earthworm biology and ecology

THE HIGHER TAXA (LOS TAXA SUPERIORES) Phylum ANNELIDA

Class CLITELLATA !

hermaphroditic annelids, without parapodia but with metameric segmentation internally as well as externally,

!

with gonads very few and in definite segmental as well as intrasegmental locations,

!

with special ducts for discharge of genital products,

!

at maturity with a definitely located tumescence of the epidermis (clitellum) to secrete a cocoon in which eggs are deposited to be fertilized and develop therein without a free larval stage,

!

with spermathecae for storage of gametes received during exchange of sperm with a copulatory partner.

Remarks: Departures from the above norm, in the Clitellata, are known. Parthenogenesis in many groups, asexual budding, etc. in the microdriles, marsupial-like brood chambers in certain leeches, lack of spermathecae in some leeches and certain megadriles (Criodrilidae) and total absence of various sexual organs in all known specimens of some oligchaetes are common. A detailed discussion can be found in the supplied reprint [Reynolds, J.W. 1974. Are oligochaetes really hermaphroditic amphimictic organisms? Biologist 56(2): 90-99.] Oligochaetes are more closely related to the leeches than to the polychaetes. This has long been recognized by notable authors (Avel, Gates, Michaelsen, Pickford and Stephenson). Michaelsen (1919, 1928) was the first to recognize this and he combined two orders (Oligochaeta and Hirudinea) in a class that he named Clitellata. Acceptance of that grouping requires the Oligochaeta to be a subclass or an order rather than a class, unless the Clitellata is to become a subphylum of the Annelida.

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J. W. Reynolds

Earthworm Biology and Ecology

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Order OLIGOCHAETA !

annelids with spacious coelomic cavities containing coelomocytes of various sorts,

!

with a closed vascular system comprising at least a dorsal and a ventral trunk,

!

with setae typically in each segment except the peristomium and the periproct;

!

ovaries behind the testis segments

Remarks: Here also there are departures from what now seems to be the norm of oligochaete structure as characterized above. The branchiobdellids not only have lost their setae but have developed a sucker posteriorly as well as certain other leech-like characters. Experts in this group (Holt, 1965) prefer to place this group as an order within the Clitellata. There are competing higher classification systems. One is presented earlier on p. 16 (Reynolds & Cook, 1993). Another version will be published soon (Coates, Gelder, Madill, Reynolds and Wetzel, 1996): Phylum Annelida Superclass Aphanoneura Class Acanthobdellae - bristle leeches Order Acanthobdellida Class Branchiobdellae - crayfish worms Order Branchiobdellida Class Hirudinea - leeches Order Gnathobdellida Order Pharyngobdellida Order Rhyncobdellida Superclass Clitellata Class Oligochaeta Order Enchytraeida - potworms Order Haplotaxida Order Lumbriculida Order Opisthopora - earthworms Order Randiellida Order Tubificida 42

J. W. Reynolds


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Family ACANTHODRILIDAE 1880 1884 1888 ...

Acanthodrilidae (part1) Claus, Grundzüge der Zoologie (ed. 4) 1: 479. Acanthodrilidae (part1) + Pontodrilidae + Plutellidae Vejdovský, System und Morphologie der Oligochaeten (Prag), p. 63. Acanthodrilidae (part1) + + Rosa, Boll. Mus. Zool. Univ. Torino (3), 41: 9. Michaelsen and Pickford reduced to subfamily Acanthodrilinae

1959 1959 1982 ...

Acanthodrilidae Gates, Bull. Mus. Comp. Zool. Harvard 121: 255. Acanthodrilidae (part1, Megascolecidae) Lee, Earthworm Fauna of N.Z., p. 32. Acanthodrilidae Julka, Rec. Zool. Surv. India 80: 130.

In recent years, Fragoso and James have gone to employing Tribes Acanthodrilini and Dichogastrini in the family Megascolecidae. Bouché, Csuzdi, Julka, Reynolds, Righi and Zicsi have not followed this practice. 1

excluding the species with meroic excretory systems which are placed in the Megascolecidae.

Diagnosis: !

Digestive system, with an intestinal origin behind xiii

!

Vascular system, with hearts behind xi

!

Excretory system of holoic nephridia

!

Spermathecae, diverticulate (in front of the testis segments except in 1 species of Plutellus)

!

Clitellum, multilayered, including female pore segment

!

Ovaries fanshaped and with several egg-strings

!

Seminal vesicles, trabeculate

!

Prostates, tubular and of ectodermal origin

!

Males pores, in xvii or xviii

!

Dorsal pores, present or lacking

!

Setae, eight per segment

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J. W. Reynolds

Remarks: The Acanthodrilidae as now recognized is defined morphologically by characters common to all included species and genera.

Species recorded from Argentina: Acanthodrilus doello-juradoi Cordero, 1942 Microscolex collislupi Michaelsen, 1910 Microscolex dubius (Fletcher, 1887) Microscolex michaelseni Beddard, 1895 Microscolex phosphoreus (Dugès, 1837) Notiodrilus bovei (Rosa, 1889) Notiodrilus silvestrii Rosa, 1901


Córdoba

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Family GLOSSOSCOLECIDAE 1921

Glossoscolecidae Michaelsen, Arch. Naturg. 86(A): 141.

Diagnosis: !

Digestive system, with one esophageal gizzard in vi (or its homoeotic equivalent), with paired, extramural calciferous glands in some of segments vii-xiv (or their homoeotic equivalents)

!

Vascular system, with dorsal and ventral trunks, a supra-esophageal, paired extraesophageals median to the hearts, and a subneural adherent to the parietes

!

Excretory system, nephridia, macroic, in intestinal region holoic, vesiculate

!

Spermathecae, adiverticulate (with few exceptions anterior to the testis segments)

!

Clitellum, saddle, multilayered

!

Ovaries, in xiii (= metagynous)

!

Seminal vesicles, frequently lacking

!

Prostates with muscular ducts, none

!

Male pores behind female pores

!

Dorsal pores, none

!

Setae, sigmoid, simply pointed

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J. W. Reynolds

Remarks: Beddard (1895: 623) in his monograph admitted that there was "no one character found in all the Glossoscolecidae which is absolutely characteristic of the family." Nevertheless, there has been no disagreement since 1900 as to the content of the taxon but only as to its hierarchical status, phylogeny, and relations to other taxa (Gates, 1972: 53). The range, restricted to a definite portion of the Americas, rather than the anatomy, presumably provides the explanation for a situation so unusual in the classical system (i.e. Michaelsen-Stephenson system). The only characters without exceptions or qualifications in definitions of the group by Stephenson (1923, 1930) were: (1) a single esophageal gizzard, and (2) calciferous glands present. Both characters, it is noteworthy, are solely somatic. Neither character, nor a combination of the two, is diagnostic, being present alone or in combination in other families. A third character, spermathecae adiverticulate, was included by Beddard (1895) and Michaelsen (1900) in their definitions but is equally applicable to various other families. Although by 1918 Michaelsen had realized that internal structure as well as number and location of calciferous glands was of some importance in defining genera, he continued to ignore vascular and excretory systems.

Species recorded from Argentina: Anteoides rosae Cognetti, 1902 Diaguita michaelseni Cordero, 1942 Enantiodrilus borelli Cognetti, 1902 Glossoscolex bergi (Rosa, 1900) Glossoscolex bonariensis Cordero, 1942 Glossoscolex corrientus Righi, 1984 Glossoscolex forguesi (Perrier, 1881) Glossoscolex uruguayensis uruguayensis Cordero, 1943 Glossoscolex uruguayensis ljungstromi Righi, 1978 Opistodrilus borelli Rosa, 1895 Pontoscolex corethrurus (Müller, 1847) Rhinodrilus parvus (Rosa, 1895)


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Family LUMBRICIDAE 1876

Lumbricidae (part.), Claus, Grundzüge der Zoologie (ed. 3) 1: 416. Lumbricius and Helodrilus)

(included only

Diagnosis: !

Digestive system, with an intramural calciferous gland1, a terminal esophageal valve reaching into xv, an intestine beginning with a crop followed by a gizzard2, a sacculated and unsacculated portion and ending in an intestine with typhlosolate and atyphlosolate regions3, but without intestinal caeca and supra-intestinal glands.

!

Vascular system, with complete dorsal, ventral, subneural (and lateroneural) trunks, the latter adherent to the nerve cord

!

Excretory system, nephridia, holoic, vesiculate, ducts passing into parietes in region of B

!

Spermathecae, adiverticulate, pores at intersegmental levels

!

Clitellum, saddle, multilayered, always behind xvii

!

Ovaries, in xiii, bandlike, each terminating distally in a single eggstring

!

Seminal vesicles, 2-4 pairs in ix-xii

!

Prostates with muscular ducts, none

!

Male pores (in xv) behind female pores (in xiv)

!

Dorsal pores, present

!

