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Additional file 2.

Additional file 1.

Petroscolex centenarius Csuzdi, Szederjesi & Sherlock, sp. nov. (Figures 2, 3) Holotype. NHMUK 1997.1594 clitellate adult. DR Congo, Kivu, Irangi. Leg. Gazana N���Doli, 20.02.1973. Paratypes. HNHM AF/3555 1 ex. clitellate adult, NHMUK 1997.1595 1 ex. clitellate adult broken in two pieces. Locality and date same as of the Holotype. NHMUK 1997.1548 ��� 1549 1 ex. clitellate adult, 1 ex. juvenile, HNHM AF/3539 1 ex. clitellate adult, DR Congo, Kivu, Irangi. Leg. Gazana N���Doli, 08.03.1973. NHMUK 1997.1526 ��� 1529 1 ex. clitellate adult, 1 ex. aclitellate adult, 2 ex. juvenile, NHMUK ANEA 2019.7368 1 ex. clitellate adult, HNHM AF/5754 1 ex. clitellate adult, DR Congo, Kivu, Irangi. Leg. Gazana N���Doli, 09.03.1973. NHMUK 1997.1530 1 ex. aclitellate adult, DR Congo, Kivu, Irangi. Leg. Gazana N���Doli, 07.06.1973. Etymology. The specific epithet refers to the 100th birthday of Prof. Pietro Omodeo. Description. Holotype: Preserved length 160 mm, diameter after clitellum 5 mm, segment number 196. Paratypes: 150���190 mm in length, 4���5 mm in diameter. Segment number 153���206, anteclitellar segments frequently multiannulate, segments ii���xi carinate. Colour preserved brownish, alive unknown. Head epilobous 1/2 open, dorsal pores lacking. Setae: ab distant cd paired, setal ratio after clitellum aa:ab:bc:cd:dd = 4:3:3.6:1:12. Nephridial pores begin on segment ii, aligned somewhat dorsal to setal line c. Clitellum annular on segments ��xiii�����xvii. Prostatic pore midventral on a spherical papilla in xiii, near to the 12/13 intersegmental furrow which sometimes emerges as a penis-like organ. Female pores small dots on xiv, close to the intersegmental furrow 14/15 just below setal line d. Spermathecal pore single midventral on xviii (Fig. 2 A���C). Internal characters: Muscular gizzard in vi, large, cylindrical. Septa 7/8���10/11 moderately, 11/12 slightly thickened. Calciferous glands paired in xiv, large mouflon-horn shaped. Small chylus-sacs in ix, x, xi. Dorsal blood vessel simple throughout. Hearts in x���xi, moniliform. Nephridial system holoic with thin biramous nephridial bladders. Typhlosole lacking. Proandric. Testes and sperm funnels in x enclosed in a coiled sperm-reservoir. Seminal vesicles large in xi. Ovaries in xiii enclosed in an ovarian capsule pendant from septum 12/13. This ovarian capsule laterally continuing in a short, hardly recognizable ovarian duct joining the fertilisation chamber and bearing a pendant ovisac. From the fertilisation chamber a straight oviduct leads to the female pore in xiv. The ovarian capsule medially communicates on both sides with a delicate ovo-spermathecal duct running alongside the bursa copulatrix and joining apically to the pear-shaped, highly muscular spermathecal atrium (Fig. 3A). This atrium opens midventrally on segment xviii. The left and right ovarian apparatus connected by a small intercommunicating duct running above the intestine (Fig. 3A). Prostates a pair of long spiral tubes, joining the bursa copulatrix (copulation pouch) via a common Y-shaped duct. Bursa copulatrix situated above the spermathecal atrium and completely covers it dorsally. Bursa copulatrix highly muscular, somewhat spindle-shaped, oriented headwards and opening (through the prostatic pore) on a penislike protuberance on xiii (usually retracted). The two male ducts run backward along both sides of this copulatory pouch, and join to it subapically near each other (Fig. 3B). Penial setae lacking. Discussion. Eudrilidae is the only earthworm family with internal fertilisation (Clausen 1965; Jamieson 1967; Sims 1969), which requires intercommunication of the ovarial and spermathecal systems resulting in a common ovo-spermathecal apparatus. This ovo-spermathecal apparatus is highly variable among different taxa both in its structure and external opening (Sims 1987). The external opening of the spermatheca, usually located near the front of the clitellum, is frequently shifted back: for example, in Beddardiella Michaelsen, 1910 (spermathecal pore at 16/17), Keffia Clausen, 1963 (in xix), or in Parapolytoreutus Segun, 1980 (as far back as segment 22/23). Sometimes it shows wide variation even inside a genus: for example, in Buettneriodrilus aequatorialis Michaelsen, 1935 the spermathecal pore is in 12/13, while in Buettneriodrilus armatus (Michaelsen, 1913) it is in segment xix (Zicsi & Csuzdi 1986; Sims 1987). The position of the prostate pores is less variable; in the genera within the family described to date, it can be found in xvii, 17/18 or in xviii. Petroscolex gen. nov. is the first genus in Eudrilidae with prostatic pore in xiii, anterior to the clitellum. The new genus is also unique in the