Transcript Founding an Online Nomenclator of Clitellata: Oligochaeta

Summary of IOTM3
(International Oligochaeta Taxonomy Meeting 3)
Cyprus Presentations
by Rob Blakemore
COE fellow,
Soil Ecology Research
Group,Yokohama National University
IOTM Presentions
(Posters and Powerpoints)
1. COSMOPOLITAN EARTHWORMS. 2nd Edition (Blakemore, 2006).
2. Acanthodrilidae, Octochaetidae, Exxidae, & Megascolecidae revisited.
3. Review of pheretimoid (Pheretima auct.) taxa.
4. 'Wikiworm' - a universal matrix of megadriles x region x ecology.
5. Ecological Strategies of Earthworms.
6. A mystery worm from Tasmania.
7. Megascolex (Promegascolex) mekongianus Cognetti, 1922: its extent,
ecology and allocation to Amynthas (Oligochaeta: Megascolecidae)
8. Some General Observations!
1. Cosmopolitan Earthworms – an Eco-Taxonomic Guide
to the Peregrine Species of the World
Second Edition (2006)
This new CD Guide concerns the Biology, Ecology,
and Distribution of >120 Exotic Species found
most commonly around the World.
Species are figured and identified using keys.
All Species, Genera and Families are reviewed.
The Guide provides an essential tool for Students,
Ecologists and Soil Scientists.
.First edition (Blakemore, 2002) recommended
text for Soil Quality, species determination by
ISO/FDIS23611-1:2006.
Blakemore, R.J., (2006). Cosmopolitan Earthworms – an Eco-Taxonomic
Guide to the Peregrine Species of the World. (2nd Edition). VermEcology,
Japan. Pp. 600 + 150 figs.
Blakemore, R.J., (2002). Cosmopolitan Earthworms – an Eco-Taxonomic
Guide to the Peregrine Species of the World. VermEcology, Kippax, Australia.
Pp. 426 + 80 figs.
2a. Phylogeny of Acanthodrilidae, Octochaetidae,
Exxidae, & Megascolecidae revisited.
"A Darwinian classification, by using two criteria, similarity and common descent, leads to the
recognition of classes (taxa) of similar entities.." "..almost any method of weighing is
preferable to using unweighed characters"...
"..morphological characters, the product of large numbers of genes, are usually quite reliable"
- Ernst Mayr & W.J. Bock (2002: www.blackwell-synergy.com/servlet).
Table of weighted key characters for type-species of (sub-)family type-genera
(Derived states: Hearts >11, Non-acanthodriline, Non-tubular prostates, Non-holoic nephridia)
2b. Phylogeny of Acanthodrilidae, Octochaetidae,
Exxidae, & Megascolecidae revisited.
Simple phylogram above from basic weighted morphology data below – using PHY-FI
(Fredslund, 2006)
2c. Phylogeny of Acanthodrilidae, Octochaetidae,
Exxidae, & Megascolecidae revisited.
Support for the
current phylogeny,
using weighted
morphology, is that
resulting tree (on
left) is a good fit with
cladogram from
molecular analyses,
as presented at
IOTM2 in Romania
(Blakemore, 2005:
Fig. 1 - modified
slightly + EXXIDAE).
Strict comparisons yet require
DNA testing, ideally of (typespecimens of) the type-species
of the type-genera.
2d. Phylogeny of Acanthodrilidae, Octochaetidae,
Exxidae, & Megascolecidae revisited.
MEGASCOLECIDAE is separated from other
families in title above and is diagnosed purely by
its
derived,
non-acanthodriline
i.e.,
megascolecine, male field, irrespective of any
other character - see figure on right (from Lee
1959, Lee et al. 2000).
A possible sub-family with tubular prostates
(and holoic nephridia), is based on tribe
Argilophilini, (type American Argilophilus
marmoratus ornatus Eisen, 1893) by Fender &
McKey-Fender (1990) - although this name
competes for priority with Vejdovsky's (1884:
63) PONTODRILIDAE and/or PLUTELLIDAE
[types Indo-Australasian Pontodrilus litoralis
Perrier, 1874, and Australian Plutellus
heteroporus Perrier, 1873 that is now
(Blakemore, 1994) restricted to central coastal
New South Wales, respectively].
2e. Phylogeny of Acanthodrilidae, Octochaetidae,
Exxidae, & Megascolecidae revisited.
