WORMS love EARTH - earthworms & their soil

Unravelling some Kinki earthworms (Annelida: Oligochaeta: Megadrili: Megascolecidae) - Part II

2011-03-09T13:35:00.001-05:00

Robert J. Blakemore

Opusc. Zool. Budapest (2010) 41(2), 191–206

Abstract.

Metaphire tanbode sp. nov. is found in rice paddy in Kinki plain at Lake Biwa and Amynthas yamade sp. nov. is from Hira range to the West. M. tanbode belongs to the M. hilgendorfi / A. tokioensis species-complex, while montane A. yamade is comparable to both Amynthas aeruginosus-group and Duplodicodrilus schmardae-group. Genetic barcoding (mtDNA COI) via types is attempted. Taxonomic ‘housekeeping’ requires replacement of invalid homonyms: e.g. Pheretima montana Ishizuka, 1999 (non type-species P. montana Kinberg, 1867) is renamed Amynthas nonmontanus; others are A. nonsilvestris, A. noninvisus, A. nonmonticolus and A. nonsetosus, noms. et combs. novae. Thus Pheretima Kinberg, 1867 s. stricto remains unrecorded from Japan while prior Amynthas Kinberg, 1867, and its derivative Metaphire Sims & Easton, 1972, are abundant and diverse. Family and generic level definition and placement of Oriental pheretimoids are restated for the benefit of current workers and for novice field-ecologists. Surveys of below-ground biodiversity of rice paddy in Lake Biwa is compared to more natural habitats around Lake Pedder in Western Tasmanian Wilderness Area, and co-incidentally, both have 21 recorded earthworm species. Thus claims from various countries of less than six species per location are contraindicated by thorough eco-taxonomic methods yielding more representative results.

Impact of above- and below-ground invertebrates on temporal and spatial stability of grassland of different diversity

2011-03-09T11:46:00.000-05:00

Nico Eisenhauer, Alexandru Milcu, Eric Allan, Norma Nitschke, Christoph Scherber, Vicky Temperton, Alexandra Weigelt, Wolfgang W. Weisser and Stefan Scheu

Journal of Ecology (2011) 99, 572–582

Summary

1. Recent theoretical studies suggest that the stability of ecosystem processes is not governed by diversity per se, but by multitrophic interactions in complex communities. However, experimental evidence supporting this assumption is scarce.

2. We investigated the impact of plant diversity and the presence of above- and below-ground invertebrates on the stability of plant community productivity in space and time, as well as the interrelationship between both stability measures in experimental grassland communities.

3. We sampled above-ground plant biomass on subplots with manipulated above- and belowground invertebrate densities of a grassland biodiversity experiment (Jena Experiment) 1, 4 and 6 years after the establishment of the treatments to investigate temporal stability. Moreover, we harvested spatial replicates at the last sampling date to explore spatial stability.

4. The coefficient of variation of spatial and temporal replicates served as a proxy for ecosystem stability. Both spatial and temporal stability increased to a similar extent with plant diversity.Moreover, there was a positive correlation between spatial and temporal stability, and elevated plant density might be a crucial factor governing the stability of diverse plant communities.

5. Above-ground insects generally increased temporal stability, whereas impacts of both earthworms and above-ground insects depended on plant species richness and the presence of grasses. These results suggest that inconsistent results of previous studies on the diversity–stability relationship have in part been due to neglecting higher trophic-level interactions governing ecosystem stability.

6. Changes in plant species diversity in one trophic level are thus unlikely to mirror changes in multitrophic interrelationships. Our results suggest that both above- and below-ground invertebrates decouple the relationship between spatial and temporal stability of plant community productivity by differently affecting the homogenizing mechanisms of plants in diverse plant communities.

7. Synthesis. Species extinctions and accompanying changes in multitrophic interactions are likely to result not only in alterations in the magnitude of ecosystem functions but also in its variability complicating the assessment and prediction of consequences of current biodiversity loss.

Earthworm Invasion as the Driving Force Behind Plant Invasion and Community Change in Northeastern North American Forests

2011-03-09T11:34:00.002-05:00

VICTORIA A. NUZZO, JOHN C. MAERZ, AND BERND BLOSSEY

Conservation Biology (2009) 23(4) 966–974

Abstract

Identification of factors that drive changes in plant community structure and contribute to decline and endangerment of native plant species is essential to the development of appropriate management strategies. Introduced species are assumed to be driving causes of shifts in native plant communities, but unequivocal evidence supporting this view is frequently lacking. We measured native vegetation, non-native earthworm biomass, and leaf-litter volume in 15 forests in the presence and absence of 3 non-native plant species (Microstegium vimineum, Alliaria petiolata, Berberis thunbergii) to assess the general impact of non-native plant and earthworm invasions on native plant communities in northeastern United States. Non-native plant cover was positively correlated with total native plant cover and non-native earthworm biomass. Earthworm biomass was negatively associated with cover of native woody and most herbaceous plants and with litter volume. Graminoid cover was positively associated with non-native earthworm biomass and non-native plant cover. These earthworm-associated responses were detected at all sites despite differences in earthworm species and abundance, composition of the native plant community, identity of invasive plant species, and geographic region. These patterns suggest earthworm invasion, rather than non-native plant invasion, is the driving force behind changes in forest plant communities in northeastern North America, including declines n native plant species, and earthworm invasions appear to facilitate plant invasions in these forests. Thus, a focus on management of invasive plant species may be insufficient to protect northeastern forest understory species.

Dietary flexibility aids Asian earthworm invasion in North American forests

2011-03-09T11:30:00.000-05:00

WEIXIN ZHANG, PAUL F. HENDRIX, BRUCE A. SNYDER, MARIROSA MOLINA, JIANXIONG LI, XINGQUAN RAO, EVAN SIEMANN, AND SHENGLEI FU

Ecology (2010) 91(7) 2070–2079

Abstract

On a local scale, invasiveness of introduced species and invasibility of habitats together determine invasion success. A key issue in invasion ecology has been how to quantify the contribution of species invasiveness and habitat invasibility separately. Conventional approaches, such as comparing the differences in traits and/or impacts of species between native and/or invaded ranges, do not determine the extent to which the performance of invaders is due to either the effects of species traits or habitat characteristics. Here we explore

the interaction between two of the most widespread earthworm invaders in the world (Asian Amynthas agrestis and European Lumbricus rubellus) and study the effects of species invasiveness and habitat invasibility separately through an alternative approach of ‘‘third habitat’’ in Tennessee, USA. We propose that feeding behaviors of earthworms will be critical to invasion success because trophic ecology of invasive animals plays a key role in the invasion process. We found that (1) the biomass and isotopic abundances (d13C and d15N) of A. agrestis were not impacted by either direct effects of L. rubellus competition or indirect effects of L. rubellus-preconditioned habitat; (2) A. agrestis disrupted the relationship between L. rubellus and soil microorganisms and consequently hindered litter consumption by L. rubellus; and (3) compared to L. rubellus, A. agrestis shifted its diet more readily to consume more litter, more soil gram-positive (Gþ) bacteria (which may be important for litter digestion), and more nonmicrobial soil fauna when soil microorganisms were depleted. In conclusion, A. agrestis showed strong invasiveness through its dietary flexibility through diet shifting and superior feeding behavior and its indirectly negative effect of habitat invasibility on L. rubellus via changes in the soil microorganism community. In such context, our results expand on the resource fluctuation hypothesis and support the superior competitor hypothesis. This work presents additional approaches in invasion ecology, provides some new dimensions for further research, and contributes to a greater understanding of the importance of interactions between multiple invading species.

Re-integrating earthworm juveniles into soil biodiversity studies: species identification throughDNAbarcoding

2011-03-09T11:25:00.000-05:00

B. RICHARD, T. DECAE¨NS, R. ROUGERIE, S. W. JAMES, D. PORCO and P. D. N. HEBERT

Molecular Ecology Resources (2010) 10, 606–614

Abstract

Species identification of earthworms is usually achieved by careful observation of morphological features, often sexual characters only present in adult specimens. Consequently, juveniles or cocoons are often impossible to identify, creating a possible bias in studies that aim to document species richness and abundance.DNA barcoding, the use of a short standardized DNA fragment for species identification, is a promising approach for species discrimination. When a reference library is available, DNA-based identification is possible for all life stages. In this study, we show that DNA barcoding is an unrivaled tool for high volume identification of juvenile earthworms. To illustrate this advance,we generated DNA barcodes for specimens of Lumbricus collected from three temperate grasslands in western France. The analysis of genetic distances between individuals shows that juvenile sequences unequivocally match DNA barcode clusters of previously identified adult specimens, demonstrating the potential of DNA barcoding to provide exhaustive specimen identification for soil ecological research.

