Saturday, October 29, 2011 ~Sunday, October 30, 2011 The style of the quotation from sims and Easton shown in p18 was changed.
This file was cut off from Chapter 3 on November 9, 2011.
Jump to Sims & Easton
Easton
Blakemore
Others Gamou et al.
------
Appendix 1A Sims & Easton 1972
A similarity matrix was obtained from a consideration of 56 characters, it was examined by the method of principal co-ordinates analysis firstly to which a minimum spanning tree was applied, then secondly by using a five nearest neighbours' procedure. From the arrangement of the species, assemblages were detected which permitted the recognition of eight genera (one being divided into subgenera): Archipheretima, Pithemera gen. nov., Ephemitra gen. nov., Metapheretima, Planapheretima, Amynthas, Metaphire nom. nov., Pheretima (Pheretima) and Pheretima (Parapheretima). Following research into the literature, most known species were assigned to genera and to nominal species-groups within the genera, only a few being listed as species incertae sedis (Sims & Easton (1972) Biol. J. Linn. Soc., 4: 169-268 [p169])
hawayanus-group
Amynthas
/agrestis
/Goto & Hatai, 1899: 17
Amynthas
/communissimus *
/Goto & Hatai, 1899: 65
=sieboldi lenzi
/Micaelsen. 1899b:9
Amynthas
/hilgendorfi
/Michaelsen, 1892: 235 (part)
illotus-group
Amynthas
/irregularis
/Goto & Hatai, 1899: 14
morrisi-group
Amynthas
/koreanus
/Kobayashi, 1938a: 119
pomellus-group
Amynthas
/vittatus
/Goto & Hatai, 1899: 74
sieboldi-group
Amynthas
/yamadai
/Hatai, 1930: 644
Amynthas
/yunoshimensis
/Hatai, 1930: 655 (part)
tokioensis-group
Amynthas
/gomejimensis
/Ohfuchi, 1937a: 18
Amynthas
/hilgendorfi
/Michaelsen, 1892: 235 (part)
parvicystis
/(Goto & Hatai, 1899: 18)
Amynthas
/rukugo
/Beddard, 1892b: 763
Amynthas
/tappensis
/Ohfuchi, 1935: 409
Amynthas
/tokioensis
/Beddard, 1892b: 762
Amynthas
/yunoshimensis
/Hatai, 1930: 655 (part)
youngi-group
Amynthas
/hilgendorfi
/Mihchaelsen, 1892: 235 (part)
zebrus-group
Amynthas
/hilgendorfi
/Mihchaelsen, 1892: 235 (part)
Provisional list of Genus Amynthas :Appendix 1f is shown in p234-237 of Sims and Easton 1972.
glandularis-group
Metaphire
/glandularis
/Goto & Hatai, 1899: 18
Metaphire
/levis
/Goto & Hatai, 1899: 20
Metaphire
/servinus
/Hatai & Ohfuchi, 1937: 1
Metaphire
/soulensis
/Kobayashi, 1938a: 131
Metaphire
/vesiculata
/Goto & Hatai, 1899: 21
Metaphire
/hataii
/Ohfuchi 1937a: 13
Provisional list of Genus Metaphire :Appemdix 1g is shown in p237-239 of Sims and Easton 1972.
The species which looked like Amynthas tokioensis hakozakiensis was chosen from provisional list and it showed in the table.
Appendix 1B R. W. SIMS AND E. G. EASTON 1972 P172
Variation in expression in these and other numerous features
provide the characters for delineating species
1. Body-shape.
2. Setae.
3. Dorsal pores.
4. Clitellum.
5. Male pores.Paired, usually on segment xviii exceptionally on segments xix or xx. They provide a common opening for the vasa deferentia and the prostatic ducts. They discharge either directly onto the ventral body surface (usually through papillae) (Fig. IA) or open into paired copulatory pouches (Fig. 1B).They are minute but where they discharge into copulatory pouches, the external openings to the latter are usually referred to as "male pores" and these are large with tumid lips. The distances apart of the paired pores are variable.
6. Copulatory pouches. These structures which are sometimes known as bursae copulatrices, when present, form the ectal portion of the posterior male system. Often each is a simple pouch-like invaginationof the ventral body-wall but secretory pads, secretory diverticula which are tubular (diverticulate "muscular" glands), stalked glands, glandular papillae, a penis or even penial setae may be associated with it (Fig 1C, D). Occasionally nephridia are present on the coelomic surfaces of the pouches.
7.Female pore.
