Phaenocora unipunctata ( Ørsted, 1843 ) Bendl, 1908

Houben, Albrecht M., Steenkiste, Niels Van & Artois, Tom J., 2014, Revision of Phaenocora Ehrenberg, 1836 (Rhabditophora, Typhloplanidae, Phaenocorinae) with the description of two new species, Zootaxa 3889 (3), pp. 301-354 : 343-344

Phaenocora unipunctata ( Ørsted, 1843) Bendl, 1908

( Fig. 7 )

Derostoma unipunctatum ØRSTED 1843: 560; Ørsted 1844: 66, plate 2 Fig. 25; Schmidt 1848: 36 –38, plate 2 Figs 5, 5 a, 5b; Schultze 1851: 44; Vejdovský 1880: 503; Graff 1882: 367 –368; Braun 1885: 221 –226, plate 4 Figs 21–22; Lippitsch 1889: 147 –167, TextFig. p 156, plate 8 Figs 1 –17; Fuhrmann 1894: 217, 220–221, 274–280; Vejdovský 1895: 113 –114, 116, 118, 122, 124–128, plate 5 Figs 34–36; Fuhrmann 1900: 730; Dorner 1902: 493; Sekera 1904: 441; Brinkmann 1906: 134 –136 table 5 Figs 22–23.

Derostoma schmidtianum Schultze 1851: 44 –46, 50–52, plate 4 Fig. 6–9, Braun 1885: 221; Vejdovský 1895: 124.

Derostomum schmidtianum Schmidt 1858: 26 .

Phaenocora unipunctata Bendl 1908 , 129–130; Bendl 1909: 294, 297–298, Fig. 2; Graff 1909: 93, Fig. 189; Wahl 1910: 42 –44 plate 2 Fig. 1 ; Hofsten 1911: 4, 32–33; Hofsten 1912: 553, 559, 582, 624, 625; Graff 1913: 134 –137, 138 Figs 136a, 136b; Wilhelmi 1913: Fig. 67; Böhmig 1914: 91; Meixner 1915: 535, 537–538, 541; Hofsten 1918: 35, TextFig. 8C, plate XXVIII Fig. 23, 24; Luther 1918: 49; Nasonov 1919: 620, 621, 633, 1192; Beklemischev 1921: 641 –643, plate 2 Figs 12, 13; Luther 1921: 9, 11, 25; Reisinger 1923: 9, 26, 57–58, Figs 28, 32; Reisinger 1924: 251; Nasonov 1926: 869 –870; Beklemischev 1927: 205; Beklemischev 1929: 534, 538; Sekera 1930: 100; Stummer-Traunfels & Meixner 1930 (and references therein): 3410, 3456, 3460; Steinböck 1932: 210, 212; Gilbert 1935: 284, 286, 294–295, 302, 315, 318, 340–345, 348, 352–353, 369, 377, TextFigs 1C, 3Aa, 3Ab, tables 1, 2; Beauchamp 1936: 149 –151; Gilbert 1938a: 212, 214; Weise 1942: Fig. 22 Marcus 1946: 66 –68, 70, 75; Steinböck 1948: 22 –23; Steinböck 1949: 232, 242, 250–251; Rixen 1961: 500, Fig. 45; Luther 1963: 127 –130, Fig. 42, plate I Fig. L; Steinböck 1966: 131, 175; Young 1970: 221 –222 Fig. 4A, table 1; Young 1973a: 208 –209, 212, 219 tables 1, 5; Young & Harris 1973: 85 –87; Mack-Fira 1974: 249, 262; Eaton & Young 1975: 66; Heitkamp 1981: 30, table 1; Heitkamp 1982: 141 –142; Müller & Faubel 1993: 379, 387, tables 1, 2; Noreña-Janssen 1995: 243–244, Fig. 16 ; Gamo & Noreña-Janssen 1998: table 1; Noreña et al. 1999: table 1; Korgina 2002: table 1; Noreña et al. 2004: 234; Willems et al. 2006: 3, 8, 11; Noreña et al. 2007: 1998, table 2, 3; Noreña et al. 2008: 10 , Fig. 8 .