Cryptic speciation of hormogastrid earthworms revealed by mitochondrial and nuclear data

2011-03-09T11:21:00.001-05:00

Marta Novo, Ana Almodóvar, Rosa Fernández, Dolores Trigo, Darío J. Díaz Cosín

Molecular Phylogenetics and Evolution (2010) 56, 507–512

Abstract

Species delimitation of earthworms has been difficult to determine with certainty due to their structural simplicity. We sequenced fragments of COI, 16S, t-RNAs and 28S for 202 Hormogastridae individuals from the central Iberian Peninsula and three outgroup taxa. A morphological constancy was found but a high genetic diversity suggests the presence of five cryptic allopatric species. Results showed a pattern of isolation by distance and a positive but weak correlation between some soil properties (coarse sand and total loam content) and genetic distances, which indicates that these populations may have been shaped genetically but not morphologically, by the environment.

High mitochondrial DNA sequence diversity in the parthenogenetic earthworm Dendrobaena octaedra

2011-03-09T11:17:00.002-05:00

KE Knott and J Haimi

Heredity (2010) 105, 341–347

Abstract

Apomictic parthenogens are clonal organisms with limited genetic opportunity for increasing diversity beyond mutation. However, such species can be successful and have been shown to harbor more genetic diversity than might be expected. Here we surveyed diversity of the cytochrome oxidase subunit I gene from the mitochondrial genome of the earthworm Dendrobaena octaedra, an apomictic parthenogen. Diversity estimates made previously from allozyme markers for this species were high, but could have been affected by a detection bias, namely variable expression of alleles in the polyploid genome. We found similarly high mtDNA diversity over three localities in Finland, each represented by two sites; either with or without anthropogenic soil metal contamination. A single haplotype was most common over all sites, and over two thirds of all haplotypes sampled were similar to it in sequence (only varying by 1–3 substitutions). However, more divergent rare haplotypes were also found in one locality, Imatra, in southeast Finland close to the Karelian Isthmus. Metal contamination in the soil did not have a significant effect on genetic diversity, although metal exposure is known to be detrimental to the worms. There was no evidence for cryptic species within D. octaedra and phylogenetic analyses showed some structure of lineages that may have diverged in historical glacial refugia. Other mechanisms, such as rare genetic exchange with closely related species or high mutation and dispersal rates may explain high genetic diversity in D. octaedra.

DNA Barcoding Reveals Cryptic Diversity in Lumbricus terrestris L., 1758 (Clitellata): Resurrection of L. herculeus (Savigny, 1826)

2011-03-09T11:11:00.002-05:00

Samuel W. James, David Porco, Thibaud Decaens, Benoit Richard, Rodolphe Rougerie, Christer Erseus

PLoS ONE (2010) 5(12), e15629. doi:10.1371/journal.pone.0015629

Abstract

The widely studied and invasive earthworm, Lumbricus terrestris L., 1758 has been the subject of nomenclatural debate for many years. However these disputes were not based on suspicions of heterogeneity, but rather on the descriptions and nomenclatural acts associated with the species name. Large numbers of DNA barcode sequences of the cytochrome oxidase I obtained for nominal L. terrestris and six congeneric species reveal that there are two distinct lineages within nominal L. terrestris. One of those lineages contains the Swedish population from which the name-bearing specimen of L. terrestris was obtained. The other contains the population from which the syntype series of Enterion herculeum Savigny, 1826 was collected. In both cases modern and old representatives yielded barcode sequences allowing us to clearly establish that these are two distinct species, as different from one another as any other pair of congeners in our data set. The two are morphologically indistinguishable, except by overlapping size-related characters. We have designated a new neotype for L. terrestris. The newly designated neotype and a syntype of L. herculeus yielded DNA adequate for sequencing part of the cytochrome oxidase I gene (COI). The sequence data make possible the objective determination of the identities of earthworms morphologically identical to L. terrestris and L. herculeus, regardless of body size and segment number. Past work on nominal L. terrestris could have been on either or both species, although L. herculeus has yet to be found outside of Europe.

Phylogenetic analysis of New Zealand earthworms (Oligochaeta: Megascolecidae) reveals ancient clades and cryptic taxonomic diversity

2011-03-09T11:00:00.002-05:00

Thomas R. Buckley, Sam James, Julia Allwood, Scott Bartlam, Robyn Howitt, Diana Prada

Molecular Phylogenetics and Evolution (2011) 58, 85–96

Abstract

We have constructed the first ever phylogeny for the New Zealand earthworm fauna (Megascolecinae and Acanthodrilinae) including representatives from other major continental regions. Bayesian and maximum likelihood phylogenetic trees were constructed from 427 base pairs from the mitochondrial large subunit (16S) rRNA gene and 661 base pairs from the nuclear large subunit (28S) rRNA gene. Within the Acanthodrilinae we were able to identify a number of well-supported clades that were restricted to continental landmasses. Estimates of nodal support for these major clades were generally high, but relationships among clades were poorly resolved. The phylogenetic analyses revealed several independent lineages in New Zealand, some of which had a comparable phylogenetic depth to monophyletic groups sampled from Madagascar, Africa, North America and Australia. These results are consistent with at least some of these clades having inhabited New Zealand since rifting from Gondwana in the Late Cretaceous. Within the New Zealand Acanthodrilinae, major clades tended to be restricted to specific regions of New Zealand, with the central North Island and Cook Strait representing major biogeographic boundaries. Our field surveys of New Zealand and subsequent identification has also revealed extensive cryptic taxonomic diversity with approximately 48 new species sampled in addition to the 199 species recognized by previous authors. Our results indicate that further survey and taxonomic work is required to establish a foundation for future biogeographic and ecological research on this vitally important component of the New Zealand biota.

Prey choice by carabid beetles feeding on an earthworm community analysed using species- and lineage-specific PCR primers

2011-03-09T10:53:00.004-05:00

R. ANDREW KING, IAN P. VAUGHAN, JAMES R. BELL, DAVID A. BOHAN and WILLIAM O. C. SYMONDSON

Molecular Ecology (2010) 19, 1721–1732

Abstract

The carabid beetle Pterostichus melanarius is a major natural enemy of pests, such as aphids and slugs in agricultural systems. Earthworms are a dominant non-pest component of the diet of P. melanarius which help sustain the beetles during periods when the pest population is low or absent. In this study we wanted to test whether this predator exercises prey choice among different earthworm species or ecological groups. High levels of genetic diversity within morphological species of earthworm necessitated the development of primers that were specific not just to species but lineages and sublineages within species as well. Gut samples from beetles were analysed using multiplex-PCR and fluorescent-labelled primers. Calibratory feeding trials were undertaken to calculate median detection times for prey DNA following ingestion. Extensive testing demonstrated that the primers were species-specific, that detection periods were negatively related to amplicon size and that meal size had a highly significant effect on detection periods. Monte Carlo simulations showed that, in general, worms were being predated in proportion to their densities in the field with little evidence of prey choice, other than probable avoidance of the larger, deep-living species. There was no evidence that epigeic species were being taken preferentially in comparison with endogeic species. There was also no evidence that defensive secretions by Allolobophora chlorotica reduced predation pressure on this species by P. melanarius. We concluded that any management system that increases earthworm densities generally, regardless of component species, is likely to be optimal for increasing numbers of this beneficial beetle predator.