Setae, sigmoid and single pointed, eight per segment, in regular longitudinal ranks, in genital tumescences elongated but slender and longitudinally grooved ectally

1

Always in the region of x-xiv? Mainly in xvii of anthropochorous species. In the remainder of the family? 3 Sometimes called a rectum or proctodeum. 2

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J. W. Reynolds

Remarks: The history of the systematics and nomenclatural problems with the Lumbricidae is presented elsewhere in the text (pp. 58-65) from Reynolds, J.W. (1995) [Status of exotic earthworm systematics and biogeography in North America. Pp. 1-27 In Hendrix, P.F. (ed.), Earthworm ecology and biogeography.]

Species recorded from Argentina: Allolobophora georgii Michaelsen, 1890 Aporrectodea rosea (Savigny, 1826) Aporrectodea trapezoides (Dugès, 1828) Aporrectodea turgida (Eisen, 1873) Bimastos beddardi sophiae Mercandal & Barrio, 1988 Bimastos parvus (Eisen, 1874) Dendrodrilus rubidus (Savigny, 1826) Eisenia foetida (Savigny, 1826) Eiseniella tetraedra (Savigny, 1826) Octodrilus complanatum (Dugès, 1828) Octodrilus transpadanus (Rosa, 1884) Octolasion cyaneum (Savigny, 1826) Octolasion tyrtaeum (Savigny, 1826)


Córdoba Córdoba Córdoba Córdoba Córdoba Córdoba Córdoba Córdoba

Córdoba Córdoba

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Family MEGASCOLECIDAE 1891 1895 1900 1921 1930 1959

Megascolecidae (part1), Rosa, Ann. Naturhist. Hofmus. Wien 6: 379. Megascolecidae (part1), Beddard, Monogr. Order Oligochaeta, p. 357. Megascolecinae (Megascolecidae), part1, Michaelsen, Das Tierreich, 10: 161. Megascolecidae (part1), Michaelsen, Arch. Naturgesch. 86(A): 141. Megascolecinae (Megascolecidae), part1, Stephenson, The Oligochaeta, p. 828. Megascolecidae, Gates, Bull. Mus. Comp. Zool. Harvard Coll., 112: 255.

1

excluding all species without truly racemose prostates

Diagnosis: !

Digestive system, with intestinal origin behind ovarian segment

!

Vascular system, with a supraesophageal trunk or trunks, extra-esophageals median to the hearts

!

Excretory system, nephridia, meroic usually avesiculate

!

Spermathecae, in front of gonadal segments, with seminal chambers

!

Clitellum, annular, multilayered, female pore segment always included

!

Ovaries, in xiii, fanshaped, with numerous eggstrings

!


!

Prostates, present

!

Male pores (xviii frequently), behind female pores

!


!

Setae, perichaetin, sigmoid, with simply pointed tip

morphs

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with reference to parthenogenesis, a group of individuals that reproduce parthenogenetically or that have the option to do so, and which share a common anatomy as a result of the degradations, deletions, or other changes from structure of the ancestral amphimictic population as a result of isolation because of the uniparental reproduction.

J. W. Reynolds

H morph

Designates the biparental population (species is defined from H morph rather than from any of its degraded derivatives). Full complement of reproductive organs each of which is capable of functioning in a normal manner. Sperm are mature and, when opportunity permits, exchanged in copulation or at least passed on to partner even if none are received in return.

A morph

Athecal (= without spermathecae).

R morph

Anarsenosomphic (= without male terminalia, i.e. Genital markings and GM glands, lacking; male deferent ducts reduced or lacking)

I morph

First order intermediate morphs. These differ from the A only in having a spermatheca, sometimes normally developed, otherwise more or less rudimentary, at one of the normal H sites. (I1 understood)

I2 morph

Second order intermediate morphs differ from the H in absence of some genital markings and associated glands, on one or both sides of the body, or in absence of male terminalia only on one or both sides of the body, or in various combinations of the two organ deletions. Intermediate between H and R morphs.

I3 morph

Third order intermediate morphs differ from the R morph in absence of one or more of the spermathecae. Intermediate between R and AR morphs.

I4 morph

Fourth order intermediate morph between H and AR morphs.

I5 morph

Fifth order intermediate morph betwenn A and AR.

Ip morph

Pseudo-intermediate morphs are characterized by presence of male terminalia (on other than normal segment), on one or both sides, presence of a spermatheca, presence of a genital marking and its associated gland on one or both sides where the male terminalia should be.

AR morph

This is a fourth order intermediate characterized by being aprostatic (but still with prostatic ducts) and athecal; prostates may be rudimentary and testis sacs may be lacking.

ARZ morph

This is a rare terminal morph known only from two specimens of Metaphire houlleti characterized by being athecal, anarsenosomphic parthenogenetic morph without testes.

Hp morph

This is a hermaphroditic parthenogenetic morph in which the organs such as spermathecae, seminal vesicles, testes, and prostates remain juvenile.

Z morph

This is a parthenogenetic morph without testes.


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Remarks: The morphological and phyletic heterogeneity of the classical Megascolecidae was somewhat simplified in the 1920's by Michaelsen's separating off, as the Acanthodrilidae, all subfamilies characterized time remained morphologically undefinable by any single, significant "character-in-common". The classification by Michaelsen (1900), regarded as a triumph of arrangement which brought order into confusion and that constituted a remarkable advance in our understanding of the group, was retained with but minor modification by Stephenson (1923, 1930) in both his monographs. Stephenson believed that the obvious systematic difficulties were capable of resolution and that they were due to nature itself rather than to a phylogenetic and superficially circumscribed classification. One result of this belief in subsequent years (1930-1960) was the neoclassical modifications, which were not based on new knowledge of somatic anatomy, but simply amounted to reshuffling species and genera into greatly increased phyletic heterogeneity. The racemose and of mesoblastic origin (Gates, 1972: 131-2).

Species recorded from Argentina: Amynthas diffringens (Baird, 1869) Amynthas hawayanus (Rosa, 1891) Amynthas morrisi (Beddard, 1892) Chilota bicinctus (Beddard, 1895) Chilota corralensis var. belgica ernosvitov, 1934 Chilota patagonicus (Kinberg, 1867) Metaphire californica (Kinberg, 1867) Metaphire posthuma (Vaillant, 1868) Yagansia beddardi (Rosa, 1895) Yagansia gracilis (Beddard, 1895) Yagansia longiseta (Beddard, 1895) Yagnasia neuquina Alvardo, 1971 Yagnasia papillosa (Beddard, 1895)

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Córdoba Córdoba Córdoba

Córdoba

nomen nudum

J. W. Reynolds


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Family OCNERODRILIDAE 1966

Ocnerodrilidae Gates, Ann. Mag. Nat. Hist. (13), 9: 53.

Diagnosis: !

Digestive system, with extramural calciferous glands in ix-x

!

Vascular system, with extra-esophageal trunks that pass into the calciferous glands, branch therein, and after uniting posteriorly emerge to join the supra-esophageal trunk

!

Excretory system, nephridia, holoic, avesiculate

!

Spermathecae, present or lacking (A, AR, ARZ morphs)

!

Clitellum, annular, multilayered, range xiii-xx

!

Ovaries, in xiii, fanshaped to almost circular, with numerous eggstrings

!

Seminal vesicles, in ix and xii

!

Prostates with muscular ducts, present

!

Male pores (xvii), behind female pores (xvi)

!

Dorsal pores, lacking

!

Setae, lumbricin, sigmoid, eight per segment

Morphs

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A, AR, ARZ present

J. W. Reynolds

Remarks:

Species recorded from Argentina Belladrilus jimi Righi, 1984 Belladrilus (Santomesia) emiliani Righi, 1984 Eukerria ascuncionis (Rosa, 1895) Eukerria eiseniana (Rosa, 1895) Eukerria garmani argentinae Jamieson, 1970 Eukerria halophila (Beddard, 1892) Eukerria mcdonaldi (Eisen, 1893) Eukerria saltensis (Beddard, 1895) Eukerria santafeina Ljungström, 1971 Eukerria stagnalis (Kinberg, 1867) Eukerria subandina (Rosa, 1895) Eukerria tucumana Cordero, 1942 Eukerria weyenberghi Cordero, 1942 Gatesia unica Jamieson, 1962 Ilyogenia comondui Eisen, 1900 Ilyogenia paraguayensis (Rosa, 1895) Ocnerodrilus occidentalis Eisen, 1878 Quenchua michaelseni Jamieson, 1962


Córdoba Córdoba

Córdoba Córdoba

Córdoba

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Family OCTOCHAETIDAE 1900 1959

Octochaetinae + + (Megascolecidae), Michaelsen, Das Tierreich 10: 319. Octochaetidae, Gates, Bull. Mus. Comp. Zool., Harvard Coll., 121: 254.

Diagnosis: !

Digestive system, with or without gizzards (0, 1, 2)

!

Vascular system, with or without subnural trunk

!

Excretory system, meroic (enteronephric and exonephric)

!

Spermathecae, pregonadal and diverticulate

!