position of calciferous glands in xiv, not xii or xiii as in other species of the subfamily. Sims (1987) distinguished two groups of genera based on this character, one group with calciferous glands in xii, distributed in West Africa, and another one with calciferous glands in xiii, distributed across tropical Africa. Petroscolex gen. nov. with its calciferous glands in segment xiv is more similar to the latter group because all the genera with backward shifted spermathecal pores possess calciferous glands in xiii. The distribution of the new genus supports this as well, the type locality Irangi, Kivu is at the eastern border of the Congo-basin. A third peculiarity of the new species is the proandric condition of the testes. In Eudrilidae the male apparatus is mostly holandric, although a few genera show metandric reduction: Metascolex Michaelsen, 1903, Polytoreutus Michaelsen, 1890 or Okudrilus Csuzdi & Sherlock, 2015. Petroscolex gen. nov. is the first eudrilid genus with proandric reduction., Published as part of Csuzdi, Csaba, Rota, Emilia, Szederjesi, T��mea, Sherlock, Emma, Brown, George G., Chang, Chih-Han, Cosin, Dario Diaz, Fragoso, Carlos, Jamieson, Barrie G. M., Hong, Yong, James, Samuel W., Paoletti, Maurizio G., Pavl��cek, Tom��s, Plisko, Danuta, Pop, Victor V. & Shen, Huei-Ping, 2019, Description of a new Central African earthworm, Petroscolex centenarius gen et sp. nov. (Crassiclitellata, Eudrilidae), celebrating the 100 th birthday of Pietro Omodeo, pp. 501-508 in Zootaxa 4674 (5) on pages 504-507, DOI: 10.11646/zootaxa.4674.5.1, http://zenodo.org/record/3468756, {"references":["Clausen, M. W. (1965) Description of two new species of Libyodrilus Beddard, 1891, with remarks on the genus and on the female system in the family Eudrilidae. Videnskabelige Meddelelser Dansk Naturhistorisk Forening, 128, 273 - 292.","Jamieson, B. G. M. (1967) A taxonomic review of the African megadrile genus Stuhlmannia (Eudrilidae, Oligochaeta). Journal of Zoology London, 152, 79 - 126. https: // doi. org / 10.1111 / j. 1469 - 7998.1967. tb 01640. x","Sims, R. W. (1969) Internal fertilization in Eudrilid earthworms with the description of a new Pareudriline genus and species (Oligochaeta) from Ghana. Journal of Zoology, 157, 437 - 447. https: // doi. org / 10.1111 / j. 1469 - 7998.1969. tb 01714. x","Sims, R. W. (1987) A review of the Central African earthworm family Eudrilidae (Oligochaeta). In: Bonvicini Pagliai, A. M. & Omodeo, P. (Eds.), On earthworms. Selected symposia and monographs, UZI, 2. Mucchi, Modena, pp. 359 - 388.","Zicsi, A. & Csuzdi, Cs. (1986) Weitere Angaben zur Regenwurmfauna des Kongo-Gebietes (Oligochaeta: Eudrilidae und Glossoscolecidae). Acta Zoologica Hungarica, 32, 385 - 412.","Csuzdi, Cs., Sherlock, E., Talla Kouete, M. & Dohertty-Bone, T. M. (2015) Four new earthworm species from the highlands of Cameroon with description of a new genus Okudrilus gen. n. (Oligochaeta: Eudrilidae & Acanthodrilidae). African Invertebrates, 56, 25 - 38. https: // doi. org / 10.5733 / afin. 056.0103"]}

Petroscolex centenarius Csuzdi, Szederjesi & Sherlock, sp. nov. (Figures 2, 3) Holotype. NHMUK 1997.1594 clitellate adult. DR Congo, Kivu, Irangi. Leg. Gazana N’Doli, 20.02.1973. Paratypes. HNHM AF/3555 1 ex. clitellate adult, NHMUK 1997.1595 1 ex. clitellate adult broken in two pieces. Locality and date same as of the Holotype. NHMUK 1997.1548 – 1549 1 ex. clitellate adult, 1 ex. juvenile, HNHM AF/3539 1 ex. clitellate adult, DR Congo, Kivu, Irangi. Leg. Gazana N’Doli, 08.03.1973. NHMUK 1997.1526 – 1529 1 ex. clitellate adult, 1 ex. aclitellate adult, 2 ex. juvenile, NHMUK ANEA 2019.7368 1 ex. clitellate adult, HNHM AF/5754 1 ex. clitellate adult, DR Congo, Kivu, Irangi. Leg. Gazana N’Doli, 09.03.1973. NHMUK 1997.1530 1 ex. aclitellate adult, DR Congo, Kivu, Irangi. Leg. Gazana N’Doli, 07.06.1973. Etymology. The specific epithet refers to the 100th birthday of Prof. Pietro Omodeo. Description. Holotype: Preserved length 160 mm, diameter after clitellum 5 mm, segment number 196. Paratypes: 150–190 mm in length, 4–5 mm in diameter. Segment number 153–206, anteclitellar segments frequently multiannulate, segments ii–xi carinate. Colour preserved brownish, alive unknown. Head epilobous 1/2 open, dorsal pores lacking. Setae: ab distant cd paired, setal ratio after clitellum aa:ab:bc:cd:dd = 4:3:3.6:1:12. Nephridial pores begin on segment ii, aligned somewhat dorsal to setal line c. Clitellum annular on segments ½xiii–½xvii. Prostatic pore midventral on a spherical papilla in xiii, near to the 12/13 intersegmental furrow which sometimes emerges as a penis-like organ. Female pores small dots on xiv, close to the intersegmental furrow 14/15 just below setal line d. Spermathecal pore single midventral on xviii (Fig. 2 