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M. hilgendorfi /A. tokioensis

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1 M. hilgendorfi /A. tokioensis
[Eng.] A more detailed description of samping site is shown in AmyMetaKouen.MPP. OIG: Ohmuta Industrial region Grassy place, [Oh/9] 9MR: Meadow along road, [Oh/1, Ta/2, T/3, I/1, O/10] 17FP: Forest park, [Oh/9, Kkk/1 ] 10MI: Meadow in improved land, (Several were organic poor soil) , [D/4, C/1, ] 5 SSF: Small shrine forest, (many were organic poor soil) [Oh/5, T/3, W/1, I/1, C/1, B/2, A/2, U/1] 19MDA: Meadow where dicotyledonous plant is abundant, (many were organic rich soil) [D/32] 32SNU: Shrine forest near urban area (Ever greem forest), [Ft/2, K/4, D/2] 8ENS: Shrine forest Ever green forest near sea, [Ma/1, Fh/1, M/10, O/6] 18 Low: Low mountain (Independent peak of altitude 500m or less) (Ever green forest). [Fh/1, T/4, Kkk/2. Kkm/3, Kyn/7] 17 Sch: Schrine forest (Ever green forest), [Fh/5, Mu/4, H/3] 11San Sangun mountainous district, Mt. Sangun (Alt. 936m) – Mt. Homan (Alt. 829m) (Ever green forest) [D-C/ 26, S/5] 31Kos: Mt. Kosyo (Alt. 900m) (Ever green forest), [A/5] 5Sef: Sefuri mountainous district Mt. Sefri (Alt m) (Ever green-summer green forest), [ C/2, Fs/2, N/1, Saga Pref.Tos/1] 6Sya; Mt. Syaka (Alt. 1231m) (Ever green-summer green forest) [Y/2] 2Hik: Mt Hikosan (Alt. 1200m) (Ever green-summer green forest, [S/2] 2 Additional sampling site Kum; Kumamoto Pref. Kyushu mountainous district (Mt. Kunimi Alt. 1739m (Ever-summer) 3 [JPN.]調査/採集地点のより詳しい記述はAmyMetaKouen.PPMに示している。

2 Amynthas tokioensis [Eng.] Diagnosis: Two pairs of spermathecal pores ca body circumference apart in furrows 6/7/8. One or more small genital markings paired just in front of the setal arcs anteriorly on segment 7 and/or 8, all markings with glands internally. Male pores are superficial. Male pore ca circumference apart with 7-16 setae between male pores. One genital marking present on setal line near male pore. Diverticulum is equal (Kubuki and Syaka) or shorter (Ohmuta) to Ampulla in length. Prostatic gonad is large occupying segments separating two or more plates, and Prostatic duct is U (Ohmuta and Syaka) or loop (Kubuki) shape. The duct is slender in early and is stout in later. There is compact glandular mass just before the end of Duct. Remark:  Many features shown in above are same to that of Amynthas tokioensis shown in Fig.1 and Fig.2 of "Review of Japanese earthworms (Annelida: Oligochaeta) after Blakemore (2003) [JPN.] Fig. 1 Amynthas tokioensis (Beddard, 1892). Ventral view of spermathecal pore. B. Ventral view of male pore. C. Spermathecal pore. (The Ampulla in 6/7 has come off from the duct.). D. Prostatic gonad and duct. E. Caeca

3 Amynthas vittatus [Eng.]
Remark: A. tokioenssis and Amynthas vittatus can be differenciated only by the differences of the number, and position of genital papillae on segments 7 , 8 and 18. Blakemore (2005) described Banded colouration due to pale seta lines, of Amynthas vittatus. Colour banded colouration is not clear in many individuals in present study. Blakemore (2005) listed A. vittatus in synonymous with A. tokioensis [JPN.] Fig. 2 Amynthas vittatus (Goto & Hatai 1898) Ventral view of spermathecal pore. (A’. Twice close-up of right spermathecal pore in 7/8th segment) B. Ventral view of male pore. (B’. Twice close-up of right male pore). C. Spermathecal pore. D. Prostatic gonad and duct. E. Caeca

4 Amynthas tokioensis v. Akizuki
[Eng.] There are two pairs of genital markings of pre setal arc of body surface of segment 8. There are several glands near the spermathecae and mass glands near mid ventral wall of segment 8. There is two pair of genital marking in pre and another two pair post, of the setal arcs of segment 18 (Four pairs in total). Remark: The feature of this specimen is in “mass glands on the center of ventral wall of segment 8” and “Two pairs of genital markings in pre- and post of the setal arcs on segmemt 18”. The arrangement of genital marking of this taxa, partially, looks like that of Pheretima verticosa (Ishizuka1999). Blakemore 2005 considers Pheretima verticosa (Ishizuka 1999) to be synonym of Amynthas tokioensis (Beddard, 1892) [JPN.] Fig. 3 Amynthas tokioensis v. Akizuki : A. Ventral view of spermathecal pore. B. Ventral view of male pore. C. Spermathecal pore. D. Prostatic gonad and duct. E. caeca.