Clitellum, annular, multilayered, xiv-xvi +/-(xiii-xviii)

!

Ovaries, in xiii, fanshaped to almost circular, with several eggstrings

!


!

Prostates tubular and of ectodermal origin

!

Male pores (xvii), behind female pores (xvi)

!


!

Setae, lumbricin and perichaetin

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J. W. Reynolds

Remarks:

Species recorded from Argentina Dichogaster bolaui (Michaelsen, 1891) Dichogaster saliens (Beddard, 1892)


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THE LOWER TAXA (LOS TAXA INFERIORES) An example of the problems encountered in classifying the lower taxa was summarized in a review paper presented in Georgia (USA) during July 1993. The problems encountered with the Lumbricidae are greater than in many of the other families, primarily because more people have become involved over the years. This paper is presented as an example of the types of characters used in earthworm classification and how they have been employed and modified over the years. The Status of Exotic Earthworm Systematics and Biogeography in North America - a case study El status de las lombrices exoticas, sistemática y biogeografia en Norte Ameérica - un estudio detallado [from Reynolds, J.W. 1995. In Earthworm ecology and biogeography in North America, Hendrix, P.F. (ed.) Pp. 1-27. Boca Raton, FL: Lewis Publ., 244 p.]

Abstract This paper traces the taxonomic and nomenclatural history of major contributions to the Lumbricidae from 1758 through 1993. Although this history is primarily based on European reports, significant North American accounts are included. The distribution of exotic Lumbricidae are covered briefly in the text, together with a species list, including general distributional patterns. There is a detailed presentation of regional surveys for Canada and the United States, with supporting references. The concluding portion of the paper deals with current research and suggested priorities for future research activity. I. Introduction Which species are the exotic earthworms of continental North America, north of Mexico? In the most recent checklist on the subject (Reynolds and Wetzel 1994), in twelve families, excluding the Enchytraeidae, there are 147 species, of which 45 or 31 % cent are introduced. These introduced, or exotic, earthworms are found in primarily two families, i.e. 25 species of Lumbricidae and 14 species of pheretimoid Megascolecidae. This is a very small number, when you consider that the global number of species in each of these groups exceeds 400. This paper will deal with the species in the family Lumbricidae, except for the genera Bimastos and Eisenoides, which will be covered in the presentation on endemic species.

Any discussion of the Lumbricidae must consider the European literature, the centre of origin of this family, and the area exhibiting the greatest species diversity. All historical discussion must begin with Linneaus, who in his 10th edition of Systema Naturae (1758, p. 647) described Lumbricus terrestris: LUMBRICUS Corpus teres, annulatum. longitudinaliter exasperatum, poro laterali. . . . Lumbricus humanus Habitat in Humo, adscendit noctu [.] Corpus annulis ciciter centrum conftans, annuloque majore cartilagineo cinctum, trifariam retrorfum aculeatum; quarto latere inermi . . . .

In the 10th and subsequent editions of Systema Naturae, he still placed all species in the genus Lumbricus. There was an emended definition in the 12th edition (1767, p. 1077), which has solidified the opinion that Linnaeus did describe L. terrestris, "exit supra terram tempore nocturno pro copula." It was not until 1826, when Savigny was working in Paris, that a second genus was described for the terrestrial earthworms. The mention of the genus Enterion is primarily of historical interest. Although most of the 20 or so species described by Savigny still are valid and recognizable today, the genus name has long disappeared, and its species are found in nine of the familiar genera in North America: Allolobophora, Aporrectodea, Dendrobaena,

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Dendrodrilus, Eisenia, Eiseniella, Lumbricus, Octolasion and Satchellius. In 1845, the genus Helodrilus was described by Hoffmeister. There has never been a species of this genus recorded from North America. It was not until 1874 that additional new genera were described. Eisen (1874a, b) added Allolobophora, Allurus (= Eiseniella) and Dendrobaena. Many authors over the years have classified the species of Allolobophora and Helodrilus of Hoffmeister interchangeably. Malm described the new genus Eisenia in 1877. In 1885, Örley described the new genera Aporrectodea and Octolasion. In later years, Bouché (1972) and Perel (1976) employed Bouché's synonym Nicodrilus for Aporrectodea. The first North American genus to be described in the Lumbricidae was Bimastos Moore (l893). Some years earlier (1873-4), Eisen did describe lumbricid species from initial collections from North America. Bimastos, in the strict sense, is a North American genus, and over the years many European authors have placed species of Bimastos in Allolobophora and occasionally Helodrilus. In 1893, one of the first thorough revisions of the Lumbricidae was made. Rosa divided his species of Allolobophora into two subgenera (Allolobophora and Notogama), plus the genera Eophila (described in another paper in 1893), Dendrobaena and Octolasion. In the development of taxonomy of earthworms, there have been significant exchanges in attributing the species to one genus or another, and separating and regrouping them into new genera and subgenera. Rosa began a new period, in which the problems of classifying the species into single genera within the Lumbricidae began to be treated from a complex viewpoint. Beddard (1895), in his monograph, defined the Lumbricidae on five somatic characters and four genital organs. Unfortunately, inexactness then, and on many subsequent occasions, reduced the value of some characters, more particularly the conservative and potentially useful somatic characters. Michaelsen's Das Tierreich (1900) was the first major revision of all the Oligochaeta at the time. In it, he renamed Allurus as Eiseniella. Since 1900, Eiseniella and Lumbricus have undergone virtually no changes with respect to the classification of species and subspecies. In Michaelsen's system, he followed that proposed by Rosa in 1893, with only a few minor changes in the status of certain genera and subgenera. Michaelsen classified the genus Eophila (Rosa) as a synonym for the genus and subgenus Helodrilus. He also classified the genera Allolobophora and Dendrobaena of Eisen and Bimastos of Moore as subgenera of Hoffmeister's Helodrilus. Until the revision of Pop (1941), few revisions were made to Michaelsen's classification system. Smith (1917) relied on the reports and collections of Eisen (1874a, b) and the taxonomy of Michaelsen (1900). In his introduction, Smith surveyed the literature and species described from North American specimens and collections. Smith included only three genera in his Lumbricidae: Lumbricus, Octolasion, and Helodrilus, with its subgenera Allolobophora, Bimastos, Dendrobaena, Eisenia and Eiseniella. The spellings, of course, conformed to Michaelsen (1900). It is interesting to note that no new species have been described since Smith's report for Lumbricidae from


59

North America (cf. Species List, infra). In his discussion of Helodrilus (Bimastos), he described the species welchi which was the last lumbricid species to be described from North America. Svetlov (1924) and ernosvitov (1935) continued to apply the same lumbricid taxonomic system as established by Rosa and Michaelsen. The taxonomic systems of Rosa and Michaelsen were imperfect and incomplete, based primarily on the number and arrangement of spermathecae. Until the later work by Gates (1972a, c), all lumbricid taxonomic classification systems relied primarily on the highly variable genital characteristics, vis-à-vis the more conservative somatic anatomy. There were some minor differences between Svetlov and ernosvitov and these were in their placing of species in genera and subgenera. Again, practically no changes were made in the species of Eiseniella and Lumbricus. Stephenson (1930), in his well known book The Oligochaeta, reviewed some of the history of lumbricid taxonomy, but followed the same system as Michaelsen's (1900) Das Tierreich. The first major revision to the classical system of Rosa and Michaelsen came when Pop (1941) pointed out the imperfections of their basic taxonomic characters, i.e. number of seminal vesicles and the position of spermathecae. A summary of Pop's contributions: ! ! ! ! ! !

eliminated the genera Bimastos and Eophila, modified the delimitation of the genera Allolobophora, Eisenia and Dendrobaena, removed from Eisenia species with separate (distinct) setae and placed them into the genus Dendrobaena, and species with widely-paired setae into the genus Allolobophora, removed from Dendrobaena all species with closely-paired setae, considering body colour and setal arrangement, he classified the species of Bimastos and Eophila as Allolobophora, Dendrobaena or Eisenia. Pop was the first to use critically the somatic characters of body pigmentation and setal arrangement in an earthworm classification scheme.

Pop was the first to consider the structure of muscle fibres in the longitudinal musculature, but he did not employ this character consistently throughout his taxonomic system. He did establish a new basic taxonomic system for lumbricid earthworms which correctly classified a large number of species. Notwithstanding this valuable contribution, Pop did place into the genus Allolobophora species that belong to different genera and subgenera. The important result of Pop's system was that it introduced a new approach to solving taxonomic problems regarding earthworms, by forcing authors to re-evaluate old taxonomic characters and consider new ones. It was almost 15 years before any major new effort was made to revise Pop's system. Omodeo (1956) made an attempt at a general solution to the taxonomic classification of earthworms, by employing new characters in the diagnosis of genera. He took an interesting approach to the study of generic biogeography, by comparing present distribution with paleogeographical maps. On the basis of these comparisons, he determined the distribution types of a single species. Omodeo's solution to the taxonomic classification of species was based on the available data at the time. Currently, we attribute species he classified as Eophila in different genera. Omodeo introduced into earthworm taxonomy several new characters, e.g. shape of calciferous glands, chromosome number and structure of longitudinal muscle fibres, etc. Omodeo described a larger number of new genera and subgenera: ! ! !