A–C). Internal characters: Muscular gizzard in vi, large, cylindrical. Septa 7/8–10/11 moderately, 11/12 slightly thickened. Calciferous glands paired in xiv, large mouflon-horn shaped. Small chylus-sacs in ix, x, xi. Dorsal blood vessel simple throughout. Hearts in x–xi, moniliform. Nephridial system holoic with thin biramous nephridial bladders. Typhlosole lacking. Proandric. Testes and sperm funnels in x enclosed in a coiled sperm-reservoir. Seminal vesicles large in xi. Ovaries in xiii enclosed in an ovarian capsule pendant from septum 12/13. This ovarian capsule laterally continuing in a short, hardly recognizable ovarian duct joining the fertilisation chamber and bearing a pendant ovisac. From the fertilisation chamber a straight oviduct leads to the female pore in xiv. The ovarian capsule medially communicates on both sides with a delicate ovo-spermathecal duct running alongside the bursa copulatrix and joining apically to the pear-shaped, highly muscular spermathecal atrium (Fig. 3A). This atrium opens midventrally on segment xviii. The left and right ovarian apparatus connected by a small intercommunicating duct running above the intestine (Fig. 3A). Prostates a pair of long spiral tubes, joining the bursa copulatrix (copulation pouch) via a common Y-shaped duct. Bursa copulatrix situated above the spermathecal atrium and completely covers it dorsally. Bursa copulatrix highly muscular, somewhat spindle-shaped, oriented headwards and opening (through the prostatic pore) on a penislike protuberance on xiii (usually retracted). The two male ducts run backward along both sides of this copulatory pouch, and join to it subapically near each other (Fig. 3B). Penial setae lacking. Discussion. Eudrilidae is the only earthworm family with internal fertilisation (Clausen 1965; Jamieson 1967; Sims 1969), which requires intercommunication of the ovarial and spermathecal systems resulting in a common ovo-spermathecal apparatus. This ovo-spermathecal apparatus is highly variable among different taxa both in its structure and external opening (Sims 1987). The external opening of the spermatheca, usually located near the front of the clitellum, is frequently shifted back: for example, in Beddardiella Michaelsen, 1910 (spermathecal pore at 16/17), Keffia Clausen, 1963 (in xix), or in Parapolytoreutus Segun, 1980 (as far back as segment 22/23). Sometimes it shows wide variation even inside a genus: for example, in Buettneriodrilus aequatorialis Michaelsen, 1935 the spermathecal pore is in 12/13, while in Buettneriodrilus armatus (Michaelsen, 1913) it is in segment xix (Zicsi & Csuzdi 1986; Sims 1987). The position of the prostate pores is less variable; in the genera within the family described to date, it can be found in xvii, 17/18 or in xviii. Petroscolex gen. nov. is the first genus in Eudrilidae with prostatic pore in xiii, anterior to the clitellum. The new genus is also unique in the position of calciferous glands in xiv, not xii or xiii as in other species of the subfamily. Sims (1987) distinguished two groups of genera based on this character, one group with calciferous glands in xii, distributed in West Africa, and another one with calciferous glands in xiii, distributed across tropical Africa. Petroscolex gen. nov. with its calciferous glands in segment xiv is more similar to the latter group because all the genera with backward shifted spermathecal pores possess calciferous glands in xiii. The distribution of the new genus supports this as well, the type locality Irangi, Kivu is at the eastern border of the Congo-basin. A third peculiarity of the new species is the proandric condition of the testes. In Eudrilidae the male apparatus is mostly holandric, although a few genera show metandric reduction: Metascolex Michaelsen, 1903, Polytoreutus Michaelsen, 1890 or Okudrilus Csuzdi & Sherlock, 2015. Petroscolex gen. nov. is the first eudrilid genus with proandric reduction.

Petroscolex Csuzdi, Szederjesi & Sherlock, gen. nov. Type species. Petroscolex centenarius sp. nov. Diagnosis. Eudrilinae with single midventral prostatic pore in xiii and single midventral spermathecal pore in xviii. Female pores paired in xiv close to 14/15 near d. Oesophageal gizzard in vi, intestinal gizzards absent. Dorsal blood vessel simple throughout. Paired calciferous glands in xiv and unpaired chylus-sacs in ix, x, xi. Male genital apparatus proandric with sperm reservoirs in x and vesicles in xi. Excretory system holoic, vesiculate. Ovo-spermathecal apparatus paired with a dorsal interconnecting duct. Penial setae lacking. Etymology. The new genus is named in honour of Prof. Pietro Omodeo. Remarks. The new genus is unique among Eudrilidae due to the forward shift of the prostatic pore and the position of calciferous glands in segment xiv.

