an addition of section 5. Name and rank of new taxa in method 2012/6/15
ssp. hakozakiensis - v. hakozaki, ssp. munakataensis- v. munakata
ssp. homanensis - v. homan
Metaphire hilgendorfi / Amynthas tokioensis species-complex (OLIGOCHAETA: MEGASCOLECIDAE) from Fukuoka Prefecture and its surrounding Kyushu Japan
Y. S.
Preface
The present author researched earthworm's ecology in the period from 1967 to 1974. At that time, the taxonomy of the earthworm in Japan was not advanced enough. The present author was not able to give the scientific name to most abundant earthworms. The taxonomical characteristic of (the) dominant earthworm is as follows; Spermathecal pores widely pairedin 6/7/8and male pores are large disc on segment18, and the intestinal caeca are manicate. Genital marking is scarce. The dominant earthworm was collected for the first time in Hakozaki shrine forest Fukuoka city Kyushu Japan. Then, it was called tentatively "Pheretima sp. (H-1)". "Pheretima sp. (H-1)"was frequently accompanied with degraded morph thatlacked a male pore though clitellum was possessed. Other external feature, except the absence of male organ, of degraded morph looked like to those of "Pheretima sp. (H-1)". Earthworm taxonomy at that time gave the scientific species name to these degraded morphs such as “Pheretima irregularis”, “Pheretima hilgendorfi”, Pheretima agrestis”. One of the reasons why the present author did not contribute my ecology research to the science magazine is in "Several abundant earthworm were the un-descriptions".
After 1974, the present author has collected the earthworm in various habitats of the Fukuoka prefecture and its surroundings, Kyushu Japan. In these earthworm collections, there were several typesthat genital marking around a male pore is different though it looked like "Pheretima sp. (H-1)".
Recentlythe earthworm taxonomy of Japan has been arranged by Blakemore 2003 2005（After Blakemore 2003）. It came to be able to examine my collection by the idea of " Metaphire hilgendorfi/ Amynthas tokioensis species-complex" advocated by Blakemore 2003 & 2005.
At first, several new taxa having full sexual organ being like to Amynthas tokioensisare described (chapter one). In chapter two, the emergence frequency of degraded morphs belonging to Metaphire hilgendorfi / Amynthas tokioensis species complex is studiedand discuss the relation between the earthworm having full sexual organ and the degraded morph lacking male organ.In chapter three, these new taxa are located toMetaphire hilgendorfi / Amynthas tokioensisspecies complex that Blakemore advocates and discussed the relation between complete morph and degraded morph.
Chapter One
Several taxa belonging to Metaphire hilgendorfi/Amynthas tokioensis species complex having full sexual organ in Fukuoka Kyushu Japan,in relation to "After Blakemore 2003"
[Abstract:Metaphire hilgendorfi/Amynthas tokioensis species complex consists of parthenogenetic taxa (Degraded morph) and the taxafully equipped with the reproductive system(Complete morph). Only the specimen fully equipped with the reproductive system among them was examined. Fivenew varieties were collected from Fukuoka Prefecture, besides four known taxa: Amynthas tokioensis, Amynthas vittatus, Metaphire vesiculataand Metaphire hilgendorfi. New taxa are Amynthas tokioensis v. akizuki (four pair of genital marking in pre and post setal line of segment 18), Amynthas tokioensis v. hakozaki (large disc of male pore) and Amynthas tokioensisv. munakata (middle phase between A.tokioensis and A.tokioensis v.hakozaki), Amynthas tokioensis v. homan(three pair of spermathecal pore in segments 5/6/7/8 with superficial male pore) and Metaphirehilgendorfi v. sefiri (male pore in hollow).
[Keyword: Blakemore, Metaphire hilgendorfi/Amynthas tokioensisspecies complex,Degraded morph, Complete morph, Fukuoka prefecture]
Introduction
Blakemore (2003, 2005) said, Resolution of the Metaphire hilgendorfi (/Amynthas tokioensis)species-complex is one of the most pressing and seemingly intractable problems in Japanese (and Korean) earthworm systematics. 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.