Aporrectodea trapezoides

2010-08-08T19:22:00.001-04:00

Apr 24, 2009. Bank of Tigris River, Diyarbakir, Turkey

Apr 15, 2010. JHU, Baltimore, MD, USA

Aporrectodea rosea

2010-08-08T19:13:00.000-04:00

Apr 22, 2010. SERC, MD, USA

Octolasion lacteum

2010-08-08T19:05:00.000-04:00

Apr 16, 2010. SERC, MD, USA

Nov 6, 2010. SERC, MD, USA

Octolasion cyaneum

2010-08-08T18:58:00.000-04:00

Apr 16, 2010. SERC, MD, USA

Nov 17, 2009. SERC, MD, USA

Eisenoides loennbergi

2010-08-08T18:51:00.002-04:00

Mar 18, 2010. SERC, MD, USA

Apr 16, 2010. SERC, MD, USA

Aporrectodea caliginosa

2010-08-08T18:41:00.002-04:00

Apr 16, 2010. SERC, MD, USA

Mar 18, 2010. SERC, MD, USA

Allolobophora chlorotica

2010-08-08T11:49:00.000-04:00

Mar 18, 2010. SERC, MD, USA

Lumbricus friendi

2010-08-08T11:30:00.001-04:00

Mar 18, 2010. SERC, MD, USA

Nov 11, 2009. SERC, MD, USA

Lumbricus rubellus

2010-08-08T11:25:00.002-04:00

Nov 6, 2009. SERC, MD, USA

Lumbricus terrestris

2010-08-08T11:17:00.001-04:00

Lumbricus terrestris. Mar 22, 2010. JHU, MD, USA

台灣已命名蚯蚓增加到79種

2010-08-08T00:49:00.001-04:00

台灣又發現了四種新種蚯蚓，蚯蚓物種數逼近80大關! 詳見由特有生物研究中心團隊發表於倫敦自然史博物館出版的Journal of Natural History的論文。

Four new species of Amynthas earthworms (Oligochaeta: Megascolecidae) from the Central Mountain Range of southern Taiwan

Authors

Chu-Fa Tsai, Huei-Ping Shen, Su-Chen Tsai

Journal

Journal of Natural History (2010) 44, 1251–1267

Abstract

Four new species of terrestrial earthworms belonging to the genus Amynthas (Megascolecidae: Oligochaeta) are described from the Central Mountain Range in southern Taiwan. Two octothecate species, Amynthas pavimentus sp. nov. and Amynthas biorbis sp. nov., were found at elevations of 2000–2700 m and 412–1000 m, respectively, on both east and west slopes. Amynthas pavimentus has patched genital papillae between setal line and intersegmental furrow in VIII–IX and XVII–XX. Amynthas biorbis has a pair of large papillae adjacent to male porophores in XVIII. The other two species are the octothecate Amynthas libratus sp. nov., found at 1000 m, and the sexthecate Amynthas hongyehensis sp. nov., found at 950–1000 m, on the eastern slope. Amynthas libratus is related to A. pavimentus but has horizontal rows of papillae along setal lines or intersegmental furrows in VII–IX and XVII–XX. Amynthas hongyehensis has two pairs of large, presetal papillae medial to male porophores in XVIII and XIX.

Rob's new paper

2010-08-08T00:34:00.002-04:00

[Robert J. Blakemore's comments on DNA barcoding.]

Neotypification of Drawida hattamimizu Hatai, 1930 (Annelida, Oligochaeta, Megadrili, Moniligastridae) as a model linking mtDNA (COI) sequences to an earthworm type, with a response to the ‘Can of Worms’ theory of cryptic species

Author

R.J. Blakemore, E.K. Kupriyanova, M.J. Grygier

Journal

ZooKeys 41: 1–29 (2010)

Abstract

A neotype is designated for the large and ecologically interesting species of Japanese earthworm, Drawida hattamimizu Hatai, 1930. Its morphological redescription is unambiguously combined with the neotype’s sequence of the Cytochrome Oxidase I (COI) locus of the mitochondrial DNA, the fi rst time an earthworm species’ type has been thus profi led. Probably it is an introduced exotic rather than a translocated native, with a patchy distribution that appears only partly defi ned in Japan where it is both a restricted and an endangered listed species. Brief comparison of sympatric Drawida japonica (Michaelsen, 1892) to the type-species Drawida barwelli (Beddard, 1886) – and this latter from Shiga appears as a new record for Japan – allows the diagnosis of Drawida Michaelsen, 1900 to be amended slightly. Th e contentious issue of molecular ‘cryptic species’ is queried in relation to the lack of molecular data from type-specimens, the unique name-bearing references employed in zoological nomenclature. Without such reference, neither eco-taxonomic nor genomic studies of earthworm taxa can progress. In this regard, questions are raised concerning the molecular identities and provisional divergences of cosmopolitan generotypes Allolobophora chlorotica chlorotica (Savigny, 1826), the Aporrectodea caliginosa (Savigny, 1826) species-complex sensu Blakemore (2002), and of ecotoxicological standard test-species icon Eisenia fetida fetida (Savigny, 1826). Resurrection of their respective synonyms is mooted. Resolution of relationships within and between earthworm genera and families without DNA testing of the representative type-species and type-genera is flagged as another crucial concern.

A paper in SSB

2010-08-08T00:07:00.006-04:00

都什麼年代了還有人在搞這種一顆16S rRNA tree一顆COI tree的遊戲，而且還拿不同科的蚯蚓來分析，跑出來連Lumbricidae是單系群都不支持，更要命的是竟然還登在SSB上，不知道有多少人要被這篇文章誤導了，真搞不懂審paper的人是怎麼審的。對DNA條碼有點研究的人都知道，COI提供的phylogeny不管支持度再怎麼高，都只能當做working hypothesis來看待。2003年以來，已有不少的研究說明了COI在蚯蚓中僅適用於研究近緣種，拿來分析一個科實在是...。另外文章裡對學名的處理亂七八糟，完全無視ICZN的規範。

Are the sexual, somatic and genetic characters enough to solve nomenclatural

problems in lumbricid taxonomy?

Journal

Soil Biology & Biochemistry 41 (2009) 2257–2271

Author

Marı´a Jesu´ s Iglesias Briones, Paloma Mora´n, David Posada

Abstract

Traditionally, phylogenetic relationships within Lumbricidae have been investigated by using morphological characters which show high variability between individuals. Furthermore, the lack of agreement in their ranking as diagnostic characters for taxonomic and phylogenetic purposes has led to situations in which the same species receives different names in different parts of the world or may be included in different genera depending on the classification system proposed. Although the recent use of molecular tools in metazoan phylogeny has questioned our traditional understanding of animal classification, these techniques have not been yet fully exploited in earthworm evolutionary biology.
Here we construct molecular phylogenies to contrast them with the traditional morphological classification within the Lumbricidae. We have put special emphasis on clarifying the phylogenetic relationships of those species commonly found in the European Atlantic area, trying to highlight some of the nomenclatural problems associated with certain species and to provide a better understanding of this group for earthworm ecologists.We obtained DNA sequences for two fragments of the mitochondrial COI and 16S genes for thirty-six earthworm taxa belonging to the family Lumbricidae and seven species belonging to six outgroup families according to Sims and Gerard (1999)’s terminology (Criodrilidae, Glossoscolecidae, Megascolecidae, Acanthodrilidae, Eudrilidae and Hormogastridae). We interpreted the results in the light of the available information on genital, somatic and genetic characters published over the last 100 years and performed a detailed anatomical study of certain species for which there are contradictory descriptions.

In agreement with previous results, our study concludes that the 16S rDNA and COI sequence fragments have a limited discriminatory value above the genus level. In general, the relationships suggested by 16S were more in agreement with morphology than COI but the two genes proved to be useful at the shallowest taxonomic levels. Accordingly, both Lumbricus and Octolasion genera were shown to form distinctive clades despite the distinct geographical distribution of some of their members (e.g. Lumbricus friendi versus Lumbricus terrestris). In contrast, in the case of the heterogeneous Dendrobaena, Aporrectodea and the so-called ‘‘catch-all genus Allolobophora’’, the phylogenetic resolution was very limited. In addition, both markers seem to be suitable in revealing ecological similarities within species complexes. This was particularly true in the case of the Octolasion species and those included in the Ap. caliginosa complex, which suggest that occupying different niches in the soil may act as an isolating mechanism of their populations and could lead to speciation.

