Ortmann- Ajkai , A . & Farkas , J., 28. 03. 2011; P . 99.11. slideNo. 742. onespecimen, betWeenvillageSellyeandBogácsa, 45°52’24.6”N, 17°48’01.1”E, 104 ma.s.l., meadoWWithyoung Quercus plantation, leg. Dózsa- Farkas, K ., Ortmann- Ajkai , A . & Farkas , J., 28. 03. 2011; P . 99.12 slide No. 583 one specimen, Tivadar, 48°03’48”N, 22°31’18”E, 108 ma.s.l., meadoWnearTiszariver, leg. Dózsa- Farkas, K . & Farkas , J., 8. 10. 2000; P . 99.13 slide No. 719 one specimen, Markaz, Mátra Mts, 47 o 50 ’ 47 ” N, 20 o 03’08 ” E, 450 ma.s.l., oakWoodland, leg. Dózsa- Farkas, K . & Farkas , J., 31. 03. 2003; P . 99.14. slideNo. 981 oneadultspecimen, Szőce , Őrség, 46 o 54’02”N, 16 o 34’29”E, 229 ma.s.l., sphagnumbog, leg. Dózsa- Farkas, K ., Farkas , J ., Tóth , Z . & Hock , F., 31. 03. 2014; P . 99.15. slideNo. 984 oneadultspecimen, Szentgyörgyvölgy , Őrség, 46 o 45’14”N, 16 o 25’01”E, 238 m a.s.l., hayfield, leg. Dózsa- Farkas, K ., Farkas , J ., Tóth , Z . & Hock , F., 31. 03. 2014 .

Etymology – NamedafteritssimilarityWith F. connata .

Diagnosis – TheneWspeciescanberecognizedbythemainfolloWing combination of characters: (1) large size (15–23 mm in vivo), segments 45–65; (2) tWochaetae (rarelyone) perbundle; (3) bodyWallthick; (4) fiveprecli- tellarnephridia; (5) coelomo- mucocytesc/btype, lenticytessmall; (6) clitel- lumsaddle- shaped, hyalocytesandgranulocytesarrangedintransverseroWs dorsally; (7) bursalslitT-shaped; (8) largeseminalvesicle; (9) spermfunnel cylindrical, approximately 3–4 times as long as Wide in vivo, (2–3 times When fixed), collarasWideasthefunnelbody, spermca 500 μmlong; (10) subneural gland present in XIII and XIV; (11) the most important characteristic feature, apartfromthesubneuralglands, largeextrasexualglandsin 4/5 andinXI, (rarely in VII and XIII); (12) spermathecae separate entally, ampullae Wide, WithtWolargediverticulaonshortstalks.

Description – Large Worm, holotype 13.1 mm long, 690 μm Wide at VIII and 710 μm attheclitellum (fixed), segments 66. Bodylengthofparatypes 14–23 mm, Width 450–540 μm at VIII and 570–700 μm at the clitellum (in vivo), length of fixed specimens 11.4–19.9 mm, Width 470–690 μm at VIII, 530–710 μm at the clitellum, segments 48–66. Chaetae tWo perbundlethroughout, occasionally 1, formula: (1),2 – 2,(1): 2 – 2. Lengthofchaetaeabout 55–60 μm, diameter 6.5–7 μm preclitellarly and 60–70 × 6.5–8 μm postclitellarly. Detached chaetaeoftenpresentincoelom. Headporeat 0/I. DorsalporesfromVII. LightbroWn epidermalglandcellsarrangedin 2–3 transverseroWspersegments (Fig. 3A). BodyWall thick, about 45–50 μm, cuticleabout 3 μm). ClitelluminXII–1/2XIIIsaddle- shaped, the hyalineandgranularglandcellsarrangedindensetransverseroWsdorsally (Figs 2A–B). Brain ( Fig. 1A) egg-shaped, about 2–2.5 timeslongerthanWide (about 180–200 μmlong). Oesophagealappendages (peptonephridia) ( Fig. 1D) longWithWidelumenandWith 3–5 terminalbranches. Pharyngealglandsalluniteddorsally, allWithventrallobes. Chlorago- cytesfromV, darkbroWn (invivo). DorsalbloodvesselfromXIX–XXI, bloodcolourless. Fivepairsofpreclitellarnephridiafrom 6/7 to 10/11. Coelomo- mucocytes (length 40–57 μm invivo) typec/b (accordingtoMÖLLER 1971), finegranularmatrixandclearlyvisible nucleus, sometimesWithafeWrefractilevesicle, lenticytessmall (3–5 μmlong) andscarce (Figs 2C–D). ChyluscellsbetWeenXVI–XVIII, occupyingthreesegments. Seminalvesicle largeinX–XI. Spermfunnels ( Figs 1B, 2E) cylindrical, 3–4 timesaslongasWide, about 500–600 μmlong (invivo). Infixedspecimensthelengthoffunnelsare 225–400 μm. Collar as Wide as funnel body. Spermatozoa about 350–500 μm long, heads 120–130 μm (in vivo). Vasdeferens 8–9 μmWide. Malecopulatoryorgans (Fig. 2F) onaverage 156 μmlong, 84 μmWideand 67 μmhigh, bursalslitT-shaped (Fig. 2H). Well- developedsubneuralglands inXIII–XIV (Fig. 2G). Characteristicextrasexualglands (copulatoryglands) at 4/5 nearthe specimens, scalebares = 100 μm)