Phoenocora unipunctata (incorrect subsequent spelling) Cognetti de Martiis 1916: 193–194, 198–199, 201, 207, 209–212, 217–219, 224–225, 228–229, 233, 235–236.

Derostoma unipunctata Hofsten 1912: 624 ; Luther 1921: 9.

Phaenocora unipunctata (?) Nasonov 1919: 622.

Phaenocora unipunctata acaudata Weise 1942: 145 , 165, Fig. 21.

Known distribution: Widely distributed: many localities in Europe (also see Luther 1963 for localities and references): Münchenstein and Reinach ( Switzerland) ( Fuhrmann 1894); Bulychevo ( Russia) ( Nasonov 1919); United Kingdom and Ireland ( Young 1970); Hatch Mere, Cheshire ( United Kingdom) ( Young 1973a); Romania (see Mack-Fira 1974 for localities and references); Seeburger lake, Seeburg ( Germany) ( Heitkamp 1981); environment of Göttingen ( Germany) ( Heitkamp 1982); in the vicinity of Hamburg ( Germany), along the Elbe estuary (see Müller & Faubel 1993 for localities and references); Along the Zapatón river and the Salor river ( Spain) ( Noreña et al. 1999); upper Volga river basin ( Russia) ( Korgina 2002); Diepenbeek ( Belgium), De Maten ( Willems et al. 2006), Ason River near Santa Ulalla Sanctuary (Cantabria, Spain), 43°20’N; 03°26’W; Artificial spring in Ramales de la Victoria (Cantabria, Spain), 43°16’N; 03°27’W ( Noreña et al. 2007); Djursholm, trench at Ekebysjön ( Sweden) (SMNH no. 93018); Tingvallavatn ( Iceland) (SMNH nos 92993–92998, 96991, 96995).

Asia: Eastern Russia, Indonesia and the Tien-Shan area ( China / Kyrgyz Republic border) at about 3500 m altitude (see Luther 1963 for localities and references); Keren Carmel ( Israel); Temporary, artificial pool in Hai Bar Carmel Nature Reserve ( Israel); Lahav, northern Negev ( Israel) ( Noreña et al. 2008).

South America: Cordoba, ( Argentina), series of ponds along from Marina to Angelica; Rio Negro ( Argentina), small pond near to lake Nahuel Huapi (Noreña-Janssen 1995).

New localities: Vijlen ( The Netherlands) Sediment collected on 28 March 2004, inundated in the laboratory in June 2004.

Material examined: Nine serially-sectioned specimens from Vijlen (six sagitally-, two horizontally- and one transversely-sectioned specimens). One sagitally-sectioned specimen designated neotype (sections of one specimen located on two slides: SMNH Type-8674a and Type-8674b), the others reference material (sections of eight specimens located on sixteen slides: HU nos VI.3.43–VI.3.50; VI.4.01–VI.4.08). Eight transverselysectioned specimens from Tingvallavatn, Iceland ( SMNH nos 92993–92998, 96991, 96995), one whole mount from Djursholm, trench at Ekebysjön, Sweden ( SMNH no. 93018).

Diagnosis: Animals up to 3 mm long, with a small tail. Eyes black, yellowish or reddish up to red, clearly present or only weakly visible. Pigmentation absent or sometimes present as small spots of brown pigment. Zoochlorellae often present. The male copulatory organ is of the duplex-type IIIA. The female genital system is of the UNIPUNCTATA - type. A small intestinal bursa is connected directly to the gut, a long genito-bursal duct and a short female genital canal are present. The oviduct opens into the junction of the genito-bursal duct and the female genital canal.

Descriptive notes: The studied animals are about 1.5 mm long (measured on sagitally-sectioned specimens). The testes are somewhat lobular and are situated laterally. The vitellaria are situated at the ventral side.

The male copulatory organ ( Fig. 7 A) is of the duplex-type IIIA. A clear sphincter is present between the seminal vesicle and the prostate vesicle ( Fig. 7 A: sph). The unarmed penis papilla is lined with a thin epithelium.