Importantly, our results provide good support for removing Levinsen’s L. eiseni from the Lumbricus genus, although its taxonomic position is still problematic and further studies are required, using slower markers and by including a greater number of the forms of each genus in the analyses. Similarly, the anatomical, genital and genetic characters investigated here do not allow the full clarification of the taxonomical status of Dendrobaena/Eisenia hortensis and D./E. veneta; however, in order to maintain the nomenclature stability, we propose to retain them in Dendrobaena until further analyses are carried out on test type specimens (e.g. paratypes or topotypic material). Furthermore, more research is also needed to solve the great heterogeneity of the genus Dendrobaena.

Our review also highlights the need for more collaborative work between alpha taxonomists and molecular phylogeneticists to make any meaningful progress in earthworm classification, so that the material included in the phylogenetic trees is properly identified and the results interpreted adequately. Furthermore, in order to avoid further taxonomical ambiguities, soil ecologists are advised to provide the most accurate identification of the species used in their studies or at least a good description of the specimens used together with the geographical location and the soil characteristics where the material was collected. Preserving some material for further taxonomic and genetic reference is also advised for amending possible mistakes.

We finally conclude that anatomical or genetic information alone cannot define any lumbricid genus and that gathering morphological, biological, physiological, ecological and genetic evidence together provides the best tool for solving the current ‘‘unequal chaos’’ of lumbricid taxonomy and for full elucidation of phylogenetic relationships within the Lumbricidae.

Two new species and some new records from eastern Taiwan

2010-02-09T15:48:00.000-05:00

台灣又多了兩種蚯蚓了!

Earthworm fauna of Eastern Taiwan, with descriptions of two new species (Oligochaeta: Megascolecidae)

Journal
ZOOTAXA (2010) 2341, 52–68
Authors
YU-HSI WANG & HSI-TE SHIH

Abstract
The earthworm fauna of eastern Taiwan was studied, and two new species, viz. Amynthas shinjiroi sp. nov. and Pithemera tao sp. nov. are described. Other species from our collection included Dichogaster (Diplothecodrilus) saliens (Beddard, 1893), Amynthas aspergillum (Perrier, 1872), A. gracilis (Kinberg, 1867), A. aff. hupeiensis (Michaelsen, 1891), A. robustus (Perrier, 1872), A. aff. morrisi (Beddard, 1892), Pithemera bicincta (Perrier, 1875), P. lanyuensis Shen & Tsai, 2002, and Pontoscolex corethrurus (Müller, 1856). Pithemera lanyuensis was first collected on Lanyu (Orchid Island), but its distribution is now extended to the main island of Taiwan. The finding of the exotic Pontoscolex corethrurus in eastern Taiwan shows it has probably invaded the entire island of Taiwan, and its impacts upon the ecology and agriculture need to be studied in the future.

"Population Genetics" of Hormogaster elisae

2010-02-09T14:54:00.002-05:00

有不少人跟我討論過我們在投paper時，相對於來自拉丁語系的國家的作者，似乎比較吃虧。我們的英文是很爛，有時候比較複雜的東西不一定能簡單扼要的說清楚，但不見得會受到歧視。我自己從沒有因為說中文不受到重視或是被歧視，國外reviewer的意見其實頗為中懇，對我的幫助也很大。但我常看到的是一篇paper出現這樣的情形 : 明明就沒有達到這個期刊的標準卻被登出來了。有時候這會讓人很失望，進而影響投稿到這個期刊的意願......。

[82 Hormogaster elisae specimens were used to understand past demography of this species. However, at least 4 species are in the so called 82 "Hormogaster elisae" specimens. Thus, the connection between major haplotype groups, which actually are different species, in the TCS network seems meaningless to me: how could we connect two nodes with 66 pseudo-nodes (hypothesized haplotypes)? Further, the mismatch distributions were calculated using relatively few specimens when compared with other genetic studies. For instance, what could the mismatch distributions calculated using 9, 12, 12 and 11 specimens tell us about the populations in Anchuelo, Pardo, Sevilla N and Siguero, respectively? Could we say that some of them experienced range expansion? On the other hand, this is an interesting paper and is the first one that uses minimum spanning network-related approaches to study earthworms.]

High genetic divergence of hormogastrid earthworms (Annelida, Oligochaeta) in the central Iberian Peninsula: evolutionary and demographic implications

Journal
Zoologica Scripta (2009) 38, 537–552
Authors
MARTA NOVO, ANA ALMODO´ VAR & DARI´O J. DI ´ AZ-COSI´N
Abstract

Hormogastridae earthworms are highly important for the functioning of the Mediterranean soil system. However, little is known about the species distribution and genetic diversity of these soil invertebrates. In the present study, the genetic differentiation and gene flow were studied among populations of hormogastrids from the central Iberian Peninsula. A 648-bp portion of the mitochondrial cytochrome c oxidase I gene was sequenced for 82 individuals from 7 localities, resulting in the identification of 38 haplotypes exclusive to localities. All of the individuals were morphologically identified as Hormogaster elisae, but the high genetic divergence found among populations (up to 20.20%) suggests the occurrence of more than one cryptic species within this region. Further analysis of the phylogenetic relationships revealed six different evolutionary lineages coincident with geographical location, including the two nearest populations Molar and Reduena as one evolutionary unit. From these results, at least three new species could be inferred, in addition to the morphospecies H. elisae s.s. Partitioning of genetic variance among populations indicated that isolation by distance was the primary agent for differentiation of the investigated hormogastrid populations. Our data suggest that the evolutionary lineages for H. elisae s.l. originated between the late Miocene and the early Pleistocene, but that mtDNA genealogies coalesce on amore recent scale of a few thousand years.

Earthworm Fauna of Taiwan 一書重要訊息

2009-12-31T08:05:00.002-05:00

二手書店販售的Earthworm Fauna of Taiwan (台灣蚯蚓誌) 多為印刷廠流出之瑕疵品
印刷品質不佳 請勿購買
請參照內文66頁或其他頁面的印刷品質辨識真正由台大出版中心出版的台灣蚯蚓誌

下圖為瑕疵品, 照片黑, 模糊不清

下圖為正版, 顏色淡, 圖像銳利, 對比強且清楚

phylogeography of a Japanese earthworm

2009-11-12T17:27:00.004-05:00

台灣蚯蚓COI基因超過10%的種內變異是個很有趣的現象。有些國外的學者認為這裡面應該有隱蔽種的存在。也許是研究做得還不夠多，國外的物種出現這麼高的遺傳變異的例子很少，但這在台灣的特有種蚯蚓身上實在是非常普遍，而偏偏台灣特有種的比例又不低，形態變異有大有小，時常造成分類上的困擾。Minamiya等人的這個研究指出了在Metaphire sieboldi內有接近16%的遺傳變異，與台灣的情形相當，對蚯蚓種內的高遺傳變異提供了支持。

[An interesting paper about a beautiful Japanese gigantic earthworm. The results show high intraspecific variation of up to 15.9% for mitochondrial COI gene, which is equivalent to those found previously in many species in Taiwan. This pattern suggests that high intraspecific genetic variation may be a common phenomenon in earthworms. The authors also used molecular clock of 4.8% per million years for COI estimated by Chang et al. (2008) to calculate the divergence time of populations, and the results showed consistency between geological data and clade distribution, providing evidence for this slightly high molecular clock.]

A phylogeographic study of the Japanese earthworm, Metaphire sieboldi (Horst, 1883) (Oligochaeta: Megascolecidae): Inferences from mitochondrial DNA sequences

Journal
European Journal of Soil Biology 45 (2009) 423–430
Authors
Yukio Minamiya, Jun Yokoyama, Tatsuya Fukuda
Abstract

To clarify the historical phylogeography of a Japanese earthworm, nucleotide sequence variations of Metaphaire sieboldi were investigated with respect to its geographic distribution. Sequencing of mitochondrial DNA (mtDNA), including the cytochrome oxidase subunit I (COI) region and the 16S rDNA region, of 72 samples including 4 outgroup taxa, and phylogenetic analyses using neighbour-joining (NJ) and Bayesian inference (BI) methods indicated that M. sieboldi forms a monophyletic group. This monophyletic group was divided into seven subgroups; most subgroups comprised individuals from several areas, except for southern Shikoku and southern Kyushu. The phylogenetic tree demonstrated that the Shikoku and Kinki populations were paraphyletic at the basal node, and individuals from Kyushu and western Shikoku were positioned at the advanced group. Estimated phylogeographic events based on mtDNA analyses include (1) M. sieboldi originated in Shikoku and/or Kinki, (2) individuals from Chugoku and Kyushu have a common ancestor from Shikoku, and (3) individuals from western Shikoku originated from Kyushu progenitors (about 0.4–0.5 mya) after separation by rising sea levels.