Petroscolex Csuzdi, Szederjesi & Sherlock, gen. nov. Type species. Petroscolex centenarius sp. nov. Diagnosis. Eudrilinae with single midventral prostatic pore in xiii and single midventral spermathecal pore in xviii. Female pores paired in xiv close to 14/15 near d. Oesophageal gizzard in vi, intestinal gizzards absent. Dorsal blood vessel simple throughout. Paired calciferous glands in xiv and unpaired chylus-sacs in ix, x, xi. Male genital apparatus proandric with sperm reservoirs in x and vesicles in xi. Excretory system holoic, vesiculate. Ovo-spermathecal apparatus paired with a dorsal interconnecting duct. Penial setae lacking. Etymology. The new genus is named in honour of Prof. Pietro Omodeo. Remarks. The new genus is unique among Eudrilidae due to the forward shift of the prostatic pore and the position of calciferous glands in segment xiv., Published as part of Csuzdi, Csaba, Rota, Emilia, Szederjesi, T��mea, Sherlock, Emma, Brown, George G., Chang, Chih-Han, Cosin, Dario Diaz, Fragoso, Carlos, Jamieson, Barrie G. M., Hong, Yong, James, Samuel W., Paoletti, Maurizio G., Pavl��cek, Tom��s, Plisko, Danuta, Pop, Victor V. & Shen, Huei-Ping, 2019, Description of a new Central African earthworm, Petroscolex centenarius gen et sp. nov. (Crassiclitellata, Eudrilidae), celebrating the 100 th birthday of Pietro Omodeo, pp. 501-508 in Zootaxa 4674 (5) on page 504, DOI: 10.11646/zootaxa.4674.5.1, http://zenodo.org/record/3468756

DENDROBAENA GANGLBAUERI (ROSA, 1894) Allolobophora (Dendrobaena) ganglbaueri Rosa, 1894: 1. Dendrobaena ganglbaueri ��� Zicsi 1991: 176. Diagnosis: Length 35���55 mm, diameter 2���3 mm, setae distant. Colour red-violet. First dorsal pore in 7/8 or 8/9. Clitellum on 23, 24���29; tubercles on 25���27. Male pore small on segment 15. Nephridial pores aligned in setal line b. Four pairs of vesicles in 9���12; spermathecae two pairs in 9/10, 10/11 d. Calciferous glands with diverticula in 11. The last pair of hearts in 11, with a pair of small extraesophageal vessels in 12. Nephridial bladders biscuit-shaped. Material examined: HNHM/15595 1 ex., Croatia, Ivan��ćica, ��. Garai, J. Kontsch��n, D. Mur��nyi, 1 April 2006. HNHM/16415 1 ex., Hungary, Szentmargitfalva, Cs. Csuzdi, 10 April 2004. V5011 1 ex. holotype, Slovenia, Črna Prst, L. Ganglbauer. V5012 1 ex., Austria, Schneealpe, R. Sturany., Published as part of Szederjesi, T��mea, Pop, Victor V., Pavl����ek, Tom����, M��rton, Orsolya, Kr��zsik, Vir��g & Csuzdi, Csaba, 2018, Integrated taxonomy reveals multiple species in the Dendrobaena byblica (Rosa, 1893) complex (Oligochaeta: Lumbricidae), pp. 500-516 in Zoological Journal of the Linnean Society 182 on pages 507-508, DOI: 10.1093/zoolinnean/zlx049, http://zenodo.org/record/5714532, {"references":["Rosa D. 1894. Allolobophora Ganglbaueri ed A. Oliveirae nuove specie di Lumbricidi europei. Bollettino dei Musei di Zoologia ed Anatomia Comparata della R. Universita di Torino 9: 1 - 3.","Zicsi A. 1991. Uber die regenwurmer Ungarns (Oligochaeta: Lumbricidae) mit bestimmungstabellen der arten. Opuscula Zoologica Budapest 24: 167 - 191."]}