8.Spermathecal pores. Variation in the pores is determined largely by the distribution, numbers and sizes of the spermathecae. They occur mainly in a single intersegmental furrow or a series of consecutive furrows between 4/5 and 8/9 (exceptionally 9/10), infrequently the pores are intersegmental. Usually paired but they may be single or numerous. When there is more than one, the distances that the pores are apart is also variable. Additionally they may be minute (and difficult to see) or large with tumid lips. (The occurrence of large spermathecal pores can be correlated with large "male pores".)
9.Papillae.
10.Septa.
11.Coelomic sacs. .
12. Gizzard.
13. Oesophageal pouches.
14. Calciferous glands.
I5 Intestine.
16. Lateral hearts.
17. Testes.
18. Seminal vesicles.
19. Spermathecae.
20. Nephridia. ------The excretory systems of these worms are complex and their comparative study is a sufficiently large task to constitute a separate investigation. In view of the large number of species involved this aspect of the morphology of the worms is clearly beyond the scope of the present work. As it was often not possible to obtain adequate information on nephridia, for the purposes of this investigation the characters of the excretory system had to be omitted. (Either the descriptions of species seldom contained detailed observations of the nephridia, or much of the material examined was inadequate due to previous dissection or poor preservation.)
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.
Key to the species and nominal species-groups of the genus Amynthas
1 Spermathecal pores intersegmental' 9
9(1) First spermathecal pores behind 4/5 15
23(15) First spermathecal pores at 6/7 24
24 More than one thecal segment 25
25Two thecal segments 26
26 Bithecate 27
27Holandric tokioensisgroup
Key to the species andspecies-groups of the genus Metaphire
" Athecatc examples of several species assigned to the genus Metaphire have been recorded. At the present time it is not possible to distinguish between athecate specimens of Metaphire and Pheretima (Pheretima).
1 First spermathecal pores behind 4/5 ` 3
3(1) First spermathecal pores behind 5/6 24
24(3) First spermathecal pores at 6/7 25
25More than one thecal segment 27
27(25) Two thecal segments 28
28Intestinal caeca, structure multiple glandularis-group
Appendix 2 Easton 1981
Easton’s remark: Nineteen species and subspecies are included within the hilgendorfi species complex, they exhibit a wide range of variation in the expression of several characters, especially in the arrangement of genital markings, number of spermathecal furrows and the degree of development of the male pores. Insufficient data are currently available either to establish the validity of the component taxa or to recognize discrete subgroups within the complex. (Bull Br. Mus. nat. Hist. (Zool.) 40 (2): 33-65 [p33])
Easton 1981? Or 1982
/syn
Amynthas hilgendorfi species-complex
Amynthas
/hilgendorfi
/Michaelsen, 1892 : 235
/Rokugo, irregularis, schizopora Beddard, 1900
Amynthas
/rokugo
/Beddard, 18926 : 756.
Amynthas
/tokioensis
/Beddard, 18926 : 759
Amynthas
/vittata
/Goto & Hatai, 1898 : 74
Amynthas
/schizopora
/Goto & Hatai, 1898 :76.
Amynthas
/irregularis
/Goto & Hatai, 1899 : 13.
Amynthas
/agrestis
/Goto & Hatai, 1899 : 17
Amynthas
/glandularis
/Goto & Hatai, 1899 : 18.
Amynthas
/communissima
/Goto & Hatai, 1899 : 23.
/syn. sieboldi: Goto & Hatai, 1898
Amynthas
/sieboldi lenzi
/Michaelsen, 1899 : 9.
Amynthas
/ambigua
/Cognetti, 1906 : 782.
Amynthas
/yunoshimensis
/Hatai. 19306 : 655
Amynthas
/iappensis
/Ohfuchi, 1935 : 409.
Amynthas
/gomejimensis
/Ohfuchi; 1937a : 18.
Amynthas
/gucheoensis
/Song & Paik, 1970.
/not recorded from Japan
Amynthas
/jiriensis
/Song & Paik, 1971.
/not recorded from Japan
Amynthas
/koreana
/Kobayashi, 1938a
/not recorded from Japan
Amynthas
/shinkeiensis
/Kobayashi, 1938a.
/not recorded from Japan
Metaphire
Metaphire
/hataii
/Ohfuchi, 1937
Metaphire
/servina
/Hatai & Ohfuchi, 1937
Metaphire
/yamardai
/HaTai, 1930
Metaphire
/soulensis
/Kobayashi,- 1938a
/syn. Metaphire yamardai??