Presentchapter examines the earthworm that has a complete reproductive system at the beginning of series. Several New taxa like to “Pheretima sp. (H-1)”are described. And, the dominant taxa in my ecological study from 1967 to 1974:“Pheretima sp. (H-1)” is located in Metaphire hilgendorfi /Amynthas tokioensis species complex.
Material and Method
1. Sampling and preservation
The earthworm was collected from Fukuoka Prefecture from 1974 to 2004 in about 182 sites of various vegetations (195 times in total). In each sampling, the earthworm was collected during one hour while removing litter and soil to 5-7cm depths with a small shovel. The earthworm collected was moved to my experiment room and were put into water. They were anesthetized by adding drop-by-drop absolute alcohol slowly. The earthworm anesthetized was put between the glass sticks, and was fixed with 10% formalin solution for a day. Later, the specimens were kept in 5 % formalin solution. The specimens were identified, and their body length and body widths were measured.
The earthworm specimen obtained in the population study from 1967 to 1974 was preserved in 70% alcoholic liquid by stopping the cork. Tens of years pass, an alcoholic liquid for preservation disappears, and the majority of specimens have dried up. Small number of specimens escaped drying, and these were used for a part of presentstudy.
2. Observation of taxonomic feature
The following character are observed and recorded.
External feature:Body-shape, body length, body width, body colour, segment number, setae number on 10thand 20thsegments. The position and the shape of spermathecal pore, male pore and Genital marking.Body circumference apart of spermathecal pores and of male pore. The shape of genital marking on body surface.
Internal feature: The shape of spermatheca, prostatic gonad and prostatic duct, and the shape of caeca and Inner gland corresponding to genital marking on body surface.
3. Observation and Illustrations
Dorsal dissections were performed under binocular microscope, specimens were pinned on a plastic tray containing waters. Most feature and anatomical detail are symmetrical. In the sketch of surface, externals of right male pore and spermathecal pore, in observer view, were chiefly sketched. In the sketch of internal, Left organs, in observer view, weresketched.
All specimens were sketched in using the photograph.Olympus C-2100 Ultra zoom (2,100,000 pixels),with macro conversion lens (MCON-40). The camera is fixed to the photographing stand. The photograph <A4 size> is printed out. The line is retouched on the photograph with the pencil. The photograph is put on BANKO trace stand B4. It draws to the tracing paper in earthworm's morphologic characteristic, while confirmingthe position and the shapes of Vas deferens, and Prostate gland and duct with eyes under binocular microscope (Nikon SMZ 1B).
4. Referred document and Abbreviation
The research chiefly referred to Blakemore 2003-2005.Blakemore 2010 changed a part of the composition of species complex as follows: (1) Change from Metaphire agrestis to Amynthas agrestis and Metaphire hataii was restored from Metaphire agrestis, (2) Metaphire soulensis is restored to Metaphire hilgendorfi spp-complex.These changes are described clearly in each part.
Metaphire hilgendorfi/ Amynthas tokioensis species complex is often abbreviated with Metaphire hilgendorfi spp-complex.
5. Name and rank of new taxa
Blakemore (201X) stated, The Metaphire hilgendorfi (Michaelsen,?1892)/Amynthas tokioensis (Beddard, 1892) parthenogenetic/clonal spp-complex has since snowballed into >60 names and its resolution remains the hottest yet seemingly intractable problem in Oriental (and Cosmopolitan) earthworm systematics.Several new taxa of this species complex were collected also in Fukuoka Kyushu Japan. However, these new taxa are characterizedonly by the difference of signs on body surface and its inner glands. I hesitate to describe these new taxa as new species or new sub species disregarding the point of Blakemore (201X).
Various categories such as variety, form, and morph, race, mutant, and seasonal form might be distinguished and be named from subspecies further below. However, these categories are put outside the application of Animal Nomenclature (Koji Yokokawa, ICZN 45.5 and 45.6. ).
Nomenclature does not determine the inclusiveness or exclusiveness of any taxon, nor the rank to be accorded to any assemblage of animals, but, rather, provides the name that is to be used for a taxon whatever taxonomic limits and rank are given to it(principle (2) of Zoological nomenclature ). The name and rank of taxa in Metaphire hilgendorfi / Amynthas tokioensis parthenogenetic/clonal spp-complexare in a troublesome situation as the point of Blakemore (2011). In such situation, I report newly taxa collected in Fukuoka Kyushu Japan only as "variety" which are put outside the application of Animal Nomenclature.