Proposed restoration of the sub-family PONTODRILINAE
with
tubular
prostates
leaves
the
residue
in
MEGASCOLECINAE diagnosed by derived non-tubular (i.e.
tubuloracemose or racemose) prostates agreeing with the Sri
Lankan type, Megascolex caeruleus Templeton, 1844.
Figure on right-side shows prostates, tubular only in top row;
all others are derived “non-tubular” forms.
(Top left is paired prostates of Acanthodrilidae).
(Top right is Pontodrilus litoralis – Pontodrilinae).
(Low right special euprostate of Eudrilus eugeniae - Eudrilidae).
2f. Phylogeny of Acanthodrilidae, Octochaetidae,
Exxidae, & Megascolecidae revisited.
Further, secondary and subordinate division may be
"convenient"
within
the
two
sub-families
MEGASCOLECINAE and PONTODRILINAE on basis
of such features as holoic vs. non-holoic nephridia (on
right) or lumbricine vs. non-lumbricine setae (below).
2g. Phylogeny of Acanthodrilidae, Octochaetidae,
Exxidae, & Megascolecidae revisited.
Due to the “well known dependence of the conformation of the alimentary tract on food and
environment” (Stephenson, 1930: 720), the position within OCTOCHAETIDAE [type N.Z.
Octochaetus multiporus (Beddard, 1885)] of sub-family BENHAMIINAE, currently
defined by its arrangement of calciferous glands, is not fully resolved.
Neither is the status, validity nor extent within ACANTHODRILIDAE (type New
Caledonian Acanthodrilus ungulatus Perrier, 1872) of the polygiceriate DIPLOCARDIINAE/-IDAE (type North American Diplocardia communis Garman, 1888) – that possibly
merits separate family status. See REGULATION TABLE below:
3a. Review of Oriental pheretimoid (Pheretima
auct. family Megascolecidae) taxa
Mt Kinabalu Pheretima darnleiensis (above left), Japanese Metaphire sieboldi
(above right). Do you know the worm on right? 
Red line encircles endemic “Pheretima domain” in map below.
ANSWER: Amynthas rodericensis.
3b. Review of Oriental pheretimoid (Pheretima
auct. family Megascolecidae) taxa
Currently pheretimoids number ca. 920 (sub)species from a total of >1,300 taxa,
including numerous synonyms, invalid names and lapsae.
The Pheretima-group previously consisted of about 800 nominal species for which Sims
& Easton (1972) and Easton (1979) thought about half (ca. 400) were valid. Forty
species are COSMOPOLITAN EXOTICS. All taxa are checklisted (Blakemore, 2004,
2005, 2006).
Twelve genera are described (see next slide for summary).
Funding support is require to complete compilation of a searchable and key-able
database of morphological characters plus distributional data for each taxon, while full
taxonomic resolution probably requires DNA 'fingerprinting', preferably of (neo)types,
especially for the numerous parthenogenetically degraded polymorphs.
3c. Review of Oriental pheretimoid (Pheretima
auct. family Megascolecidae) taxa
Characters of Pheretima-group genera incl. Dendropheretima and Isarogoscolex from Philippines by James (2004)
3d. Review of Oriental pheretimoid (Pheretima
auct. family Megascolecidae) taxa
Species similar to Japanese Metaphire schmardae schmardae (Horst, 1883) having large
eversible copulatory pouches with duplicated paired male intromittant organs or pseudopenes (and manicate/multiple intestinal caeca) merit placement in a newly proposed genus.
Thus there is progression from superficial male pores (Amynthas) to non-superficial
(Metaphire/Pheretima) with proposed new genus most derived in its complex eversible pores.
4a. 'Wikiworm' a universal matrix of
Earthworms x Region x Ecology
This presentation proposes extension of an existing program in order to fill gaps in
knowledge and to present online bioinformatics about ecology and taxonomy of
megadrile earthworms, information that is currently scattered, outdated, or
otherwise unavailable.
Responding to the ‘Taxonomic Impediment’ and 'Biodiversity Crisis', this proposal
complements stated aims of groups such as CBD/GTI, Diversitas, GBIF, BioNET,
IUCN/SSG to provide nomenclators/lists such as Nomenclaror Zoologicus, ION,
Species 2000, Tree-of-Life and Taxonomicon.
This basic information offsets an initiative under auspices of ICZN to complete
“ZooBank” database by 2008. Currently incomplete, it has 7,585 Oligochaeta
taxa including megadrile and microdrile spp and genera (but many mistakes).