5 Amynthas tokioensis v. Hakozaki
[Eng.] Male pore of A. tokioensis v. Hakozaki is characterized by large disc and two genital papilae that touch it. These features can be clearly distinguished from Amynthas tokioensis. However, other features look like well with each other. There is a small genital marking mid ventral in pre-setal line on more likely segments 7 and/or 8, of some individuals. There is no internal gland corresponding to external genital marking. Male pores are large disc (superficial). There must be two genital papillae (penial setae?) along the circumference of large disc. Male pores ca circumference apart with 8-10 setae between male pores. Prostatic gonad greatly extends over  The distribution of Amynthas tokioensis v. Hakozaki has been almost limited to old grass field and the forest having a rich ground flora in the plains part of Fukuoka City, Kasuga City, Dazaifu City, and Chikushino City, Kyushu Japan. A. tokioensis v. Hakozaki is distributed around the distribution region of A. tokioensis and A. vittatus. It is thought that A. tokioensis v. Hakozaki is variety or ecotype of Amynthas tokioensis. Etymology. Named after the locality [JPN.] Fig. 4 Amynthas tokioensis v. Hakozaki. A. Ventral view of spermathecal pore. B. Ventral view of male pore. C. spermathecal pore. D. Prostatic gonad and duct. E. Caeca.

6 Amynthas tokioensis v. Munakata
Table 1 Amynthas tokioensis v. Munakata [Eng.] Male pores are large disc (superficial).  But, the disc is slightly smaller than that of A. tokioensis v. Hakozaki.   There are one or two genital papillae (penial setae?) along the circumference of large disc.   Besides, there are one or two papillae on setal line of male segment.  Remark: Male disc of A. tokioensis v. Munakara is a little smaller than that of A. tokioensis v. Hakozaki. The number of genital papillae which touches disc is always 2 in A. tokioensis v. Hakozaki, and the range of 1-3 in A. tokioensis v. Munakara and there are one or two genital papillae on setal line of male segment in earlier taxa but no genital papillae in that areas of later taxa. The diagnosis on genital marking of v. Munakata is the intermediate phase of A. tokioensis and A. tokioensis v. Hakozaki. Those differences are very small. Other features look like well for “true” Amynthas tokioensis. Therefore, A. tokioensis v. Munakata is thought to be variety or ecotype of Amynthas tokioensis besides A. tokioensis v. Hakozaki. Distribution:  The distribution region of A. tokioensis v. Munakata is evergreen forest of Munakata City which is located east of Fukuoka City outskirts. Etymology. Named after the locality. [JPN.] Fig. 5 Amynthas tokioensis v. Munakara : A. external view of spermathecal pore. B. external view of male pore. C. Spermathecal pore. D. Prostatic gonad and duct. E. caeca

7 Amynthas tokioensis v. Homan
[Eng.] Three pairs of spermathecal pores ca body circumference apart in furrows 5/6/7/8. Diverticulum is equal to Ampulla in length. Genital marking in pre seta line of ventral site of segment 7, and in furrow of segment 7/8 near spermathecal pore, and three pairs near male pore region. There are mass of capsulogenous glands near the spermathecae duct of segments 7 and 8. Male pores are superficial. Genital marking on male segment 18 are two pairs pre setal line and one pair post setal line. Remark: Amynthas tokioensis v. human shows superficial male pore with plostatic gonad and duct and genital marking, on 6-9 segments, each of which looks alike well for those of Amyntas tokioensis. [JPN.] Fig.6 Amynthas tokioensis v. Homan : A. Ventral view of spermathecal pore. B. Ventral view of male pore. C. Spermathecal pore. D. Prostatic gonad and duct. E. Caec

8 Metaphire hilgendorfi
[Eng.] Note and distribution: Most individuals of Metahire hilgendorfi collected from Fukuoka prefecture was degraded morph lacking male organ (and sometime lacking also female organ). Only the specimens (of Metaphire hilgendorfi ) collected the vicinity of the top of Mt. Sefuri (Alt m) have with full sexual organ (of male and female). Son and Paik (1969) reported that Lots of specimens from many districts of mainland seldom had male pore. However, twenty-two out of twenty three specimens from Daglets Is; Evidently had both male pore and prostate gland. (Pheretima hilgendorfi in p15-16 of Son and Paik (1969). [JPN.] Fig. 7 Metaphire hilgendorfi (Michaelsen, 1892): A. Ventral view of spermathecal pore. B. Ventral view of male pore. (B’. Twice close-up of right male pore and the circumference.). C. Spermathecal pore. D. Prostatic gonad and duct. E. Caeca

9 Metaphire vesiculata [Eng.] Genital markings are nothing. There is one internal gland near each spermathecal duct. But, it was not “Nephridia on spermathecal ducts”. Male pores within shallow copulatory pouches on segment 18 with two internal gland near spermathecal duct.  There are two internal glands near the end of duct. [JPN.] Fig Metaphire vesiculata (Goto & Hatai, 1899): A. Ventral view of spermathecal pore. B. Ventral view of male pore. C. Spermathecal pore. (The diverticulum in 7/8 has come off from the duct.). D. Prostatic gonad and duct. E. Caeca.