The diagnosis of Metaphire hilgendorfi / Amynthas tokioensis species complex has the possibility to be rewritten. Rank of taxa described as variety here will be examined again after the rewriting the diagnosis of Metaphire hilgendorfi / Amynthas tokioensis species complex.
Result
Amynthas tokioensis species -group
Amynthas tokioensis (Beddard, 1892)
Material examined: Ohmuta C. Tenmangu July 2, 1979 B.L. 138 x 5.8mm, Ohmuta C. Kubuki July 13, 1979, 138.0 x 7.3mm, Yabe V. Mt. Syaka –Gozen. July 24, 1991, 108 x 5.3 mm. Fig. 1 is the Sketch of Ohmuta Tenmangu specimen. (Measurement record: Ohmuta, Kubuki, Syaka)
Diagnosis: Two pairs of spermathecal pores ca. 0.35 -0.41body 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 pores ca. 0.35-0.41 circumference apart with 7-16 setae between male pores. One genital marking present on setal line near male pore. Intestinal caeca is manicated on intestinal canal of segment 27.
Fig. 1 Amynthas tokioensis (xxxx, 18xx): A. Ventral view of spermathecal pore. B. Ventral view of male pore. C. Spermathecal pore. (The Ampula in 6/7 has come off from the duct.). D. Prostatic gonad and duct. E. Caeca
Diverticulum is equal (Kubuki andSyaka) or shorter (Ohmuta) to Ampula in length. Prostatic gonad is large occupyingsegments 16-23 separating two or more plates, and Prostatic duct is U (Ohmuta and Syaka) or loop (Kubuki) shape. The duct is slender inearly 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)". Blakemore (2005) locates Amynthas irregularis and Amynthas levis (Goto & Hatai) which is degraded morph to the synonym of A. tokioensis. In addition, many taxa having full male organ such as Amynthas purpuratus(Ishizuka, 1999b). comb. nov., Amynthas tappensis (Ohfuchi, 1935) and Amynthas vittatus (Goto & Hatai, 1898), these are listed in synonym of A. tokioensis. Blakemore 2005 said, thus Amynthas tokioensis becomes the representative taxon of an Amtnthas tokioensis species-group currently combined, uncomfortably, within the Metaphire hilgendorfi species complex.
Amynthas vittatus [Synonymy of Amynthas tokioensis(Goto & Hatai, 1898) type?]
Material examined: Kitakyusyu C. Yahatanishi-kuGoto park 151.0 x 6.9mm, Asakura C. Akizuki Akizukijo August 3, 1979 168 x 9.0 mm, Maebaru C. Umihatchiman shrin near nagaito elementary school June 26, 2003, 129.0 x 7.0mm, Chikusino C. Mt. Tenpai,June 11, 1988, 138.0 x 6.5mm.Fig. 2 is the Sketch ofGoto. (Measurement record: Goto, AkiB?, Nagaito and Tenpai)
Fig. 2 Amynthas vittatus (xxxx, 18xx): A. 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.
Diagnosis: Two pairs of spermathecal pores ca. 0.41 -0.49 body circumference apart in furrows 6/7/8.Diverticulum is equal to Ampula in length. Genital markings in paired sets of three or four papillae, with internally small glands and several bottle like glands, linearly in segments 7 and 8 just in behind of spermathecal pores, and similar markings on setal line near male disc and along disc on segment 18, with mass glands internally. Male pore are superficial. Male pores ca. 0.35-0.42 circumference apart with 7-12 setae between male pores.Prostatic gonad are large occupying segments 16-22, and Prostatic duct is U shape and large. There is glandular mass just before duct. Intestinal caeca manicate.Colour banded colouration is not clear in many individuals.
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.
Amynthas tokioensis var. akizuki4/9/2011
Material examined:Asakura C. Akizuki, Suiyuu Shrine,July 11, 1988, 163mm x 8.0mm. Fig. 3 is the Sketch of Aki specimen. (Measurement record; Aki).