60

he divided Dendrobaena into subgenera Dendrobaena and Dendrodrilus, he divided Octolasion into subgenera Octolasion and Octodrilus, he divided Allolobophora into subgenera Allolobophora (3 groups) and Cernosvitovia, and genera Bimastos,

J. W. Reynolds

! !

Eiseniona, Eophila (3 groups), Helodrilus and Microeophila, expanded the diagnosis of Eisenia, Eiseniella and Lumbricus, Eiseniona erected as a new genus, many of its species are now in Aporrectodea.

Gates (1957) produced the first of what would become by 1980 a 26 part series of Contribution to a revision of the earthworm family Lumbricidae. This was followed in 1958 by part two. These 26 papers by Gates are the only serious attempt, to date, by a North American to revise the Lumbricidae. In 1968, Gates produced three more papers in his lumbricid revision series (Gates 1968a-c). Gates( 1969a) redefined two lumbricid earthworm genera, Bimastos and Eisenoides, the latter erected for two North American species which had for years been attributed to Allolobophora, Eisenia and Helodrilus. This was his sixth paper in his lumbricid series, which was followed a few months later by his seventh (Gates 1969b). The next major revision to lumbricid taxonomy was that of Bouché (1972). He retained a part of Omodeo's taxonomic system (genera Dendrobaena, Octolasion, Octodrilus, Eiseniella, Lumbricus) and retained the subgenera Dendrobaena and Dendrodrilus. In addition, he described the genus Kritodrilus. He included in his Eisenia (sensu Pop), two species which have subsequently been placed in different genera (Allolobophoridella eiseni and Bimastos parvus). Bouché made dramatic changes when it came to Allolobophora and Eophila. Based on species found in France, he classified them as Allolobophora, Helodrilus and the new genus Nicodrilus (now junior synonym of Aporrectodea) with subgenera Nicodrilus and Rhodonicus. The species of Allolobophora were classified as Allolobophora sensu stricto and sensu lato, which would indicate that he did not find a satisfactory solution to the classification of the species in this genus. The genus Eophila was replaced by his three new genera: Orodrilus, Prosellodrilus and Scherotheca. !

!

! ! !

The following is a summary of Bouché's (1972) contribution to the revision of the Lumbricidae: descriptions of 24 new species, 28 subspecies and 18 varieties, notwithstanding that the International Code of Zoological Nomenclature [Art. 45(a)] states that the species-group includes only the specific and subspecific categories, six new genera erected: Orodrilus, Prosellodrilus and Scherotheca (to deal with problems in Dendrobaena and Allolobophora, plus the elimination of Eophila), Ethnodrilus (primarily for ecological reasons), Kritodrilus (to classify Tétry's species calarensis) and Nicodrilus (now = Aporrectodea, to help with the Allolobophora problems). four new subgenera erected, Dendrodrilus was retained as a subgenus of Dendrobaena; species of Bimastos, Eophila and Helodrilus of others placed in various genera, and large quantities of ecological data attributable to species; extensive sampling employed, plus access to numerous collections of colleagues.

In 1972, Gates continued with two more papers (8th and 9th) in his contribution series on lumbricid revision. The first was a large and significant contribution (Gates 1972a), wherein he demonstrated, using morphological somatic characters, that the trapezoides complex, or what was frequently a collection of up to seven species, with three to four frequently included in what is termed Allolobophora caliginosa, were in fact distinct species, i.e. trapezoides (Dugès 1828), tuberculata (Eisen 1874), turgida (Eisen 1873), longa (Üde 1885), limicola (Michaelsen 1890), nocturna (Evans 1946) and icterica (Savigny 1826). Gates stated early in his introduction what was his own "esotery", which was formulated initially during his early years of working with the classical earthworm taxonomic system. After working with earthworm systematics for 45 years, he summed it up as follows: !

All individuals that are closely related enough to belong to one species should demonstrate that relationship by a considerable common anatomy not subject to individual variation (disregarding abnormality and monstrosity).


61

! !

All species so closely related as to belong to a single genus and presumably with a common ancestry, should demonstrate that relationship by a lesser amount of invariable common anatomy. All genera belonging to a single family also should prove that relationship by a still smaller amount of commonly shared, invariant anatomy. (emphasis added, Gates 1972a: 1-2)

Gates' observation was that if one carefully examined classical and neoclassical definitions of genera and families, they would reveal a lack of this invariation. Two of the characters upon which Gates placed considerable emphasis in the separation of these species of the trapezoides complex were: location of genital tumescences, typhlosole termination and segment number. Many European authors prefer to use caliginosa for turgida and several of the other species as forms or varieties of caliginosa, e.g. trapezoides and tuberculata , Perel, Sims and Zicsi). Bouché (1972) used different form names, which were soon put into synonomy, but were continued by Easton (1983) in his list. For a period of time, there was also confusion concerning Lumbricus terrestris and Allolobophora terrestris (Savigny) and with Aporrectodea longa, particularly in England. The taxonomic and nomenclatural problems which occurred in Europe, in this case, have not been encountered in North America. Gates placed these species in the genus Aporrectodea, within two years after this paper (1972a) appeared. In subsequent years, Reynolds (1975a, b, 1976a-d, 1994b, and others) using these criteria, was able to show in biogeographical surveys throughout North America, that not only were these separate species, but that they had distinct distributional patterns. In a very recent study, Bøgh (1993), using electrophoretic techniques, upheld the existence of these species (in part, for those which were included in his study). In his second paper of the year (1972b), Gates dealt with his own evaluation of Eisenoides, which was made possible by many wide ranging collections of specimens by the Tall Timbers staff. Perel (1973) compared the nephridial bladders from Bouché's genera with those of the Balkans in Omodeo's Eophila, and determined that the different (opposite) orientation indicated that the archaic species of France and those of the Balkans have resulted from independent development. In this review, Perel stated that Bouché's taxonomic system possessed no criteria allowing one to state the generic affinities of species. Although the greatest problem with Bouché's system was his overwhelming reliance on intuition, he was the first to examine intergenerically the solutions to the problems of phylogeny. He introduced "series of species", by means of which the relationships of the species from the same genus could be shown in an illustrative manner. In 1973, Gates continued with his 10th paper, in which he examined Octolasion. This paper was strongly critical of neoclassical revisionists, particularly Omodeo (1956), for failing to be precise with invariant characters, especially those of the more conservative somatic systems. He presented new information on the vascular and excretory systems and down-played the usefulness of classifying the calciferous sacs in this genus. In the following year, Gates (1974a, b) continued his series with contributions on Dendrobaena octaedra and Eisenia rosea. The first paper is significant, because of the discussion of the importance of polymorphism and parthenogenetic populations for the classification of earthworms. Gates drew on many years of experience in studying the pheretimoid earthworms (Megascolecidae), where parthenogenetic morphs in many of the hundreds of species had created a nightmare for oligochaetologists over the decades. It was a busy year for Gates when, during 1975, he published six new papers in his contribution series on the revision of the Lumbricidae. In his first paper of the year (Gates 1975a), he described the genus Satchellius with Enterion mammale Savigny, 1826 as the type. In this paper, Gates also included new somatic information to be included in new diagnoses for 13 genera of the Lumbricidae. The characters he included were: calciferous sacs, position of calciferous gland opening into esophagus, calciferous lamellae nephridial vesicles, nephropores and, in some cases, pigmentation and prostomium. The other five papers (Gates 1975b-f) had no direct bearing on North American collections or distributions, except for a couple of references to U.S. Quarantine Bureau interceptions.