DENDROBAENA BYBLICA (ROSA, 1893) Allolobophora (Dendrobaena) byblica Rosa, 1893a: 4. Helodrilus (Eophila) thyrrenus ��� Baldasseroni 1907: 48. Eisenia schelkovnikovi ��� Michaelsen 1907: 87. Helodrilus schelkovnikovi veliensis ��� Michaelsen 1910: 46. Helodrilus schelkovnikovi bakuensis ��� Michaelsen 1910: 47. Helodrilus fedtschenkoi lenkoranensis ��� Michaelsen 1910: 49. Helodrilus (Eophila) insularis ��� Chinaglia 1913: 4. Helodrilus (Dendrobaena) lacustris ��� Stephenson 1913: 55. Dendrobaena ganglbaueri var. bulgarica ��� Černosvitov 1937: 84. Eiseniella oltenica ��� Pop 1938: 135. Dendrobaena ganglbaueri var. cylindrica ��� Tzelepe 1943: 36. Dendrobaena ganglbaueri var. differentis ��� Tzelepe 1943: 34. Dendrobaena schelkovnikovi forma graeca ��� Omodeo 1955: 8. Dendrobaena galloprovincialis ��� Bartoli 1962: 438. Dendrobaena hyrcanica ��� Kvavadze & Nikolashvili 1980: 440. Dendrobaena schelkovnikovi herethica ��� Kvavadze 1985: 155. Dendrobaena lacustris saccharophila ��� Qiu & Bouch�� 2000: 156. Diagnosis: Length 20���40 mm, diameter 2.5���3 mm, setae distant. Red-violet pigmentation on the head. First dorsal pore in 10/11. Clitellum on 25���30; tubercles on 26���28. Male pore small on segment 15. Nephridial pores aligned in setal line b. Four pairs of vesicles in 9���12, small and hardly visible in 10; spermathecae two pairs in 9/10, 10/11 d or above. Calciferous glands without distinct diverticula in 10���11. The last pair of hearts in 11, with a pair of small extraesophageal vessels in 12. Nephridial bladders biscuit-shaped. Material examined: HNHM /15808 2 ex., Bulgaria, Strandcha (Bosna) Mts, 210 m, N42 ��09.080��� E27��21.918���, L. D��nyi, Z. Erőss, Z. Feh��r, J. Kontsch��n, D. Mur��nyi, 7 April 2007. HNHM /16160 1 ex., Albania, Korab Mts, 2165 m, N41��48.229��� E20��33.642���, leg. L. D��nyi, Z. Erőss, Z. Feh��r, A. Hunyadi, D. Mur��nyi, 27 June 2007. HNHM /16185 1 ex., Greece, Karpathos, 215 m, N35��39.380��� E27��09.474���, J. Kontsch��n, D. Mur��nyi, 11 November 2012. HNHM /16187 1 ex., Greece, Rhodes, 190 m, N36��20.269��� E28��03.716���, J. Kotsch��n, D. Mur��nyi, 13 November 2012. HNHM /16214 1 ex., Montenegro, Sinjajevina Mts, 1007 m, N42��57.042��� E19��10.410���, L. D��nyi, Z. Feh��r, J. Kontsch��n, D. Mur��nyi, 10 October 2008. HNHM /16353 1 ex., Israel, Har Arbel, T. Pavl��ček, 12 May 2003. HNHM /16355 1 ex., Israel, Wadi Kelt, 31��50���30���N 35��21���39���E, T. Pavl��ček, P. Cardet. HNHM /16457 1 ex., Serbia, Golubac, 78 m, L. D��nyi, J. Kontsch��n, D. Mur��nyi, 12 October 2014. HNHM /16536 1 ex., Turkey, Orontes River, Karşıyaka, 36��05���N 36��03���E, T. Pavl��ček, 14 April 2006. HNHM /16656 1 ex., Greece, Naxos, 455 m, N37��08.580��� E25��31.857���, J. Kontsch��n, D. Mur��nyi, T. Szederjesi, 6 April 2013. HNHM /16661 1 ex., Greece, Crete, 190 m, N35��06.918��� E26��13.153���, J. Kontsch��n, D. Mur��nyi, T. Szederjesi, 4 April 2013. HNHM /16924 1 ex., Turkey, Akyaka Mts, T. Pavl��ček, P. Cardet, 13 April 2014. HNHM /16936 1 ex., Turkey, G��k��eova G��l��, 37.0212��N 28.7410��E, T. Pavl��ček, P. Cardet. HNHM /16942 1 ex., Macedonia, Kožuf Mts, 1670 m, T. Kov��cs, D. Mur��nyi, 4 October 2013. V 5016 5 ex., Crete, Neapolis, A. Penther, 1906., Published as part of Szederjesi, T��mea, Pop, Victor V., Pavl����ek, Tom����, M��rton, Orsolya, Kr��zsik, Vir��g & Csuzdi, Csaba, 2018, Integrated taxonomy reveals multiple species in the Dendrobaena byblica (Rosa, 1893) complex (Oligochaeta: Lumbricidae), pp. 500-516 in Zoological Journal of the Linnean Society 182 on pages 504-506, DOI: 10.1093/zoolinnean/zlx049, http://zenodo.org/record/5714532, {"references":["Rosa D. 1893 a. Viaggio del Dr. E. Festa in Palestina, nel Libano e regioni vicine. - II. Lumbricidi. Bollettino dei Musei di Zoologia ed Anatomia Comparata della R. Universita di Torino 8: 1 - 14.","Baldasseroni V. 1907. Contributo alla conoscenza dei Lombrichi italiani. Monitore Zoologico Italiano 17: 48 - 60.","Michaelsen W. 1907. Die Lumbriciden des Kaukasischen Museums in Tiflis. Mittellungen des Kaukasischen Museums 3: 81 - 93.","Michaelsen W. 1910. Zur kenntnis der lumbriciden und ihrer