Diagnosis: Two pairs of spermathecal pores ca. 0.44 body circumference apart in furrows 6/7/8.Spermathecal pores widely pairedin 6/7/8. Diverticulum is equal to Ampula in length. 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 segment8.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). Male pores are superficial.Male pores ca. 0.38 circumference apart with 16 setae between male pores. Prostatic gonad occupied segments 16-21, separating two plates. A prostatic ductis U shape.There are glandular masses in just before Duct and mid ventral side. The intestinal caeca are manicate.
Remark: The feature of this specimen is in“mass glands on the center ofventral wall of segment 8”and “Two pairs of genital markings in pre- and postof 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).
Fig. 3 Amynthas tokioensis var. akizuki : A. Ventral view of spermathecal pore. B. Ventral view of male pore. C. Spermathecal pore. D. Prostatic gonad and duct. E. caeca.
Etymology. Named after the locality.
Note and distribution: Asakura C. Akizuki in evergreen forest of Suiyuu Shrine and Mt. Kosyo. This species was collected with Amynthas vittatus(AkiB?) in a site: Suiyuu Shrine’s forest.
Amynthas tokioensis var. hakozaki
Material examined: Dazaifu C. Hokanken, July 19, 2003, 85 x 5.7mm. Dazaifu C. Kanzeonji June 22, 2003, 157 x 7.8 mm. Fukuoka C. Hakozaki June 19, 1972, 95x 6.0mm. Fig. 4 is the Sketch ofKanzeon. Measurement record (Hokanken, Kanzeon, H1(Hakozaki)
Diagnosis: Two pairs of spermathecal pores ca. 0.40 -0.43 body circumference apart in furrows 6/7/8. Diverticulum is equal to Ampula in length. There is a small genital marking mid ventral in pre-setal line onmore likely segments 7 and/or 8, of some individuals.There is no internal gland corresponding to external genital marking.
Male poresare large disc (superficial). There must be two genital papillae (penial setae) along the circumference of large disc. Male pores ca. 0.32-0.37 circumference apart with 8-10 setae between male pores.Prostatic gonad greatly extends over segments 14-22. Prostatic duct is large in U type.There is glandular mass near the end of duct. The intestinal caeca is manicate.
Remark andDistribution: Male pore of A. tokioensisv. hakozakiis 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. The distribution of Amynthas tokioensis v. hakozakihas 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 hakozaki is distributed around the distribution region of A. tokioensis and A. vittatus. It is thought that A. tokioensisv. hakozaki is variety or ecotype of Amynthas tokioensis.
Etymology. Named after the locality.
Note: Amynthas tokioensis v. hakozaki was dominant in my ecological study 1967-1974. In the ecology research, Amynthas tokioensisv. hakozaki was called tentatively as Pheretima sp. (H-1).
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.
Amynthas tokioensisvar. munakata
Material examined: Munakata C. Munakata big shrineJune 22, 1979, BL. 135 x 6.7 mm, Munakata C. Hataoriguu schrine July 14, 1977, 130mm x 6.9mm, Oki Munakata C. Okinoshima island July 25, 1986, BL. 141x 8.5mm. Fig. 5 is the Sketch of Munakata. (Measurement records: Munakata, Kanasaki, Jin).
Diagnosis: Two pairs of spermathecal pores ca. 0.44 -0.50 body circumference apart in furrows 6/7/8.Diverticulum is equal toAmpula in length. There is a small genital marking at mid ventral in pre-setal line with several internal glands, onsegment 7 of Munakata specimen. In another specimen of this taxa :(Kanasaki), there are no genital marking in spermathecal segments, but there are a internal gland near spermathecal pore of 7 and 8 segments.
Male poresare large disc (superficial). But, the disc is slightly smaller than that of A. tokioensis v. hakozaki.Male pores ca. 0.35-0.40 circumference apart with 9-12 setae between male pores. 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.Prostatic duct is large in U or loop shape.There is glandular mass just before Duct. The intestinal caeca are manicate.
Remark: Male disc of A. tokioensis v. munakata 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. munakata 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 munakata is the intermediate phase of A. tokioensis and A. tokioensis v. hakozaki.