A longer-term objective is collaboration between earthworm workers to collate and
maintain a single unified database of distribution maps and eco-taxonomic data
for all 6,000 named megadriles [i.e. 5,500 names to December, 2000 listed on an
unpublished database by Dr Cs. Csuzdi (pers. comm.) + c. >400 subsequent new
names],
possibly
using
tools
facilitated
by
'Wikispecies'
[http://species.wikimedia.org/wiki/Main_Page - see next slide].
4b. 'Wikiworm' a universal matrix of
Earthworms x Region x Ecology
Introduction page to WIKISPECIES
4c. 'Wikiworm' a universal matrix of
Earthworms x Region x Ecology
Currently, ca. 3,600 taxa (ca. 65% of totals) are compiled on the YNU-COE website as a
series of discrete datasets for selected regions or natural taxonomic groups (current
version: bio-eco.eis.ynu.ac.jp/eng/database/earthworm/ links to Wikipedia).
Included are all 670 Holarctic LUMBRICIDAE and all 920 Oriental PHERETIMOIDS.
Most of central Africa is uncatalogued. Other omissions are the balance of the ca. 830 known + 150
new Latin America taxa, obtainable from ELAETAO (G. Brown, pers. comm. December, 2006).
5a Seeking agreement on main categories of
ecological strategies of earthworms
Lee (1959, 1985) working
with members of
several families in
NEW ZEALAND had ten
categories:


Litter species
Topsoil species
Subsoil species





Aquatic
Under logs and
stones
Under bark
In trees
(epiphytes)
etc.
Bouché (1971, 1977)
considering a few
Lumbricids in
FRANCE had:•Épigées
•Anéciques
(from Greek
“reaching up”)
•Endogées
5b Seeking agreement on main categories of
ecological strategies of earthworms
An intervening scheme by Gates (1961) classified lumbricids in Maine,
USA into ecological groups based on their seasonal activity, habitat
and diet:
1. Geophagous - feeding mainly on soil.
2. Limicolous (or limiphagous) - in mud or saturated soils.
3. Litter feeders - in leaf litter, compost or manure.
And numerous other ecological classifications and taxonomies have
been proposed.
5c Seeking agreement on main categories of
ecological strategies of earthworms
Sims & Gerard (1985; 1999: 29-30) and Coleman & Crossley (1996; 2003: 104, Tab.
4.6) list Lee's terms (with Bouche's equivalents appended). Whereas, Kladivko
(1997) followed just Lee (1985), Blakemore (2002) proposed to accept Lee
(1959) and Bouche (1977), for reasons of practicality and priority, thus:1. Litter (= Épigées) 2. Topsoil (=Endogées) 3. Subsoil (=Endogées) 4. Anéciques
Actually few species
are known to be Anecic,
examples are:
Lumbricus terrestris,
L. polyphemus
(Fitzinger, 1833), and
Okinawan Amynthas
yambaruensis (as
shown in the worm on
the top right).
5d Seeking agreement on main categories of
ecological strategies of earthworms
But really more categories are
possible as ...
EACH SPECIES IS UNIQUE
AND DIFFERENT IN IT’S:
Morphology
Ecology
Behaviour
Etc. etc.
*IT IS MOST IMPORTANT TO NAME THE SPECIES CORRECTLY & CONSISTENTLY*
6. A mystery worm from Tasmania
In 1997 some unique and unusual
specimens were unearthed in a
Tasmanian forest that seemed
intermediate between families, such
as the Criodrilidae and Lumbricidae.
Closest relationship is perhaps to
Criodrilus Hoffmeister, Helodrilus
Hoffmeister or, possibly, Drilocrius
Michaelsen, 1917.
The worm is figured (right), and it is
probably a lumbricid (non-native to
Tasmanian) apparently similar to
Dendrodrilus. But is it a new species,
or new genus? Any suggestions?
7a. Megascolex (Promegascolex) mekongianus Cognetti,
1922: its extent, ecology and allocation to Amynthas
(Oligochaeta: Megascolecidae) by Robert J. Blakemore1*, Csaba Csuzdi2, Masamichi T. Ito1, Nobuhiro Kaneko1,
Maurizio G. Paoletti3, Sergei E. Spiridonov4, Tomoko Uchida5 and Beverley D. Van Praagh6
The slender length and annulations of
the Mekong worm: measuring up to
2,900mm with more than 500 segments,
are near the maxima recorded for any
earthworm.