verbreitung. Annuaire du Musee Zoologique de l'Academie Imperiale des Sciences de St. - Petersbourg 15: 1 - 74.","Chinaglia L. 1913. Escursioni Zoologiche in Sardegna del. Dr. E. Festa III. Lombrichi. Bollettino dei Musei di Zoologia ed Anatomia Comparata della R. Universita di Torino 28: 1 - 6.","Stephenson J. 1913. Aquatic Oligochaeta from the Lake of Tiberias. Journal and Proceedings of the Asiatic Society of Bengal 9: 53 - 56.","Cernosvitov L. 1937. Die Oligochaetenfauna Bulgariens. Mitteilungen aus den Koniglich Naturwissenschaftlichen Instituten in Sofia 10: 62 - 92.","Pop V. 1938. Neue lumbriciden aus rumanien. Buletinul Societatii de Siinte din Cluj 9: 134 - 152.","Tzelepe ND. 1943. Symbole eis ten melethen ton Oligochaeton tes Hellados. University Dissertation, Athens: University of Athens.","Omodeo P. 1955. Lombrichi cavernicoli di Grecia e Turchia raccolti dal Dr. K. Lindberg. Annuario dell'Istituto e Museo di Zoologia della Universita di Napoli 7: 1 - 16.","Bartoli P. 1962. Une Nouvelle Espece de Lombricidae en Provence: Dendrobaena galloprovincialis n. sp. Bulletin de la Societe Zoologique de France 87: 438 - 442.","Kvavadze E, Nikolashvili K. 1980. On the distinctness of the species Dendrobaena schelkovnikovi (Michaelsen, 1907) and a new earthworm species from the Hirkan National Park. Izvestiya Akademii Nauk Gruzii Seriya Biologicheskaya 6: 440 - 444.","Kvavadze E. 1985. The earthworms (Lumbricidae) of the Caucasus. Tbilisi: Metsniereba [in Russian]."]}

DENDROBAENA SP. Dendrobaena byblica byblica: Szederjesi et al. 2014b: 558. Description External characters: Both specimens cut in half. Length ~ 42 mm, diameter 2.5 mm. Number of segments ~105. Pigmentation pinkish on the head. Prostomium epilobous ½ opened. First dorsal pore at intersegmental furrow 14/15. Setae distantly standing. Setal arrangement behind clitellum: aa: ab: bc: cd: dd = 1.75:1.25:1.13:1:2.5. Male pores minute on segment 15. Nephridial pores irregularly alternate between setal line b and above d. Clitellum on segments 25–30. Tubercula pubertatis on 1/ n 25, 26 –½29. Glandular tumescence on segments 9 ad, 10, 11 abcd. Internal characters: No dissepiments notably thickened. Testes and funnels paired in segments 10–11, covered by perioesophageal testis sac in segment 10 and 11. Three pairs of seminal vesicles in 9, 11 and 12. Two pairs of spermathecae in 9/10, 10/11 with external openings in setal line d. Calciferous glands in segment 11 and 12, without distinct diverticula. The last pair of hearts in 11, with a pair of extraesophageal vessels in 12. Nephridial bladders are biscuit-shaped. Crop in segments 15–16, and gizzard in segments 17–18. Typhlosolis simple lamelliform, large. The cross section of the longitudinal muscle layer of pinnate type. Material examined: HNHM /15799 1 ex., HNHM /15800 1 ex., Turkey, Istranca Mts, 253 m, N41°49.300′ E27°45.085′, L. Dányi, Z. Erőss, Z. Fehér, J. Kontschán, D. Murányi, 6 April 2007. HNHM /15813 1 ex., Turkey, Istranca Mts, 538 m, N41°44.667′ E27°39.279′, L. Dányi, Z. Erőss, Z. Fehér, J. Kontschán, D. Murányi, 6 April 2007. Remarks: The only characters that distinguish this questionable species from D. byblica are the number of vesicles (3 vs. 4) and the somewhat longer tubercula (1/ n 25, 26 –½29 vs. 26–28). To reveal its exact identity, revision of all D. byblica synonyms and similar species is necessary. However, we can’t exclude that these specimens represent a new species. DENDROBAENA CF. DAGHESTANENSIS (MICHAELSEN, 1907) Helodrilus (Dendrobaena) ganglbaueri var. daghestanensis: Michaelsen 1907: 91. Dendrobaena byblica – Pop 1949: 483 [in part].