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Bouché (1975) described a new genus, Spermophorodrilus, based on a collection from Albania. Traditionally, species belonging in this genus had been attributed to Eophila and Bimastos. (1991) erected a new type and placed the original type designated by Bouché as a synonym for another Balkan subspecies, which may have been published in a journal unavailable to Bouché. It was early 1976 when Gates officially published his transfer of Eisenia rosea to the genus Aporrectodea, giving his reasons after tracing the species path through various genera from 1826 to 1976. This was his 19 th paper in the contribution to a revision of the family Lumbricidae (Gates 1976). In 1976, Reynolds and Cook produced the original volume of Nomenclatura Oligochaetologica - a catalogue of names, descriptions and type specimens of the Oligochaeta. This was the first time that the names of all oligochaetes, along with the citation of their descriptions and the location(s) of their type specimens, were assembled in one place. Three additional supplements have been produced, which up-date the original book (Reynolds and Cook 1981, 1989 and 1993). Perel (1976, 1979) evaluated numerous taxonomic characters in her generic revision, but restricted herself to the species found in the Soviet Union. One of the most important aspects of her study was the significance of the structure and shape of the nephridial bladders, and the type of muscle fibre arrangement in longitudinal muscles. She also considered her studies on ecotypes in the revision. Perel amassed considerable data from many sources in her attempt to revise Allolobophora; unfortunately, she restricted herself to Soviet species, although she had sufficient material for a complete revision. She retained Dendrobaena, Dendrodrilus, Eisenia, Eiseniella, Octolasion and Lumbricus. She completed the diagnosis of the genus Kritodrilus. She divided Allolobophora (sensu Pop) into Nicodrilus (= Aporrectodea), Allolobophora (subgenera Allolobophora and Svetlovia [= Perelia]) and Helodrilus. Perelia was erected by Easton (1983) as a new name for Svetlovia which had been used by Olga ekanovskaya in 1975 as a genus of Tubificidae. Gates' 20th paper in his continual revision of the Lumbricidae was based on the genus Eiseniella in North America (Gates 1977). In the subsequent year (Gates 1978a, b), he followed with revisions of the genera Lumbricus and Eisenia, respectively. In his paper on the genus Lumbricus, he included a key to 14 genera of the Lumbricidae, based solely on somatic characters. It was during this discussion that he erected the new genus Murchieonia, with Allolobophora minima Muldal, 1952 (= Bimastos muldali Omodeo 1956) as type species. This genus was erected on the combination of several unique gut characters. In 1978, Zicsi described the genus Fitzingeria, including species which were previously placed by other lumbricologists in Lumbricus (Örley) and Dendrobaena (Rosa, Michaelsen and Perel). It was 1979, when Gates (1979a) presented his revision of the genus Dendrodrilus (sensu Omodeo), including a discussion of the subrubicunda and tenuis morphs. Gates considered Dendrodrilus to be monospecific, although a recent study would indicate that the subrubicunda morph may be a true species (Bøgh 1993). In late 1980, Gates produced the last two papers in his revision of the Lumbricidae (Gates 1980a, b). The first was his revision of the genus Allolobophora, as he searched for the invariant somatic characters which he felt would "enable a solution of the 'systematic chaos' in the European portion of the Lumbricidae." Based on digestive and excretory invariant somatic characters (nephridial vesicles, nephropores and calciferous glands), he supported Omodeo (1956) and subsequent authors, who accepted Al. chlorotica as type species which had been left undesignated in the neoclassical revisionists from Beddard through Michaelsen and Stephenson. His last paper, Gates (1980b) dealt with two octolasian species which have not been recorded in North America. Zicsi (1981) elevated the subgenus Cernosvitovia Omodeo, 1956 of Allolobophora to full generic rank. This has been supported by eastern European lumbricologists ever since. None of the species are found outside eastern Europe.


63

The centenary (1982) of the publication of Darwin's book The Formation of Vegetable Mould through the Action of Worms was the occasion for a symposium which attracted some 150 participants, to discuss various aspects of earthworm ecology. Two papers are of particular interest to this discussion, Sims (1983) and Easton (1983). Chapter 40 was Sims' paper, entitled The scientific names for earthworms. In his seven page paper, he devoted a page to the Allolobophora problem mentioned earlier and the caliginosa confusion. Sims also devoted considerable space to the Octolasion problems (orthography, nomenclature and taxonomy). In an attempt to obtain universality and remove confusion in earthworm taxonomy, Easton (1983) produced a checklist of valid lumbricid species through December 1981, based heavily on Bouché (1972), Perel (1979) and Zicsi (1982). Fender (1985) followed up his earlier paper (Fender, 1982) with a systematic discussion of all eleven lumbricid genera found in the eleven state area of the western United States. In this survey, Fender included considerable amounts of new material collected over the years by McKey-Fender and himself, along with a review of the literature available at the time. He devoted considerable space to evaluating the European and North American positions on Allolobophora and Aporrectodea. He recognized that Levinsen's eiseni did not belong in any as yet described genus, and did not agree with Bouché's (1972) or Gates' (1978a) positions. In his discussion of Dendrodaena, he had the opportunity to include two species not previously recorded in the West or widely in North America. Based on somatic characters (e.g. nephropores), Fender believed that neither Dendrobaena nor any other currently described genus is the proper place for this species. Fender was responsible for the major collections of species of Octolasion in the West, and these observations are included in his discussion. Zicsi (1985) continued his revision of Allolobophora with the description of the new genus Proctodrilus, containing species primarily from eastern Europe, but one or two species extending as far west as France. (1987a, b) described three new genera, viz. Creinella, Meroandriella and Alpodinaridella with two subgenera, Dinaridella and Alpodinaridella. Most of these new taxa were based on specimens from areas in and adjacent had reduced Creinella and Meroandriella to subgenera of Aporrectodea. Together revised the genus Allolobophora sensu Pop, based on 47 species found in the Balkans. They divided the species into 10 genera and 9 subgenera, including two new genera, Italobalkaniona and Karpatodinariona, with its two new subgenera, Panoniona and Serbiona. Again, these are primarily Balkan had elevated Panoniona and Serbiona to full generic status. Omodeo (1988) revised his diagnosis of Eophila, placing some of the species in Bouché's Scherotheca, on the basis of only one morphological character (number of spermathecae), and by area maps in which these genera ranged from Iberia to central Asia. Omodeo apparently did not consider newer taxonomic characters proposed by various authors in more recent times. In the following year, Omodeo (1989) described the genus Healyella, and created two new subfamilies (Lumbricinae and Spermophorodrillinae). The latter was based on its lack of tubercula pubertatis and spermathecae, but the presence of a ridged 15th segment with a male aperture, similar to the tubercula pubertastis. These are new characteristics of earthworm structure and are related to the shifting of the male aperture posterior to the 15th segment. In redefined his earlier revision of Allolobophora and included a new subgenus Allolobophoridella, with Lumbricus eiseni Levinsen, 1884 as the type. In 1991, he also included Allolobophora parva Eisen, 1874 and elevated this subgenus to full generic rank. However, he did have some reservations about whether some of the other species of Bimastos in North America should remain in Bimastos or be transferred into his new genus. His concerns centre on the presence of tubercula pubertata on some European species i.e. B. beddardi, vis-à-vis those of North America where these structures are absent, together with the shape of the nephridial bladder and its orientation (anteriorly vs. posteriorly).

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's major monograph (1991) was delayed, due to political unrest in the former Yugoslavia. In this monograph, he recognized 25 genera and 231 species. A summary of his monograph is as follows: ! ! ! ! ! ! ! !

described 21 new species, elevated subgenera Allolobophoriella, Panoniona and Serbiona to generic rank, retained Microeophila and Spermophorodrilus as genera, ignored Eophila and transferred species to various genera derived from Allolobophora and did not know what to do with some and classified them as species incertae sedis (species of uncertain taxonomic position), ignored Eiseniona Omodeo, 1956 and transferred the species to Aporrectodea and Eisenia, reduced Creinella to subgeneric rank, made a detailed analsyis of some important taxonomic characters and recognized their importance in helping to deal with phylogenetic and taxonomic problems, transferred Bimastos parvus and Lumbricus eiseni to Allolobophoridella, but admitted that new evidence based on orientation of nephridial bladders may result in this new genus being a synonym of Bimastos.

Based on the concept of and in the spirit of Easton (1983), I present (in part) the checklist of Reynolds and Wetzel (Coates et al. 1996) for the current species of Lumbricidae found in continental North America, using the hierarchical system of Reynolds and Cook (1993): ORDER HAPLOTAXIDA SUBORDER LUMBRICINA Superfamily Lumbricoidea Family Lumbricidae Subfamily Lumbricinae Allolobophora Eisen, 1874 1. Type species: Enterion chloroticum Savigny, 1826 (now = Allolobophora chlorotica) Other species: in North America, none. Allolobophoridella , 1990 2. Type species: Lumbricus eiseni Levinsen, 1884 Other species: in North America, none. Aporrectodea Örley, 1885 3. Type species: Lumbricus trapezoides Dugès, 1828 Other species: 3a. Ap. bowcrowensis Reynolds et Clapperton, 1996 4. Ap. icterica (Savigny, 1826) 5. Ap. limicola (Michaelsen, 1890) 6. Ap. longa (Üde, 1885) 7. Ap. rosea (Savigny, 1826) 8. Ap. tuberculata (Eisen, 1874) 9. Ap. turgida (Eisen, 1873) Bimastos Moore, 1893 10. Type species: Bimastos palustris Moore, 1895 Other species: 11. B. beddardi (Michaelsen, 1891) 12. B. gieseleri (Üde, 1895) 13. B. heimburgeri (Smith, 1928) 13a. B. lawrenceae Fender, 1994


nUSA; CAN

AK,OR,TN,WA

widespread AB NY,ON GA,MA,NJ,OR,PA,WA AL,CA,nUSA;eCan,BC widespread widespread widespread

DE,NJ,NC,PA,TN 11 states 9 states (s) 14 states BC

65

14. 15. 16. 17. 18.