spermathecal orifice, round (diameter about 90–100 μm, fixed) mostly on one side only, laterally (Figs 3A–B), and one oval gland in XI mid- ventrally (145–175 × 80–120 μm, fixed) (Figs 2F, 3C). In one case gland in VII instead of at 4/5 (slide No. 742). Sometimes in XIII also one oval gland (150–176 × 90 μm, fixed) just before the subneural gland of XIII. Sper- matheca (Figs 3D–H): ectal ducts as long as body diameter (about 300–450 μm long and 24–25 μm Wide, the canals Wide (6 μm) in vivo, ducts 250–330 μm long and 20–26 μm Wide in fixed specimens. One small sessile (ca 25 μm long) gland at the orifice of the spermath- Fig. 2. Micrograph of Fridericia connatiformis sp. n: A–B = clitellar gland cells dorsally, C–D = coelomocytes, E = sperm funnels, F = XI–XII ventrally (extra sexual gland marked With White arroW, male copulatory organs marked With black arroWs), G = subneural glands in XIII and XIV, H = bursal slit marked With arroW. (A, C, E, H in vivo, B, D, F, G fixed, stained;

scale bars = 50 μm)

ecalducts. DuctprojectingintoampullaWithoutentalbulb. Ampullamostlyaslongas Wide, about 140–170 μmWide (fixed), thick- Walled, containingfinepalegranules, tapering proximad, noseparationintodistalandproximalpart. Ampullarlumennot “U” form. The communicationoftheampullaeWiththeoesophagusisseparate. TWosphericaldiverticula With short stalks, thin- Walled (diameter 75–110 μm in vivo and 70–105 μm, fixed), mostly filledWithbroWnsperm (intransmittedlight) spermnotrotating, diverticulasometimes empty. OneortWomatureeggsatatime.

Fig. 3. Micrograph of Fridericia connatiformis sp. n.: A = epidermal glands in transverse roWs and an extra sexual gland in V (marked With arroW), B = tWo extra sexual glands in V (marked With arroWs), C = extra sexual glands in XI ventrally, D–H spermathecae (in D, F and H the diverticula full With sperm, in E and G the diverticula empty). (A, B, D–F in vivo, C, G fixed,

stained, H one removed spermatheca of the fixed, stained specimens; scale bars = 50 μm) largermalecopulatoryorgans. (A–B, stainedspecimens; scalebars = 50 μm)

Distributionandhabitat – DrávaFloodplain, Danube–DrávaNationalPark, Páprád, BükkhátForestReserve, riverineoak- elm- ashWoodlands, 45°52’19.4”N, 18°00’47.4”E, 109 masl; nearTiszaRiver, Tivadar, 48°03’48”N, 22°31’18”E, 108 ma.s.l., meadoW; MátraMts, Markaz, 47 o 50 ’ 47 ” N, 20 o 03’08”E, 450 ma.s.l., oakforest; Őrség: Szentgyörgyvölgyhayfield, 46 o 45’14”N, 16 o 25’01”E, 238 ma.s.l.; Szőce, sphagnumbog, 46 o 54’02”N, 16 o 34’29”E, 229 ma.s.l.

Differentialdiagnosis – DÓZSA-FARKAS (2009) classifiedthe Fridericia spe- ciesWithtWospermathecaldiverticulaingroups. IngroupII.1., thespermath- ecal diverticula are globular or hemispherical, sessile or With short stalks (11 species) ( DÓZSA-FARKAS 2009, table 3, 4). Only three species of this group have only tWo chaetae in the chaetal bundles: Fridericia cylindrica Springett, 1971 , Fridericia sohleni Rota, Healy et Erséus, 1998 , and Fridericia bisetosa (Levinsen, 1884) . All three species are smaller (10–15 mm long) than the neW species, subneuralglandsabsent, thespermfunnelsalsosmallerandonly 1.5–2× as long as Wide. Moreover in the first tWo species the collar is narroWer, and the mucocytes are of the b-type. Most similar to the neW species is F. bisetosa but the spermathecal ampullae of the neW species differs in shape and size and the diverticula are much larger ( SCHMELZ 2003). The neW species is also similar totWospeciesofgroupI.1, With’ TheproximalpartofthetWospermathecal ampullafused, onecommonopeningintooesophagus’ ( DÓZSA-FARKAS 2009), F. connata Bretscher, 1902 and F. brunensis Schlaghamersk ỳ, 2008. F. connata is similar to the neW species in size, in the number of chaetae (also not more than tWo in the bundles), in the thick preclitellar septa and someWhat in the shape ofthespermathecae, butthestalksofthespermathecaldiverticulaarelonger and With a characteristic bend toWards the ectal duct, the diverticular lumen is semicircular in top vieW, and the ampullar lumen forms a “U” around the bulbiform ental bulb of the ectal ducts ( Fig. 4A). Moreover in F. connata the tWospermathecalampullaearefused, Withonecommonopeningintothe oesophagus, andthemalecopulatoryorganslarger ( Fig. 4B). F. connata (after oWnobservationsinHungarianpopulations) mostlyhasyelloW-greencoelo- mocytes, c- typeWithsomerefractilegranules. F. brunensis haveonly 1 chaeta regularlyinposteriorlateralbundles, havetWospermathecaldiverticula, but the tWo spermathecal ampullae fused and the chylus cells in IX–XI praeclitel- larly ( SCHMELZ 2003, BRETSCHER 1902, SCHLAgHAMERSKÝ 2008). Only F. galba has similar “accessory sexual glands” ( SCHMELZ 2003, p. 54), but this species based on the shape of his spermatheca and the number of chaetae in bundles cannot be confused With F. connatiformis . The F. connatiformis differs from all Fridericia speciesbythepresenceofextrasexualglandsandsubneuralglands, itisan uniquecombinationandalsounusualthatsuchalarge Fridericia specieshas pharyngealglandsalluniteddorsally ( SCHMELZ 2003).