DENDROBAENA SP. Dendrobaena byblica byblica: Szederjesi et al. 2014b: 558. Description External characters: Both specimens cut in half. Length ~ 42 mm, diameter 2.5 mm. Number of segments ~105. Pigmentation pinkish on the head. Prostomium epilobous �� opened. First dorsal pore at intersegmental furrow 14/15. Setae distantly standing. Setal arrangement behind clitellum: aa: ab: bc: cd: dd = 1.75:1.25:1.13:1:2.5. Male pores minute on segment 15. Nephridial pores irregularly alternate between setal line b and above d. Clitellum on segments 25���30. Tubercula pubertatis on 1/ n 25, 26 �����29. Glandular tumescence on segments 9 ad, 10, 11 abcd. Internal characters: No dissepiments notably thickened. Testes and funnels paired in segments 10���11, covered by perioesophageal testis sac in segment 10 and 11. Three pairs of seminal vesicles in 9, 11 and 12. Two pairs of spermathecae in 9/10, 10/11 with external openings in setal line d. Calciferous glands in segment 11 and 12, without distinct diverticula. The last pair of hearts in 11, with a pair of extraesophageal vessels in 12. Nephridial bladders are biscuit-shaped. Crop in segments 15���16, and gizzard in segments 17���18. Typhlosolis simple lamelliform, large. The cross section of the longitudinal muscle layer of pinnate type. Material examined: HNHM /15799 1 ex., HNHM /15800 1 ex., Turkey, Istranca Mts, 253 m, N41��49.300��� E27��45.085���, L. D��nyi, Z. Erőss, Z. Feh��r, J. Kontsch��n, D. Mur��nyi, 6 April 2007. HNHM /15813 1 ex., Turkey, Istranca Mts, 538 m, N41��44.667��� E27��39.279���, L. D��nyi, Z. Erőss, Z. Feh��r, J. Kontsch��n, D. Mur��nyi, 6 April 2007. Remarks: The only characters that distinguish this questionable species from D. byblica are the number of vesicles (3 vs. 4) and the somewhat longer tubercula (1/ n 25, 26 �����29 vs. 26���28). To reveal its exact identity, revision of all D. byblica synonyms and similar species is necessary. However, we can���t exclude that these specimens represent a new species. DENDROBAENA CF. DAGHESTANENSIS (MICHAELSEN, 1907) Helodrilus (Dendrobaena) ganglbaueri var. daghestanensis: Michaelsen 1907: 91. Dendrobaena byblica ��� Pop 1949: 483 [in part]., Published as part of Szederjesi, T��mea, Pop, Victor V., Pavl����ek, Tom����, M��rton, Orsolya, Kr��zsik, Vir��g & Csuzdi, Csaba, 2018, Integrated taxonomy reveals multiple species in the Dendrobaena byblica (Rosa, 1893) complex (Oligochaeta: Lumbricidae), pp. 500-516 in Zoological Journal of the Linnean Society 182 on page 507, DOI: 10.1093/zoolinnean/zlx049, http://zenodo.org/record/5714532, {"references":["Szederjesi T, Pavlicek T, Coskun Y, Csuzdi CS. 2014 b. New earthworm records from Turkey, with description of three new species (Oligochaeta: Lumbricidae). Zootaxa 3764: 555 - 570.","Michaelsen W. 1907. Die Lumbriciden des Kaukasischen Museums in Tiflis. Mittellungen des Kaukasischen Museums 3: 81 - 93.","Pop V. 1949. Lumbricidele din Romania. Analele Academiei Republicii Populare Romane Sectiunea de Stiinte Geologice, Geografice si Biologie 1: 383 - 505."]}

DENDROBAENA LOEBLI (ZICSI, 1985) COMB. NOV. Fitzingeria loebli Zicsi, 1985: 330. ��� Szederjesi & Csuzdi 2012b: 37. Diagnosis: Length 32���60 mm, diameter 2���4 mm, setae distant. Slight pinkish pigmentation on the head. Dorsal pores not visible. Clitellum on 24�����29, 29; tubercles 25���27, ��28. Male pore on segment 25 or 26. Nephridial pores irregulated. Three pairs of vesicles in 9, 11, 12; two pairs of spermathecae in 9/10, 10/11 d. Calciferous glands in segment 11���12, without distinct diverticula. The last pair of hearts in 11, with a pair of small extraesophageal vessels in 12. Nephridial bladders biscuit-shaped. Material examined: HNHM/7505 1 ex. paratype, Turkey, Kazdag (Mt. Ida), H. Schweiger, 14 April 1965. HNHM/15757 3 ex., Greece, Sapka Mts, 651 m, N41��07.672��� E25��53.223���, L. D��nyi, Z. Erőss, Z. Feh��r, J. Kontsch��n, D. Mur��nyi, 4 April 2007., Published as part of Szederjesi, T��mea, Pop, Victor V., Pavl����ek, Tom����, M��rton, Orsolya, Kr��zsik, Vir��g & Csuzdi, Csaba, 2018, Integrated taxonomy reveals multiple species in the Dendrobaena byblica (Rosa, 1893) complex (Oligochaeta: Lumbricidae), pp. 500-516 in Zoological Journal of the Linnean Society 182 on page 508, DOI: 10.1093/zoolinnean/zlx049, http://zenodo.org/record/5714532, {"references":["Zicsi A. 1985. Regenwurmer (Oligochaeta: Lumbricidae) aus Israel und den benachbarten Landern. Revue Suisse de Zoologie 92: 323 - 331.","Szederjesi T, Csuzdi CS. 2012 b. New and little known earthworm species from Greece (Oligochaeta: Lumbricidae, Acanthodrilidae). Zootaxa 3304: 25 - 42."]}