Draft genome of Lumbricus rubellus

2009-11-08T20:43:00.011-05:00

這篇文章在Nature有專文介紹 (詳見摘要之後)。據說今年 (2009)我們會看到Lumbricus rubellus的draft genome，而如今就快年底了。其實就算拖到明年也沒關係啦! 這是件令人興奮的大代誌呢! 請大家拭目以待吧!

[This is a very good review. Let's look forward to the draft genome of Lumbricus rubellus!!]

Earthworm genomes, genes and proteins: the (re)discovery of Darwin’s worms

Journal
Proc. R. Soc. B (2009) 276, 789–797
Authors
S. R. Stürzenbaum, J. Andre, P. Kille, A. J. Morgan
Abstract

Small incremental biological change, winnowed by natural selection over geological time scales to produce large consequences, was Darwin’s singular insight that revolutionized the life sciences. His publications after 1859, including the ‘earthworm book’, were all written to amplify and support the evolutionary theory presented in the Origin. Darwin was unable to provide a physical basis for the inheritance of favoured traits because of the absence of genetic knowledge that much later led to the ‘modern synthesis’. Mistaken though he was in advocating systemic ‘gemmules’ as agents of inheritance, Darwin was perceptive in seeking to underpin his core vision with concrete factors that both determine the nature of a trait in one generation and convey it to subsequent generations. This brief review evaluates the molecular genetic literature on earthworms published during the last decade, and casts light on the specific aspects of earthworm evolutionary biology that more or less engaged Darwin: (i) biogeography, (ii) species diversity, (iii) local adaptations and (iv) sensitivity. We predict that the current understanding will deepen with the announcement of a draft earthworm genome in Darwin’s bicentenary year, 2009. Subsequently, the earthworm may be elevated from the status of a soil sentinel to that elusive entity, an ecologically relevant genetic model organism.

News about this article in Nature
The year of the worm? doi:10.1038/news.2008.1309
(first paragraph only)

The earthworm is finally inching its way into the genomic age.
Interest in the humble earthworm seems to have seen a revival of late. Stephen Stürzenbaum from Kings College London and his team are hoping to unveil its draft genome next year, and Stürzenbaum tells Nature News why he has written a review on the topic for Proceedings of the Royal Society B.

Earthworm invasion in the Annual Review of Ecology, Evolution, and Systematics

2009-11-08T15:18:00.008-05:00

這幾年蚯蚓入侵的問題漸漸受到重視。來自南美洲的黃頸蜷蚓在全球熱帶與亞熱帶地區橫行；在美國北部與加拿大，原本被更新世冰河覆蓋，幾千年來在沒有蚯蚓的環境下演化而成的森林生態系（hardwood forest）被來自歐洲的蚯蚓大舉入侵，造成地表植被巨大的改變。這篇由Paul Hendrix、Mac A. Callaham, Jr.、Sam James等知名蚯蚓專家共同撰寫的論文揭露了這個問題的嚴重性。其中的共同作者Ching-Yu Huang (黃青萸) 還是成功大學與University of Georgia畢業的台灣人喔！

[The problem of invasive earthworms has become an issue that more and more people are aware of. Its impacts are especially serious in the tropical and subtropical regions that are invaded by Pontoscolex corethrurus and in temperate regions that were previously devoid of earthworms due to galciation.]

Pandora’s Box Contained Bait: The Global Problem of Introduced Earthworms

Journal
Annu. Rev. Ecol. Evol. Syst. (2008) 39: 593–613
Authors
Paul F. Hendrix, Mac A. Callaham, Jr., John M. Drake, Ching-Yu Huang, Sam W. James, Bruce A. Snyder, Weixin Zhang
Abstract

Introduced exotic earthworms now occur in every biogeographic region in all but the driest or coldest habitat types on Earth. The global distribution of a few species (e.g., Pontoscolex corethrurus) was noted by early naturalists, but now approximately 120 such peregrine species are recognized to be widespread from regional to global scales, mainly via human activities. Species adapted to human transport and to colonization of disturbed habitats are most widespread and are the principal invasive species. We identify a number of endogenous and exogenous factors that may contribute to the successful establishment and spread of peregrine species. Quantification of these factors may help to determine why certain species become invasive while others do not. Recent advances in theory and modeling of biological invasions and in molecular techniques should prove fruitful in improving our understanding of invasive earthworms, as well as in predicting their impacts on ecosystems.

Earthworms know if the partner is virgin or not!

2009-11-08T14:55:00.006-05:00

許多無脊椎動物的雄性都有增加自己的精子在male-male competition下優勢的機制，例如產生交配栓，或是增加精子的排放量。這篇文章顯示蚯蚓這種雌雄同體、異體受精的動物也有類似的情形，牠們在發現交配的對象不是「處女」時，會將精子的射出量提高到三倍，藉以增加自己精子成功受精的數量。

[An earthworm knows if "his" mating partner is virgin or not!]

Brainless but not clueless: earthworms boost their ejaculates when they detect fecund non-virgin partners

Journal
Proc. R. Soc. B (2008) 275 (1638): 1067-1072
Authors
Alberto Velando, Julio Eiroa and Jorge Domı´nguez
Abstract

In many animals in which females store sperm, males may detect female mating status and, in order to outcompete rival sperm, increase ejaculate size when copulating with non-virgin females. Although most studies have been restricted to organisms with separate sexes, theoretical models suggest that sperm competition should also be an important selective agent shaping life-history traits in simultaneous hermaphrodites. Nevertheless, the empirical support for ejaculate adjustment in a mating opportunity is scarce in hermaphrodites. In the present study, we performed a double-mating experiment to determine whether earthworms (Eisenia andrei ) detect the mating status of their partners and whether they respond by adjusting their ejaculate. We found that earthworms triplicated the donated sperm when mating with a non-virgin mate. Moreover, such increases were greater when the worms were mated with larger (more fecund) partners, indicating that earthworms perform a fine-tune control of ejaculate volume. The results of the present study suggest that, under high intensity of sperm competition, partner evaluation is subject to intense selection in hermaphrodite animals, and donors are selective about to whom they donate how much sperm.

Earthworms affecting seedlings

2009-11-08T01:12:00.006-05:00

蚯蚓能影響種子萌發的成功率，長久下來會不會對木本植物的群聚產生影響呢？這其實是個很嚴重的問題。這幾年採集蚯蚓的記憶裡，許多次生林外來種蚯蚓數量眾多的地方，地表滿滿都是糞土，除了高大的木本植物以外一點其它的植被也沒有，如果林間出現空隙，但種子一發芽就被蚯蚓吃掉了，或是蚯蚓導致萌發率下降，時間久了很難想像會發生什麼事情。

Exotic Ecosystem Engineers Change the Emergence of Plants from the Seed Bank of a Deciduous Forest

Journal
Ecosystems (2009) 12: 1008–1016
Authors
Nico Eisenhauer, Daniela Straube, Edward A. Johnson, Dennis Parkinson, Stefan Scheu
Abstract

The anthropogenic spread of exotic ecosystem engineers profoundly impacts native ecosystems. Exotic earthworms were shown to alter plant community composition of the understory of deciduous forests previously devoid of earthworms. We investigated the effect of two exotic earthworm species (Lumbricus terrestris L. and Octolasion tyrtaeum Savigny) belonging to different ecological groups (anecic and endogeic) on the emergence of plants from the seed bank of a northern North American deciduous forest using the seedling emergence method. We hypothesized that (1) exotic earthworms change the seedling emergence from the plant seed bank, (2) L. terrestris increases the emergence of plant seedlings of the deeper soil layer but decreases that of the upper soil layer due to plant seed burial, and (3) O. tyrtaeum decreases plant seedling emergence due the damage of plant seeds. Indeed, exotic earthworms altered the emergence of plant seedlings from the seed bank and the functional composition of the established plant seedlings. Surprisingly, although L. terrestris only marginally affected seedling emergence, O. tyrtaeum changed the emergence of native plant species from the seed bank considerably. In particular, the number of emerging grass and herb seedlings were increased in the presence of O. tyrtaeum in both soil layers. Moreover, the impacts of earthworms depended on the identity of plant functional groups; herb species benefited, whereas legumes suffered from the presence of exotic earthworms. The results highlight the strong effect of invasive belowground ecosystem engineers on aboveground ecosystem characteristics and suggest fundamental changes of ecosystems by human spread earthworm species.