Pheretima (Parapheretima
Pheretima (Parapheretima)
/koellikeri
/Michaelsen, 1928
Amynthas hilgendorfi species-complex
DIAGNOSIS. Spermathecal pores absent or paired, ventrolateral in furrows 5/6/7/8 or 6/7/8 or 6/7 or 7/8. Male pores absent or superficial. Large clusters of genital markings or indistinct pigmented areas on pre- and postclitellar segments. Intestinal caeca manicate, each with about 5 diverticula, originating in segment 27.
REMARKS. Nineteen species and subspecies are included within the hilgendorfi speciescomplex, they exhibit a wide range of variation in the expression of several characters, especially in the arrangement of genital markings, number of spermathecal furrows and the degree of development of the male pores. Insufficient data are currently available either to establish the validity of the component taxa or to recognize discrete subgroups within the complex.Appendix 3A Blakemore 2003-2005
Composite diagnosis by Blakemore 2003-2005 was as follow,
Spermathecal poreswidely paired in some of 5/6/7/8, or6/7/8/9, or 6/7/8 or 6/7, or 7/8. Male poressuperficial (Amynthas), or in copulatory pouches (Metaphire). Genital markings absent, or as clusters of one or more papillated pores, or as indistinct pigmented areas on pre- and postclitellar segments. Intestinal caeca manicate originating in segment 26 or27.
[A red underline shows the part of degraded morph.]
Following Blakemore’s recent inspection of the type, Amynthas tokioensis becomes the representative taxon of an Amtnthas tokioensis species-group currently combined, uncomfortably, within the Metaphire hilgendorfi species-complex, (Blakemore 2005, Japanese earthworm, PDF file p88, pp147)
Blakemore 2005
Metaphire hilgendorfi / Amynthas tokioensis species- complex
Amynthas
/ambiguus
/Cognetti, 1906
/-55
Amynthas
/gomejimensis
/Ohfuchi, 1937
/-56
Amynthas
/koreanus
/Kobayashi, 1934
/syn. Pheretima conjugata Ishizuka 1999- synonym as per Blakemore, 2003: 43, addenda
/#57
Amynthas
/purpuratus
/Ishizuka, 1999b
/-58
Amynthas
/tappensis
/Ohfuchi, 1935
/syn. bimaculata, silvatica, ?surcata, odaesanensis Hong & James, 2001, righii Hong & James, 2001, 43, addenda; sanchongensis Hong & Hanes, 2001 Syn. nov.).
/-59
AmyntShas
/tokioensis
/Beddard, 1892
/(syns. ?Perichaeta schizopora Goto & Hatai, 1898: 76 Syn. nov., ?Perichaeta irregularis Goto & Hatai, 1899: 13 Syn. nov., Perichaeta levis Goto & Hatai, 1899; 20 Syn. nov., ?Syn. nov., ?parvicystis; ?verticosa; ? ?Amynthas yongshilensis Hong & James, 2001: 80 Syn. nov., ?A. eastoni Hong & Hames, 2001:83; ?A. boletiformis Hong & James, 2001: 84 (these last two synonyms as per Blakemore, 2003: 43, addenda); ?Amynthas paiki Hong in Hong, Lee & Kim, 2001: 266 Syn. nov.).
/#60
Amynthas
/vittatus
/Goto & Hatai, 1898
/-61
Amynthas
/yunoshimensis
/Hatai, 1930
/-62
Metaphire
/agrestis
/Goto & Hatai, 1899
/syn. hataii, striata syn. nov.
/-63
Metaphire
/communissima
/Goto & Hatai, 1899
/syn. Perichaeta sieboldi: Goto & Hatai, 1898 [non Megascolex sieboldi Horst, 1883 (= Metaphire sieboldi)]; sieboldi lenzi ; florea syn. nov.).
/-64
Metaphire
/hilgendorfi
/Michaelsen, 1892
/(syn. rokugo; ?schizopora; ?irregularis Goto & Hatai, 1899 (non Spencer,1895=Perionychella irregularis); glandularis).
/#65
Remark:The list of included species in the “Amynthas hilgendorfi species –complex” by Easton (1981) was fairly extensive and he appeared to accept the synonymies of A. hilgendorfis. by Beddard (1900) rather than those advocated by Michaelsen (1900). Studies by the current author have found specimens agreeing with M. hilgendorfi that have male pores (paired or single and sometimes displaced to segment 17 or 19) either in copulatory pouches or everted, and some other accounts (e.g. Ishizuka 2000d, 2001) have morphs with male pores that appear in copulatory pouches and, if this is taken as the normal situation, then this taxon belongs in Metaphire rather than Amynthas.(97/147 of Japanese earthworm PDF)
Metaphire
/servina
/Hatai & Ohfuchi, 1937
/-66
Metaphire
/vesiculata
/Goto & Hatai, 1899
/syn. ?koellikeri syn. nov.; ?okutamaensis syn. nov.; ?biggiberosa syn. nov
/#67
Metaphire
/yamadai
/Hatai, 1930
/syn. ?pectinifera Michaelsen, 1931 –synoynym as per Blakemore, 2003: 43, addenda).