DENDROBAENA ANNECTENS (ROSA, 1895) COMB. NOV. Allolobophora ganglbaueri var. annectens Rosa, 1895: 7. Fitzingeria annectens ��� Zicsi & Pop 1984: 246. Description External characters: Length 32���55 mm, diameter 3 ��� 4.5 mm. Number of segments 101 ��� 116. Pigmentation dark red-violet. Prostomium epilobous �� closed. Dorsal pores not visible. Setae distantly standing. Setal arrangement behind clitellum: aa: ab: bc: cd: dd = 1.39:1.22:1.67:1:1.44. Male pores minute on one of the segments 22���27. Nephridial pores aligned above setal line b. Clitellum on segments 25�����30, 30. Tubercula pubertatis on 26���28. Glandular tumescence on segments 9 b, 10, 11 c. 12 b. Internal characters: No dissepiments notably thickened. Testes and funnels paired in segments 10���11, covered by oesophageal testis sac in segment 10 and 11. Four pairs of seminal vesicles in 9���12, 10 and 11 small. Two pairs of spermathecae in 9/10, 10/11 with external openings in setal line d. Calciferous glands with well-developed diverticula in segment 11. The last pair of hearts in 11, with a pair of extraesophageal vessels in 12. Nephridial bladders are biscuit-shaped. Crop in segments 15���16, and gizzard in segments 17���18. Typhlosolis simple, large. The cross section of the longitudinal muscle layer of pinnate type. Material examined: HNHM /369 3 ex., Romania, Piatra Craiului, A. Zicsi, 5 August 1961. HNHM /7913 2 ex., Romania, Piatra Mare, A. Zicsi, 27 August 1974. HNHM /16230 1 ex., Romania, D��mbova, 458 m, N44��56.995��� E23��11.667���, Cs. Csuzdi, J. Kontsch��n, V. V. Pop, 30 October 2007. HNHM /16238 1 ex., Romania, Cerna Sat, 486 m, N45��05.396��� E22��38.134���, Cs. Csuzdi, J. Kontsch��n, V. V. Pop, 29 October 2007. HNHM /16249 1 ex., Romania, Cerna Sat, 436 m, N45��04.646��� E22��37.368���, Cs. Csuzdi, J. Kontsch��n, V. V. Pop, 29 October 2007. HNHM /16251 1 ex., Romania, Cerna Sat, 496 m, N45��07.390��� E22��40.541���, Cs. Csuzdi, J. Kontsch��n, V. V. Pop, 29 October 2007. HNHM /16558 1 ex., Romania, Băile Herculane, Cs. Csuzdi, 28 April 2004. V 5018 4 ex., V 5015 5 ex., Romania, Transylvania, L. Ganglbauer. Remarks: Dendrobaena annectens was first described as a variety of ganglbaueri until Pop (1949) synonymized both with D. byblica. Zicsi & Pop (1984) examining extensive material from the Southern Carpathians and a syntype housed in Michaelsen���s Collection (Hamburg) recognized that the male pores of this species are shifted backward to the beginning of the clitellum, therefore they removed it from the synonymy of D. byblica and placed it in Fiztingeria. However, Fitzingeria has proven to be polyphyletic. Therefore, we here place all previous Fitzingeria species including annectens in Dendrobaena., Published as part of Szederjesi, T��mea, Pop, Victor V., Pavl����ek, Tom����, M��rton, Orsolya, Kr��zsik, Vir��g & Csuzdi, Csaba, 2018, Integrated taxonomy reveals multiple species in the Dendrobaena byblica (Rosa, 1893) complex (Oligochaeta: Lumbricidae), pp. 500-516 in Zoological Journal of the Linnean Society 182 on page 508, DOI: 10.1093/zoolinnean/zlx049, http://zenodo.org/record/5714532, {"references":["Rosa D. 1895. Nuovi lombrichi dell'Europa orientale. Bollettino dei Musei di Zoologia ed Anatomia Comparata della R. Universita di Torino 10: 1 - 8.","Zicsi A, Pop VV. 1984. Neue regenwurmer aus Rumanien (Oligochaeta: Lumbricidae). Acta Zoologica Academiae Scientiarum Hungaricae 30: 241 - 248.","Pop V. 1949. Lumbricidele din Romania. Analele Academiei Republicii Populare Romane Sectiunea de Stiinte Geologice, Geografice si Biologie 1: 383 - 505."]}

There is ample evidence demonstrating the importance of maternal thyroid hormones, assessed at single trimesters in pregnancy, for child cognition. Less is known, however, about the course of maternal thyroid hormone concentrations during pregnancy in relation to child behavioral development. Child sex might be an important moderator, because there are sex differences in externalizing and internalizing behavioral problems. The current study examined the associations between maternal thyroid hormone trajectories versus thyroid assessments at separate trimesters of pregnancy and child behavioral problems, as well as sex differences in these associations. In 442 pregnant mothers, serum levels of TSH and free T4 (fT4) were measured at 12, 24, and 36 weeks gestation. Both mothers and fathers reported on their children's behavioral problems, between 23 and 60 months of age. Latent growth mixture modeling was used to determine the number of different thyroid hormone trajectories. Three trajectory groups were discerned: 1) highest and non-increasing TSH with lowest fT4 that decreased least of the three trajectories; 2) increasing TSH and decreasing fT4 at intermediate levels; 3) lowest and increasing TSH with highest and decreasing fT4. Children of mothers with the most flattened thyroid hormone trajectories (trajectory 1) showed the most anxiety/depression symptoms. The following trimester-specific associations were found: 1) lower first-trimester fT4 was associated with more child anxiety/depression, 2) higher first-trimester TSH levels were related to more attention problems in boys only. A flattened course of maternal thyroid hormone concentrations during pregnancy was a better predictor of child anxiety/depression than first-trimester fT4 levels.Keywords: Thyroid, Longitudinal, Pregnancy, Behavioral problems, Children,Sex difference