B. longicinctus (Smith et Gittins, 1915) B. parvus (Eisen, 1874) B. tumidus (Eisen, 1874) B. welchi Smith, 1917 B. zeteki (Smith et Gittins, 1915)

Dendrobaena Eisen, 1874 19. Type species: Dendrobaena boeckii Eisen, 1874 (= Enterion octaedra Savigny, 1826) (now = Dendrobaena octaedra) Other species: 20. Db. attemsi Michaelsen, 1902 21. Db. pygmaea (Savigny, 1826) Dendrodrilus Omodeo, 1956 22. Type species: Enterion rubidum Savigny, 1826 (now = Dendrodrilus rubidus) Other species: in North America, none. Eisenia Malm, 1877 23. Type species: Enterion fetidum Savigny, 1826 (now = Eisenia foetida) Other species: 24. E. hortensis (Michaelsen, 1890) 25. E. zebra (Michaelsen, 1902) Eiseniella Michaelsen, 1900 26. Type species: Enterion tetraedrum Savigny, 1826 (now = Eiseniella tetraedra) Other species: in North America, none. Eisenoides Gates, 1969 27. Type species: Allolobophora lönnbergi Michaelsen, 1894 (now = Eisenoides lönnbergi) Other species: 28. Es. carolinensis (Michaelsen, 1910) Lumbricus L., 1758 29. Type species: Lumbricus terrestris L., 1758 Other species: 30. L. castaneus (Savigny, 1826) 31. L. festivus (Savigny, 1826) 32. L. rubellus Hoffmeister, 1843 Murchieona Gates, 1978 33. Type species: Bimastos muldali Omodeo, 1956 (= Allolobophora minima Muldal, 1952 preoccupied) Other species: in North America, none.

66

9 states (s) 20 states (s) 19 states (s) IN,KS,MO 15 states

nUSA; CAN

OR,WA CA

nUSA; CAN

widespread

8 states CA

widespread

14 states (e)

13 states (se)

widespread 14 states (n); BC; eCAN VT;BC,NB,ON,PQ widespread

IN,MI,TN

J. W. Reynolds

Octolasion Örley, 1885 34. Type species: Lumbricus terrestris lacteus Örley, 1881 (= Enterion tyrtaeum Savigny, 1826; O. gracile Örley, 1885) (now = Octolasion tyrtaeum) Other species: 35. O. cyaneum (Savigny, 1826)

widespread USA; eCAN

13 states; 4 prov.

Satchellius Gates, 1975 36. Type species: Enterion mammale Savigny, 1826 (now = Satchellius mammalis) Other species: in North America, none.

NJ

III. Survey of North American Earthworm Biogeography The first attempt to present a regional view of the earthworms present in continental North America was by Reynolds (1975a, 1976a). During the past 20 years, there have been many advances and additional available data for many regions (states and provinces) in North America. The results of this additional data are presented in a recent paper (Reynolds 1994b), which includes the latest earthworm taxonomic and distribution maps, together with a discussion of the general concepts of biogeography. A table of regional earthworm surveys was presented by Reynolds et al. (1974). Since that time, this author and others have made wide-ranging collections in North America, resulting in numerous publications of the distribution of earthworms in North America. The following table (Table 1) and the list of species (supra) should give the reader a good overall impression of the biogeography of North American lumbricid earthworms. Figures 1 to 6 at the end of this chapter present distribution maps for many of the lumbricid species found in Alaska, Canada, and the continental United States. Table 1. Regional earthworm surveys in North America. Region

Alabama Alberta Arkansas Connecticut Delaware Florida Georgia Illinois Indiana Louisiana Maryland Massachusetts Michigan Mississippi Missouri Montana New Brunswick New York

Number of lumbricid species 10 67 18 15 13 132 193 19 23 84 20 18 20 10 14 8 135 20


Number of units (%) surveyed1 75 30 29 100 100 85 40 45 100 100 100 100 64 77 27 14 100 47

Reference(s)

Reynolds (1994g) Scheu & McLean (1993) Causey (1952, 1953) Reynolds (1973c) Reynolds (1973a) Reynolds (1994e) Reynolds (prep) Harman (1960) Reynolds (1994a) Harman (1952), Gates (1965, 1967) Reynolds (1974) Reynolds (1977a) Snider (1991) Reynolds (1994f) Olson (1936) Reynolds (1972a) Reynolds (1976c) Oslon (1940), Eaton (1942)

67

North Carolina Nova Scotia Ohio Ontario Oregon

21 15 22 19 18

80 100 63 986 70

Prince Edward Island Quebec (south shore) (north shore) Rhode Island Tennessee Virginia Washington

12 15 18 11 26 23 15

100 100 86 100 100 77 40

Reynolds (1994c) Reynolds (1976b) Olson (1928, 1932) Reynolds (1977b) MacNab & McKey-Fender (1947) Fender (1985) Reynolds (1975b) Reynolds (1976d), Reynolds & Reynolds (1992) Reynolds (1973b) Reynolds et al. (1974) Reynolds (1994d) Altman (1936), Fender (1985) MacNab & McKey-Fender (1947)

1

units are counties, districts, parishes, etc. 10 and 11 species of exotic pheretimoid species not included for Florida and Georgia, respectively. 4 Tandy (1969) studied only the genus Pheretima (8 species recorded for Louisiana). 5 now 14 (Reynolds and Savard 1996). 6 now 98% (Reynolds and Mayville 1994) 7 now 12 and 33% (Reynolds and Clapperton, 1996) 2,3

IV. Current Research and Priorities for Future Research One of the major problems for earthworm taxonomy in North America has been the paucity of scientists, all living in isolation. Many individuals have dabbled in earthworm research, but their contributions can frequently be counted on one hand. For two promising scientists, tragic death came early in life, before their full impact could be realized (William Murchie and Richard Tandy). Gordon Gates, the Dean of Earthworm Taxonomy and Systematics, died in 1987. Although he began publishing in 1926, it was not until the latter part of his career that he devoted much time to the earthworms of North America and the Lumbricidae. Prior to 1950, there were Frank Smith (1985-1937) and Henry Olson (1928-40), who made contributions to taxonomy and distribution, respectively. In the last few decades, Dorothy McKey-Fender and William Fender have been concentrating on the taxonomy and distribution of the native and exotic earthworm fauna of the west coast of North America, and Sam James on the endemic species of the southeastern and plains areas. Since 1972, I have collected widely throughout North America, alone and with the aid of my colleagues at Tall Timbers. The results of these collections have been published as distributional data primarily, with minimal contribution to systematics sensu stricto. Reynolds and Cook (1977, 1981, 1989 and 1993) in the Nomenclatura Oligochaetologica series have brought together in one source the necessary reference data for anyone involved in the taxonomy and nomenclature of earthworms.

V. Research Imperatives With this limited background, I suggest the following priority for research and funding efforts: A. Training of taxonomists For over two decades (Reynolds 1972b; Reynolds et al. 1974), I have stated that the scarcity of competent earthworm systematists/taxonomists was detrimental to ecologists and others. The normal institutions which employ

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J. W. Reynolds

these types of specialists and encourage their development, e.g. museums and Departments of Agriculture, have not done so in North America since Smith worked for the Illinois Natural History Survey in the early 1900's. The difficulty encountered by the Organizing Committee of this conference in obtaining sufficient people to cover all the systematic topics attests to the fact that this problem still exists. There must be a concerted effort to support this type of research, before we are left without competent specialists in our field. B. Parthenogenesis in taxonomy The other major exotic group of earthworms in North America (Megascolecidae, pheretimoid groups) has long been plagued with taxonomic problems, which resulted from widespread parthenogenesis in its species (Gates 1972c). There is parthenogenesis within the Lumbricidae. One study recently has shown that localized populations of Octolasion tyrtaeum (Jaenike et al. 1980, 1982; Jaenike and Selander 1985) exhibited parthenogenesis. Previously, taxonomic problems with some morphs of what is now Dendrodrilus rubidus may be attributable to parthenogenesis. This area of research needs more attention. C. Earthworm surveys From Table 1 in this text it is obvious, in spite of what has already been accomplished, that there are major areas of North America in which earthworm surveys are lacking. In certain areas where native species exist, there is the potential for discovery of new species. Any new species in the Lumbricidae will probably come from the native genera Bimastos and Eisenoides, or a new genus. D. Life histories It is amazing that for the nearly 8000 oligochaetes (Reynolds and Cook 1993), modern, up-dated, life history studies have been done on only a few species, i.e. less than twenty. Some of the information gathered on common Lumbricidae was done at a time when species lumping occurred, i.e. factors attributed to caliginosa which included several species and thus accounted for the range of data, vis-à-vis specific values for other species. E. Modern techniques One area which was considered for years, but only recently had any evidence to support its potential, is electrophoresis. In Bøgh (1993), he illustrated that certain species were different, i.e. Aporrectodea tuberculata is a distinct species, as well as demonstrating how to identify species from fragments. This area of research should be followed, and will probably help with areas of confusion. Acknowledgements I would like to thank Wilma M. Reynolds of the Oligochaetology Laboratory, Lindsay, Ontario for reviewing numerous early drafts of this manuscript and her helpful comments and suggestions. Also, during the Workshop, many colleagues offered useful suggestions to help improve the manuscript. Again, I am indebted to Susan Mantle of the CARMA Centre at Sir Sandford Fleming College, Lindsay, Ontario for producing the maps which appear in this text. Two colleagues in particular, William M. Fender and Samuel W. James, generously shared some of their unpublished distribution records so that these maps might reflect the most up-to-date picture of North American lumbricid distribution.