/*68
End of Metaphire hilgendorfi/Amynthas tokioensis species complex
Metaphire
/soulensis
/Kobayashi, 1938
/#75
Appendix 3A
Following my recent inspection of the type, we can now accept that the intestinal caeca are indeed manicate, and male pores (when present) are superficial; thus Amynthas tokioensis becomes the representative taxon of an Amtnthas tokioensis species-group currently combined,uncomfortably, within the Metaphire hilgendorfi species-complex, (Blakemore 2005 Japanese earthworm, p88-89)
Concerning Amynthas/Metaphire, the division is straightforward when we have clear differences such as in M. sieboldi (that has copulatory pouches with huge eversible “air-bags with glands and penes as are found in M. bipora and possibly M. malayana), but naturally there are sometimes borderline cases that may go either way. I think the answer is to simply ask “arethe male pores superficial (=Amynthas), or are they non-superficial (=not Amynthas)”. (Cited Blakemore 2005 p38-39 “Genus Amynthas Kinberg, 1867”)
The Metaphire hilgendorfi / Amynthas tokioensis species-complex (Amynthas hilgendorfi species-complex sensu Easton 1981) remains one of the most intractable and pressing problems for comprehension of the Japanese fauna as most of the component taxa, e.g. Metaphire agrestis (Goto & Hatai, 1899), are parthenogenetically degraded morphs as yet unaffiliated with their ancestral and biparental populations. (Blakemore –2003 Japanese earthworm p1, Abstract).
Resolution may be sought employing combinations of morphological and molecular (RNA, DNA) techniques to determine specific affinities while also complying with requirements of the International Code of Zoological Nomenclature (ICZN 1999). (Blakemore –2003 PDF 03-01, Japanese earthworm, p1, [Section of Abstract].
More work is obviously required to sort the parthenogenetic morphs into their respective taxa, and also to separate Amynthas species from Metaphire species, assuming that these genera are tenable within such a species complex subject to male pore degradation. However, it is possible that the manicate species Amynthas tokioensis (Beddard, 1892) if it actually has manicate caeca, is the representative of an Amynthas group that can be separated off from the Metaphire hilgendorfi species-group. (Blakemore –2003 PDF 03-01, and 2005 Japanese earthworm, Remarks on Metaphire hilgendorfi species complex in p32).
Appendix 3B Blakemore’s Saga story 2010
1/ Today, the Metaphire hilgendorfi (Michaelsen, 1892) / Amynthas tokioensis (Beddard, 1892) parthenogenetic / clonal species-complex (BLAKEMORE 2003) has snowballed into >60 names, and its resolution yet remains the hottest, most pressing and seemingly intractable problem in Oriental (and Cosmopolitan) earthworm systematics.
2/ chronological description and “hilgendorfi has priority” :
Blakemore 2010 said; Key arrangement in chronological order rather than alphabetically for reasons of priority and to allow progressive elimination of possibilities for practical morphic identification).
3/ Correction and revival.
3-a/ “-63. Metaphire agrestis (Goto & Hatai, 1899)”in Blakemore 2005 divided into Amynthas agrestis and Metaphire hatai in Blakemore 2010. Metaphire hataii (Ohfuchi,1937: 13) comb. nov. provisionally restored from Amynthas agrestis synonymy.
3-b/ Metaphire soulensis (Kobayashi, 1938) was put in to species complex in Blakemore 2010.
4/ DNA analysis (of types or neotypes from type localities, where known) is probably the only feasible means for objective and definitive resolution.///Molecular ‘solutions’ are meaningless without DNA analysis of types under the strict ICZN Principle of Typification in chronological order under its Principle of PriorityFor all these issues, molecular resolution via DNA anaylsis of types is advocated.
5/ Whether non-superficial male pore invaginations are “entirely intramural” or in “copulatory pouches” (e.g., GATES 1982) is entirely irrelevant for genetic placement, except newly for Duplodicodrilus Blakemore, 2008. The difficulty then, is to progressively link a species’ degraded morphs to their amphimictic (i.e., sexual) ancestral forms that would pres(Introduction in Saga)umably have full complements of these organs. Complicating this goal is natural interspecific variability and earthworm plasticity.