What is earthworm grunting? Why does it work?

2009-11-06T11:53:00.012-05:00

Earthworm grunting是插一根鐵棍在地上，再用一根木頭用力摩擦鐵棍的上緣發出震動與聲音，使蚯蚓從土裡面跑出來。這個方法不是對每一種蚯蚓都有用，成功率在不同地方也不一樣。這個研究顯示grunting這個動作讓蚯蚓以為「鼴鼠來了」，蚯蚓為了逃跑而爬出地表。

[A new hypothesis that explains why earthworms go out of the soil: they are escaping moles!!]

Worm Grunting, Fiddling, and Charming—Humans Unknowingly Mimic a Predator to Harvest Bait

Journal
PLoS ONE (2008) 3(10): e3472
Author
Kenneth C. Catania
Abstract

Background: For generations many families in and around Florida’s Apalachicola National Forest have supported themselves by collecting the large endemic earthworms (Diplocardia mississippiensis). This is accomplished by vibrating a wooden stake driven into the soil, a practice called ‘‘worm grunting’’. In response to the vibrations, worms emerge to the surface where thousands can be gathered in a few hours. Why do these earthworms suddenly exit their burrows in response to vibrations, exposing themselves to predation?

Principal Findings: Here it is shown that a population of eastern American moles (Scalopus aquaticus) inhabits the area where worms are collected and that earthworms have a pronounced escape response from moles consisting of rapidly exiting their burrows to flee across the soil surface. Recordings of vibrations generated by bait collectors and moles suggest that ‘‘worm grunters’’ unknowingly mimic digging moles. An alternative possibility, that worms interpret vibrations as rainand surface to avoid drowning is not supported.

Conclusions: Previous investigations have revealed that both wood turtles and herring gulls vibrate the ground to elicit earthworm escapes, indicating that a range of predators may exploit the predator-prey relationship between earthworms and moles. In addition to revealing a novel escape response that may be widespread among soil fauna, the results show that humans have played the role of ‘‘rare predators’’ in exploiting the consequences of a sensory arms race.

Use DNA barcodes to identify earthworms: a real case

2009-11-02T23:37:00.008-05:00

Eisenia fetida與E. andrei在形態上不易區分。繼2005年Perez-Losada發表於Pedobiologia的paper之後，現在有人實際應用DNA條碼來做這兩種蚯蚓的物種鑑定了。
Usefulness of DNA Barcoding in Ecotoxicological Investigations: Resolving Taxonomic Uncertainties Using Eisenia Malm 1877 as an Example

Journal
Bull Environ Contam Toxicol (2009) 82: 261–264
Authors
P. Voua Otomo, B. Jansen van Vuuren, S. A. Reinecke
Abstract

Standard test species may differ in their response to toxicants. Accurate identification of test organisms is therefore of critical importance in correctly interpreting data generated from laboratory assays. This is not always possible when species are morphologically similar or where the taxonomy of the group has recently been revised. A case in hand concerns Eisenia sp. Based on recent genetic evidence two species, Eisenia andrei and Eisenia fetida, which were previously considered a single species, are currently recognized. In these instances, DNA barcoding, demonstrated and discussed herein, provides a method to accurately identify test organisms.

A mini-review on earthworm molecular ecology. Looking forward for more research papers!!

2009-11-02T23:21:00.015-05:00

關於蚯蚓分子生態的一篇review 。蚯蚓生態、演化與分類這個領域對DNA序列分析的應用一直處於相對落後的狀態，文獻無論在質與量都遠遜於同屬環帶綱的蛭類與水棲寡毛類。這兩年狀況稍微好了些，在Molecular Phylogenetics & Evolution或Molecular Ecology兩個期刊上都有相關的文章發表，期待看到更多振奮人心的作品。

Perspectives on the application of molecular genetics to earthworm ecology

Journal
Pedobiologia 52 (2009) 191-205
Author
Lise Dupont
Summary

Despite the great interest in earthworms by ecologists and the growing use of molecular data in ecology over the past 20 years, molecular techniques have been only rarely applied to earthworm research. In the family Lumbricidae, problematic taxonomy and the variable level of ploidy within and among species are significant barriers to research on population genetics. Until 1998, allozymes were the exclusive molecular markers for earthworm population studies but they were found to have substantial limitations. The current development of more promising markers (e.g. microsatellites) provides exciting prospects for the investigation of recent population genetic events and reproductive strategies of lumbricid species. Lately, some EST projects on earthworms that will facilitate the investigation of gene expression through the development of microarrays have been undertaken. In this review, emphasis is on the different molecular techniques currently available for addressing lumbricid ecological issues. Recently developed methods in population genetics and ecological genomics are also discussed. The rapid progress in DNA sequencing technologies will facilitate both the discovery of new genetic markers, and the use of genetic and genomic tools for earthworm ecology.

DNA barcoding of earthworms

2009-11-02T23:12:00.005-05:00

蚯蚓的DNA條碼

Identifying earthworms through DNA barcodes: Pitfalls and promise

Journal
Pedobiologia 52 (2009) 171—180
Authors
Chih-Han Chang, Rodolphe Rougerie, Jiun-Hong Chen
Summary

This paper re-evaluates the use of DNA barcodes in earthworm species identification by re-analyzing sequence data for the mitochondrial cytochrome c oxidase subunit 1 (COI) gene. This re-analysis unveiled evidenced taxonomic inconsistencies which significantly affect data interpretation. When considering synonymy and misidentification in published records, our results do reveal that there is no shared COI haplotypes between morphologically distinct species and that interspecific divergence is much higher than intraspecific variation in most cases, with interspecific and intraspecific distances averaging 18.7% and 1.3% respectively. However, a few earthworm species endemic to Taiwan have deep intraspecific divergences which may represent potential cases of cryptic diversity, although incomplete lineage sorting cannot be ruled out without further study. We recognize the potential of DNA barcoding for earthworm taxonomy, but identify several issues regarding the evolution of the COI gene in these organisms which remain to be further elucidated.

Are Aporrectodea caliginosa & A. trepezoides different species?

2009-11-02T22:43:00.008-05:00

第二篇發表於Molecular Phylogenetics & Evolution上的蚯蚓paper。看來Aporrectodea caliginosa、A. trapezoides與A. tuberculata真的是三個不同的物種呢！

[The second earthworm paper in MPE. It provides DNA evidence that A. caliginosa, A. tuberculata and A. trapezoides are distinct species. ]

Phylogenetic assessment of the earthworm Aporrectodea caliginosa species complex (Oligochaeta: Lumbricidae) based on mitochondrial and nuclear DNA sequences

Journal
Molecular Phylogenetics and Evolution 52 (2009) 293–302
Authors
Marcos Pérez-Losada, Maigualida Ricoy, Jonathon C. Marshall, Jorge Domínguez
Abstract

The Aporrectodea caliginosa species complex includes the most abundant earthworms in grasslands and agricultural ecosystems of the Paleartic region. Historically this complex consisted of the following taxa: A. caliginosa s.s. Savigny, 1826, A. trapezoides Dugés (1828), A. tuberculata (Eisen, 1874), and A. nocturna Evans (1946). These four taxa are morphologically very similar and difficult to differentiate because of their morphological variability. Consequently, their taxonomic status and their phylogeneticrelationships have been a matter of discussion for more than a century. To study these questions, we sequenced the COII (686 bp), 12S (362 bp), 16S (1200 bp), ND1 (917 bp), and tRNAsAsn-Asp-Val-Leu-Ala-Ser-Leu (402 bp) mitochondrial and 28S (809 bp) nuclear gene regions for 85 European earthworms from 27 different localities belonging to the A. caliginosa species complex and four outgroup taxa. DNA sequences were analyzed using maximum parsimony, maximum likelihood, and Bayesian approaches of phylogenetic inference. The resulting trees were combined with morphological, ecological, and genomic evidence to test species boundaries (i.e., integrative approach). Our molecular analyses showed that A. caliginosa s.s. and A. tuberculata form a sister clade to A. trapezoides, A. longa, and A. nocturna, which indicates that A. longa is part of the A. caliginosa species complex. We confirm the species status of all these taxa and identify two hitherto unrecognized Aporrectodea species in Corsica (France). Moreover our analyses also showed the presence of highly divergent lineages within A. caliginosa, A. trapezoides, and A. longa, suggesting the existence of cryptic diversity within these taxa.