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Literature Cited Altman, L.C. 1936. Oligochaeta of Washington. Univ. Wash. Publ. Biol. 4(1): 1-137. Beddard, F.E. 1895. A monograph of the order of Oligochaeta. Oxford, 769 p. Bøgh, P.S. 1993. Identification of earthworms (Lumbricidae): choice of method and distinction criteria. Megadrilogica 4(10): 163-174. Bouché, M.B. 1972. Lombriciens de France écologie et systématics. Inst. Nat. Rech. Agron., Paris, 671 p. Bouché, M.B. 1975. La reproduction de Spermophorodrilus albanianus nov. gen., nov. sp. (Lumbricidae). Zool. Jb. Syst. 102: 1-11. Causey, D. 1952. The earthworms of Arkansas. Proc. Ark. Acad. Sci. 5: 31-42. Causey, D. 1953. Additional records of Arkansas earthworms. Proc. Ark. Acad. Sci. 6: 47-48. ernosvitov, L. 1935. Monographie der Tschechoslovakischen Lumbriciden. Arch. P ír. Vyzk. ech. 19: 1-86. Easton, E.G. 1983. A guide to the valid names of Lumbricidae (Oligochaeta). pp. 475-485. In: Earthworm ecology from Darwin to vermiculture (Ed. J.E. Satchell). London: Chapman and Hall, 495 p. Eaton, T.H. 1942. Earthworms of the northeastern United States: a key, with distribution records. J. Wash. Acad. Sci. 32(8): 242-249. Eisen, G. 1874a. Om Skandinaviens Lumbricider. Öfv. K. Vet. Akad. Förh. 30(8): 43-56. Eisen, G. 1874b. New Englands och Canadas Lumbricider. Öfv. Vet. Akad. Förh. 31(2): 41-49. Fender, W.M 1982. Dendrobaena attemsi in an American greenhouse, with notes on its morphology and systematic position. Megadrilogica 4(1-2): 8-11. Fender, W.M. 1985. Earthworms of the western United States. Part I. Lumbricidae. Megadrilogica 4(5): 93129. Gates, G.E. 1957. Contribution to a revision of the earthworm family Lumbricidae. I. Allolobophora limicola. Breviora, Mus. Comp. Zool. Harvard, No. 81: 1-14. Gates, G.E. 1958. Contribution to a revision of the earthworm family Lumbricidae. II. Indian species. Breviora, Mus. Comp. Zool. Harvard, No. 91: 1-16. Gates, G.E. 1965. Louisiana earthworms. I. A preliminary survey. Proc. La. Acad. Sci. 28: 12-20. Gates, G.E. 1967. On the earthworm fauna of the Great American desert and adjacent areas. Gt. Basin Nat. 27(3): 142-176. Gates, G.E. 1968a. What is Lumbricus eiseni Levinsen, 1884 (Lumbricidae, Oligochaeta)? Breviora, Mus. Comp. Zool. Harvard, No. 299: 1-9. Gates, G.E. 1968b. Contributions to a revision of the Lumbricidae. III. Eisenia hortensis (Michaelsen, 1890). Breviora, Mus. Comp. Zool. Harvard, No. 300: 1-12. Gates, G.E. 1968c. What is Enterion ictericum Savigny, 1826 (Lumbricidae, Oligochaeta)? Bull. Soc. Linn. Normandie 10th ser., 9: 199-208. Gates, G.E. 1969a. On two American genera of the family Lumbricidae. J. Nat. Hist. 9: 305-307. Gates, G.E. 1969b. Contributions to a revision of the earthworm family Lumbricidae. V. Eisenia zebra Michaelsen, 1902. Proc. Biol. Soc. Wash. 82: 453-460. Gates, G.E. 1972a. Toward a revision of the earthworm family Lumbricidae. IV. The trapezoides species group. Bull. Tall Timbers Res. Stn., No. 12: 1-146. Gates, G.E. 1972b. On American earthworm genera. I. Eiseniodes (Lumbricidae). Bull. Tall Timbers Res. Stn., No. 13: 1-17. Gates, G.E. 1972c. Burmese earthworms. An introduction to the systematics and biology of Megadrile oligochaetes with specieal reference to southeast Asis. Trans. Amer. Philos. Soc. 62(7): 1-326. Gates, G.E. 1973. The earthworm genus Octolasion in America. Bull. Tall Timbers Res. Stn., No. 14: 29-50. Gates, G.E. 1974a. Contributions to a revision of the Lumbricidae. X. Dendrobaena octaedra (Savigny, 1826) with special reference to the importance of its parthenogenetic polymorphism for the classification of earthworms. Bull. Tall Timbers Res. Stn., No. 15: 15-57. Gates, G.E. 1974b. Contributions to a revision of the family Lumbricidae. XI. Eisenia rosea (Savigny, 1826). Bull. Tall Timbers Res. Stn., No. 16: 9-30. Gates, G.E. 1975a. Contributions to a revision of the earthworm family Lumbricidae. XII. Enterion mammale

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Savigny, 1826 and its position in the family. Megadrilogica 2(1): 1-5. Gates, G.E. 1975b. Contributions to a revision of the earthworm family Lumbricidae. XIII. Eisenia japonica (Michaelsen, 1891). Megadrilogica 2(4): 1-3. Gates, G.E. 1975c. Contributions to a revision of the earthworm family Lumbricidae. XIV. What is Enterion terrestris Savigny, 1826 and what are its relationships? Megadrilogica 2(4): 10-12. Gates, G.E. 1975d. Contributions to a revision of the earthworm family Lumbricidae. XV. On some other species of Eisenia. Megadrilogica 2(5): 1-7. Gates, G.E. 1975e. Contributions to a revision of the earthworm family Lumbricidae. XVII. Allolobophora minuscula Rosa, 1906 and Enterion pygmaeum Savigny, 1826. Megadrilogica 2(6): 7-8. Gates, G.E. 1975f. Contributions to a revision of the earthworm family Lumbricidae. XVIII. Octolasion calarense Tétry, 1944. Megadrilogica 2(7): 1-4. Gates, G.E. 1976. Contributions to a revision of the earthworm family Lumbricidae. XIX. On the genus of the earthworm Enterion roseum Savigny, 1826. Megadrilogica 2(12): 4. Gates, G.E. 1977. Contribution to a revision of the earthworm family Lumbricidae. XX. The genus Eiseniella in North America. Megadrilogica 3(5): 71-79. Gates, G.E. 1978a. The earthworm genus Lumbricus in North America. Megadrilogica 3(6): 81-116. Gates, G.E. 1978b. Contributions to a revision of the earthworm family Lumbricidae. XXII. The genus Eisenia in North America. Megadrilogica 3(8): 131-147. Gates, G.E. 1979a. Contributions to a revision of the earthworm family Lumbricidae. XXIII. The genus Dendrodrilus Omodeo, 1956 in North America. Megadrilogica 3(9): 151-162. Gates, G.E. 1979b. Contributions to a revision of the earthworm family Lumbricidae. XXIV. What is Dendrobaena byblica Rosa, 1893? Megadrilogica 3(10): 175-176. Gates, G.E. 1980a. Contributions to a revision of the earthworm family Lumbricidae. XXV. The genus Allolobophora Eisen, 1874, in North America. Megadrilogica 3(11): 177-184. Gates, G.E. 1980b. Contributions to a revision of the earthworm family Lumbricidae. XXVI. On two octolasia. Megadrilogica 3(12): 205-211. Harman, W.J. 1952. A taxonomic survey of the earthworms of Lincoln Parish, Louisiana. Proc. La. Acad. Sci. 15: 19-23. Harman, W.J. 1960. Studies on the taxonomy and musculature of the earthworms of central Illinois. Champaign: Univ. Illinois, 107 p. (Ph.D. dissertation) Hoffmeister, W. 1845. Die bis jetzt bekannten Arten aus der Familie der Regenwürmer. Als Grundlage zu einer Monographie dieser Familie Brauschweig. F. Vieweg und Sohn, 43 p. Jaenike, J. and R.K. Selander. 1985. On the co-existence of ecologically similar clones of parthenogenetic earthworms. Oikos 44(3): 512-514. Jaenike, J., S. Ausubel and D.A. Grimaldi. 1982. On the evolution of clonal diversity in parthenogenetic earthworms. Pedobiologia 23(3-4): 304-310. Jaenike, J., E.D. Parker, Jr. and R.K. Selander. 1980. Clonal niche structure in the parthenogenetic earthworm Octolasion tyrtaeum. Amer. Nat. 116: 196-205. Linnaeus, C. 1758. Systema Naturae. Regnum Animale (10th ed.). 824 p. Malm, A.W. 1877. Om daggmasker, Lumbricina. Öfv. Salsk. Hortik. Vanners Göteborg Förh. 1: 34-47. Macnab, J.A. and D. McKey-Fender. 1947. An introduction to Oregon earthwqorms with additions to the Washington list. Northwest Sci. 21(2): 69-75. Michaelsen, W. 1900. Oligochaeta. In: Das Tierreich. Leif 10. Berlin: Verlag von R. Friedländer und Sohn, 575 p. Moore, H.F. 1895. On the structure of Bimastos palustris, a new oligochaete. J. Morph. 10(2): 473-496. i , N. 1987a. Alpodinaridella gen. nov. (Lumbricidae) and description of two new monotypic genera. Biol. V stn. 35(2): 61-66. , N. 1987b. Descripotion of a new genus and five species of earthworms (Oligochaeta: Lumbricidae). Scoploia 13(0): 1-11. , N. 1990. Description of a new subgenus, three new species and taxonomic problems of the genus