Clonological list
‘Species’ included in Metaphire hildgendorfi / Amynthas tokioensis species complex
Metaphire hilgendorfi (Michaelsen, 1892)
[Published September, 1892 therefore has priority over BEDDARD’s December, 1892 P. rokugo and P. tokioensis]
Amynthas tokioensis (Beddard, 1892)
Perichaeta tokioënsis Beddard, 1892b: 762. [Published December, 1892 according to MICHAELSEN, 1900: 272].
[Amynthas vittatus (Goto & Hatai, 1898) (cf. A. tokioensis account above)]
Amynthas agrestis (Goto & Hatai, 1899)
see Metaphire hataii (Ohfuchi, 1937: 13) comb. nov. provisionally restored from Amynthas agrestis synonymy].
?Pheretima striata Ishizuka, 1999b: 53, figs. 91-101. ?Amynthas minjae Hong, in Hong, Lee & Kim 2001: 264, figs. 2A-C.
[Amynthas parvicystis (Goto & Hatai, 1899) (cf. Amynthas tokioensis above)]
[Metaphire? levis (Goto & Hatai, 1899) (cf. Amynthas tokioensis above)]
Metaphire vesiculata (Goto & Hatai, 1899)
Metaphire vesiculata: SIMS & EASTON 1972: 238 (Metaphire glandularis species-group); BLAKEMORE 2003
(syn. koellikeri cf. EASTON 1981); 2005; 2007 (koellikeri restored).
[?Pheretima köllikeri Michaelsen, 1928: 8, figs. 1, 2 (et P. koellikeri). From “Japan” – now restored as
?Metaphire koellikeri].
Metaphire communissima (Goto & Hatai, 1899)
Amynthas ambiguus (Cognetti, 1906)
Metaphire? koellikeri (Michaelsen, 1928)
BLAKEMORE 2005 (restoration as priority over vesiculata).
[Amynthas? yunoshimensis (Hatai, 1930) (cf. M. hilgendorfi)]
Metaphire yamadai (Hatai, 1930)
Amynthas defectus (Gates, 1930)
Amynthas tappensis (Ohfuchi, 1935)
Metaphire hataii (Ohfuchi, 1937)
Metaphire servina (Hatai & Ohfuchi, 1937)
Amynthas? gomejimensis (Ohfuchi, 1937)
Amynthas koreanus (Kobayashi, 1938)
Metaphire soulensis (Kobayashi, 1938)
[Amynthas gucheonensis (Song & Paik, 1970) (cf. A. tokioensis)]
[Amynthas jiriensis (Song & Paik, 1971) – see A. tokioensis]
[?Amynthas ? dageletensis Hong & Kim, 2005]
Gamou et al
The worms in the first branch constantly retain male pores and some kind of genital markings, but are a minor population in field. The first branch includes A. corticis, A. tamaensis, A. carnosus, and A. robustus.The second branch is a major in the wild population, most worms of them have neither apparent genital marking nor male pore. The second branch can be divided into two sub-branches, one consists of A. hilgendorfi and A. agrestis and another consists of A. hilgendorfi and A. vittatus. The later sub-branch has a variation in 18S rRNA gene. Taken together, our results suggest that Amynthas worms have a wide variation in morphology, are genetically, closely related, and can be divided into several groups Molecular data combined with morphological observation can provide rational classification and further insight how they maintain their population even though the majority of them lost their hermaphroditism. ////the gene difference between species without male pore is small and It is doubtful these gene difference is whether it is a difference in which species can be clearly divided. (Shinobu Gamou, Tomomi Hirayama, Tomoharu Suzuki, Makoto Honda, and Seiji Matsumoto 2006).
An investigation by reverse transcription-polymerase chain reaction (RT-PCR) revealed that, whilst evcp-1 was ubiquitously expressed during all developmental stages, evcp-2 was specifically expressed in the anterior segments of mature (ciliated ) earthworms. In situ hybridization clearly demonstrated that evcp-2 is expressed in the seminal vesicles, the location of spermatogenesis, and more precisely within the cytophores surrounded by secondary spermatocytes or spermatides. Taken together, this evidence leads to the notion that evcp-2 is a likely component involved in the final modulation of spermatogenesis. (Shinobu Gamou, hinobu Gamou1 Tomoharu Suzuki1, Makoto Honda1, Seiji Matsumoto1, and Stephen R. Stürzenbaum 2006)