10 Metaphire vesiculata v. Sefuri
[Eng.] Genital markings paired just on setal line near spermathecal pores of 6/7 and 7/8. There is small internal gland corresponding to the external papillae. Male pores present in a hollow place (a depression). Then, present taxa is put in genus Metaphire.  There are two genital papillae on setal line near male pore. Remark: If "Male pores are superficial in a hollow" is excluded, Sefuri and Shika, these are almost corresponding to Diagnosis of Amynthas tokioensis. Samuel W. James (2004) stated that decision should be evaluated with an independent data set, because it is possible that the small pockets of certain Amynthas could have evolved independently of the intramural pouches of Metaphire. However, the hollow of Sefuri and Shika can already be confirmed when alive. The problem whether these belong to Amynthas or belong to Metaphire remains. [JPN.] Fig. 9 Metaphire vesiculata v. Sefuri A. Ventral view of spermathecal pore. B. Ventral view of male pore. (B’. Twice close-up of right male pore). C. Spermathecal pore. (The diverticulum in 7/8 has come off from the duct.). D. Prostatic gonad and duct. E. Caeca.

11

12 Distribution map of each complete taxa belonging to species complex
Thick white line: Amynthas tokioensis. Thin white line: Amynthas vittatus, Thick yellow line: Amynthas tokioensis v. Hakozaki, Thin yellow line: Amynthas tokioensis v. Munakata, Thick red line: Metaphire hilgendorfi and Metaphire hilgendorfi v. Sefuri, Thin red line: Metaphire vesiculata. Thick pink line: Amynthas tokioensis v. Homan, Thick pink line: Amynthas tokioensis v. Akizuki, Blue thick dot line: Metaphire pseudovittatus.

13 [Eng.] [JPN.] 3-2) It was to have clarified the contradiction shown in the above figure. Group including super ficial and copulatoly pouches is only Metaphire hilgendorfi / Amynthas tokioensis species complex. Is there a method of a little more grouping without contradiction referring to this figure? It seems to lead another contradiction when dividing with Caeca. I forecast that the shape of prostatic duct can smoothly divide the group. 一つの group内部に、super ficialとcopulatoly pouchesを含むgroupは[[Metaphire hilgendorfi / Amynthas tokioensis species complex]]だけである。 この図を参照してもう少し矛盾なくグループ分けする方法はあるだろうか?Caecaで分けると別の矛盾を導きそうに思える。prostatic ductの形状はグループをスムースにわけることができる、と私は予想する。

14 Key to the genera of the Pheretima-complex of species
1 Body depressed, setae crowded ventrally Planapheretima Body cylindrical, setae evenly distributed. 2 2 Intestinal caecum(a) absent 3 Intestinal caecum(a) present 5 3 Oesophageal pouches present. Ephemitragen. nov. Oesophageal pouches absent. 4 4 Clitellum begins on segment xiii. Archipheretima Clitellum begins on segment xiv. Metapheretima 5(2) Intestinal caecum(a), origin at or near segment xxii Pithemeragen. nov. Intestinal caeca, origin at or near segment xxvii. 6 6 Copulatory pouches absent Amynthas Copulatory pouches present. 7 7 Nephridia on spermathecal ducts. Pheretima s.s. Nephridia absent from spermathecal ducts. Metaphire nov.   Partial revision of Sims and Easton 's Key to the genera of the Pheretima-complex may be as follow, 6       Nephridia on spermathecal ducts Pheretima s.s Nephridia absent from spermathecal ducts 7   prostatic duct U shape and muscular. The terminal of duct is connected directly with parietal      Amynthas prostatic duct not U shape Prostatic duct not U shape and not muscular, Prostatic duct not U and not muscular. The duct is ended to pad on parieted Metaphire The duct is not ??????? ????????????????? The duct is ????? ????? XX ????????????????????????????       [Eng.] [JPN.] 3-3 I want to slip out a tight situation of Metaphire hilgendorfi/Amynthas tokioensis species complex. The proposal above is this essay to slip out the complexity. 上の提案はこの複雑な状態を抜け出す為の試論。その為にBlakemore博士の “Restating scope of genus Metaphire Sims & Easton, years on”を読む。 Reference Blakemore R : “Restating scope of genus Metaphire Sims & Easton, years on“. Zoology in the Middle East Volume 58, Supplement 4, 2012


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