The female genital system ( Fig. 7 B) is connected to the gut by a simple valvular apparatus ( Fig. 7 B: va), which can be closed by a weak sphincter. From this point muscles ( Fig. 7 B: mu) run toward the epithelial lining of the pharynx ( Fig. 7 B: ph) and (probably) towards the ventral epidermis (although this is not clearly observed). The valvular apparatus is directly attached to a small intestinal bursa ( Fig. 7 B: bi), which is only slightly wider than the rather long burso-intestinal duct ( Fig. 7 B: dbi). Both are often filled with sperm and lined with an irregular, nucleated epithelium. They are surrounded by muscles, which distally become stronger. The ovo-vitelloduct (od) opens between the burso-intestinal duct and the female genital canal ( Fig. 7 B: fgc). The female genital canal is rather short and it receives the shell glands ( Fig. 7 B: sg).

Remarks: According to Reisinger (1923), the pigmentation of the eye spots varied considerably among individuals of the same population, ranging from completely absent to strongly coloured. Moreover, he ( Reisinger 1923) kept animals in dark conditions and observed a diminishing eye pigmentation over generations. In addition, the presence and quantity of zoochlorellae can also differ among individuals of the same population; for instance, some populations of P. unipunctata found in Britain ( Young & Harris 1973) entirely lacked zoochlorellae.

The whole mount present in the collections of the Swedish Museum of Natural History (SMNH no. 93018) does not show any internal structure, hence we cannot confirm whether it is indeed this species. This specimen has eaten Microdalyellia rossi ( Graff, 1911) , the stylet of which can clearly be observed. Slides SMNH nos 96991 and 96995 are both sampled at the same date (23/7/1947) at Tingvallavatn ( Iceland). Slide SMNH no. 96991 is probably P. unipunctata , although this is difficult to assess because it is a transversely-sectioned specimen and the male copulatory organ is evaginated, both hampering the assessment of the exact type of copulatory organ and hence species identification. The specimen on slide SMNH no. 96995 shows no reproductive organs at all, but most probably it is the same species as on slide no. 96991, since both specimens were sampled at the same date and location (Tingvallavatn, Iceland; 23 June 1947).

Figures

FIGURE 1. Photographs of live animals A, P. evelinae. B, P. foliacea from South Africa. C, P. foliacea from India. D, P. aglobulata sp. nov.. E – H, P. gilberti sp. nov.: E – F, pigmentation of the head. G, male copulatory organ. H, female genital canal.

FIGURE 5. Phaenocora evelinae A, habitus of live animal with an invaginated male copulatory organ. B, invaginated male copulatory organ as seen on live animal. C, partly evaginated male copulatory organ as seen on live animal. D, evaginated male copulatory organ as seen on live animal. E, reconstruction on sagittal sections of male copulatory organ (caudal end at the left) F, reconstruction on sagittal sections of female genital system (caudal end at the left). (E – F: HU no. VI. 3.25).

FIGURE 7. Phaenocora unipunctata A, reconstruction on sagittal sections of male copulatory organ (caudal end at the left). B, reconstruction on sagittal sections of female genital system, original position of the ovary is marked with ... + ..... + ... (caudal end at the left). (SMNH Type- 8674).

FIGURE 8. Phaenocora variodentata A, reconstruction on sagittal sections of female genital system (caudal end at the right). B, photograph of the copulatory system. C, three photographs of different levels of the three dorsal spines. D, photograph of the cirrus with different sized spines. All on material from China (" P. sinensis ": A. SMNH Type- 8564 [SU-C, no. SZ 200903 - I- 3]; B – D. SMNH Type- 8562 [SU-C, no. SZ 200903 - I- 1]).

FIGURE 16. Reproduction of female genital system of various species. A, Phaenocora typhlops (after Luther 1921). B, P. lutheri (after Gilbert 1937). C, P. bresslaui (after Marcus 1946). D, P. chloroxantha (after Marcus 1946). E, P. highlandensis (after Gilbert 1935). F, P. polycirra (after Beklemischev 1929). G, P. rufodorsata (after Hofsten 1911).