The first earthworm paper in Molecular Phylogenetics & Evolution

2009-10-23T14:57:00.006-04:00

一直沒有辦法理解為什麼研究蚯蚓分類的人很晚才開始應用DNA序列資料。這個說法也許不是很對，因為2002、2003兩年都有蚯蚓molecular systematics相關的文章發表，這些文章的作者包含了鼎鼎大名的Victor Pop, Sam James, BGM Jamieson等人，但相對於90年代末期就已經蓬勃發展的其它無脊椎動物領域而言，確實是晚了一些。02、03年的兩篇文章試圖處理較高階層的系統分類問題，但蚯蚓算是個較難處理的類群，這兩篇paper基本上都不算是非常成功。第一篇發表在Molecular Phylogenetics & Evolution的paper遲至2008年才出現，探討的是台灣蚯蚓種群內的分類、生物地理與種化，相對於高階的系統分類研究，難度沒那麼高，重點是取樣與分析是否合理。福爾摩沙腔環蚓種群是台灣特有的大型蚯蚓，包含11個物種，在台灣島快速隆起期間( 5~2.5百萬年前)，因新形成的山脈河川造成地理隔離而種化。

[This is the first paper focusing on earthworms in Molecular Phylogenetics & Evolution. It discusses speciation of the Metaphire formosae species group as a consequence of orogenesis of the Taiwan Island 5~2.5 million years ago.]

Molecular systematics and phylogeography of the gigantic earthwormsof the Metaphire formosae species group (Clitellata, Megascolecidae)

Journal
Molecular Phylogenetics and Evolution 49 (2008) 958–968
Authors
Chih-Han Chang, Si-Min Lin, Jiun-Hong Chen
Abstract

The earthworms of the Metaphire formosae species group distributed in Taiwan are members of the Pheretima complex within the Megascolecidae. In this study, the systematics and phylogeography of this species group were investigated using DNA sequences of mitochondrial cytochrome c oxidase subunit I (COI), 16S ribosomal (r)RNA, and NADH dehydrogenase subunit 1 (ND1). The results indicated that the 13 taxa of the M. formosae species group form a clade, including a cryptic species discovered in this study. In addition, Metaphire hengchunensis (James et al., 2005) should be regarded as a subspecies of Metaphire paiwanna Tsai et al., 2000, and Metaphire bununa glareosa Tsai et al., 2000 should be elevated to specific status. Phylogeographical inferences showed that allopatric speciation occurred in this species group during the rapid uplift of the main island of Taiwan between 5.0 and 2.5 million years ago. Our analysis exposes non-monophyly within each of the genera Amynthas and Metaphire, and more generally within the Pheretima complex. Further revisions of this speciose complex are urgently needed.

Revisiting Allolobophora chlorotica

2009-10-22T13:31:00.008-04:00

在King等人用DNA序列證明Allolobophora chlorotica包含不只一個物種，且綠色跟粉紅色的個體是不同物種後，cross breeding的實驗亦提供了支持的證據。

[A related paper to King et al. 2008 in Molecular Ecology. The pink form of Allolobophora chlorotica is at least actually a different species, and is now called Allolobophora virescens (Savigny, 1826).]

Allolobophora chlorotica (Savigny, 1826): Evidence for classification as two separate species

Journal
Pedobiologia 52, 81—84 (2008)

Authors
Christopher N. Lowe, Kevin R. Butt

Summary
Breeding experiments have questioned the status of the green (g) and pink (p) colour morphs of Allolobophora chlorotica. Should they become regarded as separate species, the name Allolobophora virescens (Savigny, 1826) is available for the pink form of this earthworm.
Breeding experiments were established to investigate the status of this species utilising previously unmated laboratory-reared adults. Pairs were set up for 10 days to facilitate mating, separated and then maintained in isolation for 28 days to obtain individual reproduction results. Cocoon production and viability were first determined from intra-morphic (pXp and gXg) and inter-morphic (pXg) pairings. All pXp and gXg pairings bred true with mean cocoon production in gXg (4.6 cocoons/worm/28 days) significantly greater (P<0.05) than in pXp (3.2 cocoons/worm/28 days). In pXg pairings, both pink and green individuals produced cocoons (3.9 and 5.9 cocoons/worm/28 days, respectively). However here, viability of cocoons produced by g was very low at 6%, compared with 59% for p.
In a second experiment, previously unmated offspring from inter-morphic pairings (pg) were paired with like individuals (pgXpg) and also paired with pure bred pink (p1) and green (g1) individuals (p1Xpg and g1Xpg, respectively). All crosses produced cocoons. All cocoons produced by pgXpg crosses were non-viable. In p1Xpg and g1Xpg crosses the pg parent produced viable cocoons (64% and 79% viability, respectively) but cocoons produced by p1 and by g1 parents from these matings were 100% non-viable.
It is suggested that severely restricted viability of cocoons produced by the green morph in pXg pairings provides evidence for genetic divergence of the 2 morphs. Furthermore, back-crossing of hybrids (pg) with pure bred morphs has indicated that hybrids are male sterile.

Earthworm ID issue and DNA barcoding

2009-10-22T13:09:00.011-04:00

蚯蚓的隱蔽種是個相當令人困擾的問題。這篇文章中提到的物種都是廣泛分佈在歐洲與北美洲的物種。除了這篇文章中提到的物種之外，鼎鼎大名的night crawler (Lumbricus terrestris)也有類似的問題。一直到2007年左右蚯蚓分類學者才意識到這個問題的嚴重性，並試圖做分類上的釐清。

[The second paper focusing on earthworms in MOLECULAR ECOLOGY. Cryptic species in earthworms have been recognized as a serious problem in earthworm studies. We now know that the worms called Allolobophora chlorotica, Aporrectodea longa, Aporrectodea rosea and Lumbricus rubellus might each be composed of more than a single species. Indeed, the night crawler Lumbricus terrestris, which is not addressed in this paper, has the same problem.]

Opening a can of worms: unprecedented sympatric cryptic diversity within British lumbricid earthworms

Journal
Molecular Ecology 17, 4684–4698 (2008)
Authors
R. ANDREW KING, AMY L. TIBBLE, WILLIAM O. C. SYMONDSON
Abstract

Earthworms play a major role in many aspects of soil fertility, food web ecology and ecosystem functioning, and hence are frequently the subjects of, for example, ecological and toxicological research. Our aim was to examine the genetic structure of common earthworm species, to identify cryptic lineages or species that may be distinct ecotypes or biotypes (and hence confound current research based upon morphotypes) and to try toexplain the massive cryptic diversity that eventually emerged. We demonstrated that species such as Allolobophora chlorotica, Aporrectodea longa, Aporrectodea rosea and Lumbricus rubellus all comprise highly divergent lineages with species-level divergence at the mitochondrial cytochrome oxidase I (COI) gene. In Allo. chlorotica alone, we found 55 haplotypes for COI, with 35 of these being found in pink and 20 in green morph worms. There were no cases of the two colour morphs sharing COI haplotypes. Phylogenetic analyses of mitochondrial COI and 16S genes showed the presence of five highly divergent lineages, suggesting the presence of multiple cryptic species within Allo. chlorotica. There was no clear geographical pattern to lineage distribution and many populations were polymorphic for both mitochondrial DNA lineage and colour morph. Amplified fragment length polymorphism results, based on two primer combinations, were broadly congruent with mitochondrial DNA results with one significant exception. Despite showing over 14% divergence at COI, amplified fragment length polymorphism markers showed that the two green morph lineages may be interbreeding and therefore represent a single taxon. The cryptic diversity revealed by these results has profound consequences for all areas of earthworm research.

