Summary of IOTM3 (International Oligochaeta Taxonomy Meeting 3) Cyprus Presentations

1. Summary of IOTM3(International Oligochaeta Taxonomy Meeting 3) Cyprus Presentations by Rob Blakemore COE fellow, Soil Ecology Research Group,Yokohama National University

2. IOTM Presentions (Posters and Powerpoints) 1. COSMOPOLITAN EARTHWORMS. 2ndEdition (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!

3. 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.

4. 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)

5. 2b. Phylogeny of Acanthodrilidae, Octochaetidae, Exxidae, & Megascolecidae revisited. Simple phylogram above from basic weighted morphology data below– using PHY-FI (Fredslund, 2006)

6. 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 (type-specimens of) the type-species of the type-genera.

7. 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].

8. 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).

9. 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).

10. 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 DIPLOCARDI-INAE/-IDAE (type North American Diplocardia communis Garman, 1888) – that possibly merits separate family status. See REGULATION TABLE below:

11. 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.

12. 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.

13. 3c. Review of Oriental pheretimoid (Pheretima auct. family Megascolecidae) taxa Characters of Pheretima-group genera incl. Dendropheretima and Isarogoscolex from Philippines by James (2004)

14. 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 pseudo-penes (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.

15. 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].

16. 4b. 'Wikiworm' a universal matrix of Earthworms x Region x Ecology Introduction page to WIKISPECIES

17. 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).

18. 5a Seeking agreement on main categories of ecological strategies of earthworms Bouché (1971, 1977) considering a few Lumbricids in FRANCE had:- • Épigées • Anéciques (from Greek “reaching up”) • Endogées 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.

19. 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.

20. 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).

21. 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*

22. 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 Hoffmeisteror, 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?

23. 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.

24. 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].

25. 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 

26. 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 Metaphirefluvialoides(Huynh Thi Kim Hoi, 1998) comb. nov. from Central Highlands of Vietnam remains separate on its eversible male pores (its original figures above).

27. 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!?

28. 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.

29. Observation 2 Earthworm effect on soils and plants demonstration (5 worms, 2 weeks).

30. 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).

31. Observation 4

32. 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