7b. Megascolex (Promegascolex) mekongianus Cognetti,
1922: its extent, ecology and allocation to Amynthas
(Oligochaeta: Megascolecidae) by Robert J. Blakemore1*, Csaba Csuzdi2, Masamichi T. Ito1, Nobuhiro Kaneko1,
Maurizio G. Paoletti3, Sergei E. Spiridonov4, Tomoko Uchida5 and Beverley D. Van Praagh6
It seems Cognetti (1922) miscounted
segments of his Megascolex
(Promegascolex) mekongianus and,
believing the gizzard in "7" was
intermediate between Megascolex, with
gizzard in 5, and Pheretima, with
gizzard after 7/8, he proposed the
subgenus Promegascolex. Next, Gates
(1934: 260) redescribed the immature,
poorly preserved and abnormal type as
Pheretima mekongiana. However, Sims
& Easton (1972: 223) listed it as species
incertae sedis, excluded it from their
Pheretima-group of genera and
postulated its gizzard was "clearly in
segment 5". [Cognetti’s figures on left].
7c. Megascolex (Promegascolex) mekongianus Cognetti,
1922: its extent, ecology and allocation to Amynthas
(Oligochaeta: Megascolecidae) by Robert J. Blakemore1*, Csaba Csuzdi2, Masamichi T. Ito1, Nobuhiro Kaneko1,
Maurizio G. Paoletti3, Sergei E. Spiridonov4, Tomoko Uchida5 and Beverley D. Van Praagh6
Recently collected material from the
Laos side of River Mekong is
described that complies with the
corrected type description allowing its
new designation as Amynthas
mekongianus (Cognetti, 1922) comb.
nov.
 New material figured (RJB).
Known Mekong distribution 
7d. Megascolex (Promegascolex) mekongianus Cognetti,
1922: its extent, ecology and allocation to Amynthas
(Oligochaeta: Megascolecidae) by Robert J. Blakemore1*, Csaba Csuzdi2, Masamichi T. Ito1, Nobuhiro Kaneko1,
Maurizio G. Paoletti3, Sergei E. Spiridonov4, Tomoko Uchida5 and Beverley D. Van Praagh6
Moreover, A. fluvialis (Gates, 1939) from the Mekong in Thailand is found to be a synonym,
although Metaphire fluvialoides (Huynh Thi Kim Hoi, 1998) comb. nov. from Central Highlands
of Vietnam remains separate on its eversible male pores (its original figures above).
Observations - Why are earthworms
important?
Because they are essential for NATURAL AGICULTURAL PRODUCTION and SOIL HEALTH.
And they may play a role in reducing Global Climate Change:
If just 10,000 medium-sized farms in the U.S. converted to organic production, they would store so much
soil carbon equivalent to taking 1,174,400 cars off the road.
A 23-year side-by-side comparison study conducted by the Rodale Institute showed the carbon (C) levels of
organic soils increased while there was little change in the non organic systems. The study also showed that
organic systems used 63% less energy than required by conventional farming.
Similar findings to Rodale Institute were found in 40year organic soils in UK (see next slide).
Rising levels of CO2 in the atmosphere from burning fossil fuels is the principal cause of Global Warming.
Plants absorb CO2 from the air and sequester it into the soil as organic C compounds.
Composts innoculate the soil with humus building microbes that convert these carbon compounds into humus
(dark organic matter that also stores water) and earthworms facilitate this process (e.g. Darwin, 1881).
So perhaps we should think in terms of Global Worming!?
Observation 1
1. “Haughley Experiment” - a 40-year Organic vs. Conventional farming (Balfour, 1943, 1977; Blakemore,
1981, 1996, 2000, 2005).
2. Abundance and biomass of earthworms in side by side sections of farm.
3. Earthworm behavioural soil/food choice trials (Blakemore, 1981, 2000) & summary result.
Observation 2
Earthworm effect on soils and plants demonstration (5
worms, 2 weeks).
Observation 3
The Earth’s surface is 70% water, but we depend for our survival on soil-based
production from the remaining 30% that is dry land for >99% of all our food (and
fibres) - and just 0.6% from the Oceans (FAO, 1991).
Observation 4
THANK YOU
I (RJB) thank the presenter (Cs. Csuzdi)
and organizers of IOTM3
 Enjoy the symposium
 Hope we can meet in the near future to
continue the discussion
Till then, please check my website... 
• Cheers!

Photo is Elephant seal pups @ Macquarie island. RJB 1997