The first earthworm paper in MOLECULAR ECOLOGY

2009-10-21T22:15:00.013-04:00

這是第一篇以DNA序列資料討論蚯蚓族群遺傳的文章。美國北部五大湖區域是更新世冰河覆蓋的區域，10000年前冰河退去之前沒有任何蚯蚓，而現在生存在那個區域的蚯蚓都是外來種。牠們嚴重地影響了該區域森林(hardwood forest)下層與底層的植群結構並大大降低了植物的biomass，也對種子萌發產生不利的影響。

[I was very excited when seeing this paper last year. Someone should be doing this kind of work, and finally, there is !! ]

Genetic structure of invasive earthworms Dendrobaena octaedra in the boreal forest of Alberta: insights into introduction mechanisms

Journal

Molecular Ecology (2008) 17, 1189–1197

Authors

ERIN K. CAMERON, ERIN M. BAYNE, DAVID W. COLTMAN

Abstract

Population genetic studies can help to determine whether invasive species are established via single vs. multiple introduction events and also to distinguish among various colonization scenarios. We used this approach to investigate the introduction of Dendrobaena octaedra, a non-native earthworm species, to the boreal forest of northern Alberta. The spread of non-native earthworms in forested systems is not well understood, although bait abandonment and vehicular transport are believed to be important. Mitochondrial DNA sequencing revealed that multiple introductions of this species have occurred in northern Alberta, although individual populations may have been established by either single or multiple invaders introduced on one or more occasions. There was no relationship between genetic distances and either geographical distances or distances along road networks, suggesting that human-mediated jump dispersal is more common than diffusive spread via road networks or via active dispersal. As well, genetic diversity was significantly greater at boat launches than roads, indicating that multiple introductions may be more likely to occur at those locations. Focusing management efforts on areas where multiple introductions are likely to occur may help to reduce invasive species’ potential for adaptive evolution and subsequent rapid spread.

Amynthas robustus 壯偉遠環蚓

2009-10-20T19:50:00.008-04:00

Main campus of NTU, Taipei, Taiwan, Feb 6, 2009.

Huben (湖本), Taiwan, July 18, 2007.

Amynthas hilgendorfi

2009-10-20T19:44:00.006-04:00

Plummers Island, MD, USA, Oct. 3, 2009.

Amynthas aspergillum 參狀遠環蚓

2009-10-20T19:32:00.006-04:00

Main campus of NTU, Taipei, Taiwan, Feb 6, 2009

Earthworm Fauna of Taiwan

2009-10-20T14:29:00.002-04:00

Published on Nov. 20, 2009.

Earthworm Fauna of Taiwan: Metaphire formosae species group

2009-10-20T14:21:00.003-04:00

An "old" book published last year.

Amynthas hupeiensis 湖北遠環蚓

2009-10-20T12:47:00.006-04:00

Plummers Island, MD, USA, Oct 3, 2009.

The newest published checklist of leeches from Taiwan

2009-10-20T12:38:00.008-04:00

依據2009年發表的paper，台灣現有18種蛭類。

Two new species of Helobdella Blanchard 1896 (Hirudinida: Rhynchobdellida:Glossiphoniidae) from Taiwan, with a checklist of hirudinea fauna of the island

Journal
Zootaxa 2068: 27–46 (2009)

Authors
YI-TE LAI, CHIH-HAN CHANG & JIUN-HONG CHEN

Abstract
In this study, two new species of Helobdella leeches, Helobdella octatestisaca n. sp. and Helobdella melananus n. sp., and a newly recorded species, Helobdella europaea, are reported. Morphological characters and DNA barcode analysis were used to identify the three new species. The differences between these species and other morphologically congeneric species were also compared. In addition, the potential ecological impact of these species was discussed and a checklist of the hirudinea fauna in Taiwan, including eight species in Family Glossiphoniidae, four in Hirudinidae, two in Haemopidae, two in Haemadipsidae, and two in Salifidae, was provided.

The newest published checklist of earthworms and aquatic oligochaetes from Taiwan

2009-10-20T11:24:00.013-04:00

依據這篇2009年的paper，台灣現有98種寡毛類，包含72種蚯蚓。

A checklist of oligochaetes (Annelida) from Taiwan and its adjacent islands

Journal
Zootaxa 2133: 33–48 (2009)

Authors
CHU-FA TSAI, HUEI-PING SHEN, SU-CHEN TSAI, KANG-JIEH LIN, HWEY-LIAN HSIEH &SHAO-PIN YO

Abstract
This checklist lists 98 species and subspecies of oligochaetes (Annelida) that have been reported so far as members of the natural fauna of Taiwan and its adjacent Lanyu (Botel Tobago), Gueishan (Turtle) and Penghu (Pescadore) Islands. Terrestrial oligochaetes have been studied for over a century since 1898, particularly intensively in the past ten years. They are represented by 72 species of which 42 (58.3%) species and subspecies are endemic to Taiwan. They belong to the families Moniligastridae, Glossoscolecidae, Lumbricidae, Octochaetidae, Ocnerodrilidae, and Megascolecidae. Megascolecidae is the most dominant family with 63 species, comprising 87.5% of the terrestrial Taiwanese earthworms reported so far. Aquatic oligochaetes have been investigated for about a decade since 1995, and are represented by 26 species. Of them, 25 are in the family Naididae (including Tubificidae) and Eiseniella tetraedra in the family Lumbricidae.

WORMS love EARTH - earthworms & their soil

Unravelling some Kinki earthworms (Annelida: Oligochaeta: Megadrili: Megascolecidae) - Part II

Impact of above- and below-ground invertebrates on temporal and spatial stability of grassland of different diversity

Earthworm Invasion as the Driving Force Behind Plant Invasion and Community Change in Northeastern North American Forests

Dietary flexibility aids Asian earthworm invasion in North American forests

Re-integrating earthworm juveniles into soil biodiversity studies: species identification throughDNAbarcoding

Cryptic speciation of hormogastrid earthworms revealed by mitochondrial and nuclear data

High mitochondrial DNA sequence diversity in the parthenogenetic earthworm Dendrobaena octaedra

DNA Barcoding Reveals Cryptic Diversity in Lumbricus terrestris L., 1758 (Clitellata): Resurrection of L. herculeus (Savigny, 1826)

Phylogenetic analysis of New Zealand earthworms (Oligochaeta: Megascolecidae) reveals ancient clades and cryptic taxonomic diversity

Prey choice by carabid beetles feeding on an earthworm community analysed using species- and lineage-specific PCR primers

Aporrectodea trapezoides

Aporrectodea rosea

Octolasion lacteum

Octolasion cyaneum

Eisenoides loennbergi

Aporrectodea caliginosa

Allolobophora chlorotica

Lumbricus friendi

Lumbricus rubellus

Lumbricus terrestris

台灣已命名蚯蚓增加到79種

Rob's new paper

A paper in SSB

Two new species and some new records from eastern Taiwan

"Population Genetics" of Hormogaster elisae

Earthworm Fauna of Taiwan 一書重要訊息

phylogeography of a Japanese earthworm

Draft genome of Lumbricus rubellus

Earthworm invasion in the Annual Review of Ecology, Evolution, and Systematics

Earthworms know if the partner is virgin or not!

Earthworms affecting seedlings

What is earthworm grunting? Why does it work?

Use DNA barcodes to identify earthworms: a real case

A mini-review on earthworm molecular ecology. Looking forward for more research papers!!

DNA barcoding of earthworms

Are Aporrectodea caliginosa & A. trepezoides different species?

The first earthworm paper in Molecular Phylogenetics & Evolution

Revisiting Allolobophora chlorotica

Earthworm ID issue and DNA barcoding

The first earthworm paper in MOLECULAR ECOLOGY

Amynthas robustus 壯偉遠環蚓

Amynthas hilgendorfi

Amynthas aspergillum 參狀遠環蚓

Earthworm Fauna of Taiwan

Earthworm Fauna of Taiwan: Metaphire formosae species group

Related links

Amynthas hupeiensis 湖北遠環蚓

The newest published checklist of leeches from Taiwan

The newest published checklist of earthworms and aquatic oligochaetes from Taiwan