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Allolobophora stn. 38(1): 49-68. , N. 1991. Monograph on earthworms (Lumbricidae) of the Balkans. Slovenska Akademija Znanosti Umetnosti, Ljubljana, 757 p. , Allolobophora Eisen 1874 (emend. Pop 1941) (Lumbri cidae, Oligochaeta). Acta Mus. Mac. Sci. Nat. 1(154): 1-38. Olson, H.W. 1928. The earthworms of Ohio, with a study of their distribution in relation to hydrogen-ion concentration, moisture and organic content of the soil. Bull. Ohio Biol. Surv. 4(2), Bull. 17: 47-90. Olson, H.W. 1936. Earthworms of Missouri. Ohio J. Sci. 36(2): 102-193. Oslon, H.W. 1940. Earthworms of New York state. Amer. Mus. Nov., No. 1090, 9 p. Omodeo, P. 1956. Contributo all revisione dei Lumbricidae. Arch. Zool. It. 41: 129-212. Omodeo, P. 1988. The genus Eophila (Lumbricidae, Oligochaeta). Boll. Zool. 55(1/2): 73-84. Omodeo, P. 1989. Earthworms of Turkey. Boll. Zool. 56(1/2): 167-199. Örley, L. 1882. A palearktitus ovben elo terrikolaknak revizioja es elterjedese. Ertek. Term Magyar Akad. 15(18): 1-34. Perel, T.S. 1973. The shape of nephridal bladders as a taxomonic character in the systematics of Lumbricidae. Zool. Anz. 191(5/6): 310-317. Perel, T.S. 1976. A critical analysis of the Lumbricidae genera system. Rev. Écol. Biol. Sol 13(4): 635-643. Perel, T.S. 1979. Rasprostranenie i zakonomernosti raspredelenia do devyh ervej fauny SSSR. [Range and regularities in the distribution of earthworms of the USSR fauna] Nauka, Moskva, 268 p. Pop, V. 1941. Zur Phylogenie und Systematik der Lumbriciden. Zool. Jb. Syst. 74: 487-522. Reynolds, J.W. 1972a. A contribution to the earthworm fauna of Montana. Proc. Mont. Acad. Sci. 32: 6-13. Reynolds, J.W. 1972b. [1973] Earthworm (Annelida, Oligochaeta) ecology and systematics. In Dindal, D.L., ed. Proc. 1st Soil Microcommunities Conf. Springfield: U.S. Atomic Energy Commn., Natl. Tech. Inform. Serv., U.S. Dept. Comm., pp. 95-120. Reynolds, J.W. 1973a. The earthworms of Delaware (Oligochaeta: Acanthodrilidae and Lumbricidae). Megadrilogica 1(5): 1-4. Reynolds, J.W. 1973b. The earthworms of Rhode Island (Oligochaeta: Lumbricidae). Megadrilogica 1(6): 1-4. Reynolds, J.W. 1973c. The earthworms of Connecticut (Oligochaeta: Lumbricidae, Megascolecidae and Sparganophilidae). Megadrilogica 1(7): 1-6. Reynolds, J.W. 1974. The earthworms of Maryland (Oligochaeta: Acanthodrilidae, Lumbricidae, Megascolecidae and Sparganophilidae). Megadrilogica 1(11): 1-12. Reynolds, J.W. 1975a. Die biogeografie van Noorde-Amerikaanse (Oligochaeta) noorde van Meksiko - I. Indikator 7(4): 11-20. Reynolds, J.W. 1975b. The earthworms of Prince Edward Island (Oligochaeta: Lumbricidae). Megadrilogica 2(7): 4-10. Reynolds, J.W. 1976a. Die biogeografie van Noorde-Amerikaanse (Oligochaeta) noorde van Meksiko - II. Indikator 8(1): 6-20. Reynolds, J.W. 1976b. The distribution and ecology of the earthworms of Nova Scotia. Megadrilogica 2(8): 1-7. Reynolds, J.W. 1976c. A preliminary checklist and distribution of the earthworms of New Brunswick. N.B. Nat. 7(2): 16-17. Reynolds, J.W. 1976d. Catalogue et clé d'identification des lombricidés du Québec. Nat. can. 103(1): 21-27. Reynolds, J.W. 1977a. The earthworms of Massachusetts (Oligochaeta: Lumbricidae, Megascolecidae and Sparganophilidae). Megadrilogica 3(2): 49-54. Reynolds, J.W. 1977b. The earthworms (Lumbricidae and Sparganophilidae) of Ontario. Life Sci. Misc. Publ., Roy. Ont. Mus., 141 p. Reynolds, J.W. 1994a. The distribution of the earthworms (Oligochaeta) of Indiana: a case for the Post Quaternary Introduction Theory for megadrile migratiuon in North America. Megadrilogica 5(3): 13-32. Reynolds, J.W. 1994b. [1995] The distribution of earthworms (Annelida, Oligochaeta) in North America. Pp. 133-151 In Mishra, P.C., B.K. Senapati, N. Behera and B.C. Guru (eds.) Advances in Ecology and Environmental Sciences. New Delhi: Ashish Publ. House, 651 pp. Reynolds, J.W. 1994c. Earthworms of North Carolina (Oligochaeta: Acanthodrilidae, Komarekionidae, Lumbricidae,

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Megascolecidae, Ocnerodrilidae and Sparganophilidae). Megadrilogica 5(6): 53-72. Reynolds, J.W. 1994d. Earthworms of Virginia (Oligochaeta: Acanthodrilidae, Eudrilidae, Lumbricidae, Megascolecidae and Sparganophilidae). Megadrilogica 5(8): 77-94. Reynolds, J.W. 1994e. Earthworms of Florida (Oligochaeta: Acanthodrilidae, Eurdilidae, Lumbricidae, Megascolecidae, Ocnerodrilidae, Octochaetidae and Sparganophilidae). Megadrilogica 5(12): 125-141. Reynolds, J.W. 1994f. Earthworms of Mississippi (Oligochaeta: Acanthodrilidae, Lumbricidae, Megascolecidae, Ocnerodrilidae and Sparganophilidae). Megadrilogica 6(3): 21-33. Reynolds, J.W. 1994g. Earthworms of Alabama (Oligochaeta: Acanthodrilidae, Eudrilidae, Lumbricidae, Megascolecidae, Ocnerodrilidae and Sparganophilidae). Megadrilogica 6(4): 35-47. Reynolds, J.W. and M.J. Clapperton. 1996. New earthworm records for Alberta (Oligochaeta: Lumbricidae) including the description of a new Canadian species. Megadrilogica 6(8): 73-82. Reynolds, J.W. and D.G. Cook. 1976. Nomenclatura Oligochaetologica, a catalogue of names, descriptions and type specimens of the Oligochaeta. Fredericton: Universitry of New Brunswick, 217 p. Reynolds, J.W. and D.G. Cook. 1981. Nomenclatura Oligochaetologica Supplementum Primum, a catalogue of names, descriptions and type specimens of the Oligochaeta. Fredericton: Universitry of New Brunswick, 39 p. Reynolds, J.W. and D.G. Cook. 1989. Nomenclatura Oligochaetologica Supplementum Secundum, a catalogue of names, descriptions and type specimens of the Oligochaeta. New Brunswick Mus. Monogr. Ser. (Nat. Hist.) No. 8, 37 p. Reynolds, J.W. and D.G. Cook. 1993. Nomenclatura Oligochaetologica Supplementum Tertium, a catalogue of names, descriptions and type specimens of the Oligochaeta. New Brunswick Mus. Monogr. Ser. (Nat. Hist.) No. 9, 39 p. Reynolds, J.W. and P.N. Mayville. 1994. New earthworm records from Rainy River District in north western Ontario (Oligochaeta: Lumbricidae). Megadrilogica 6(2): 13-17. Reynolds, J.W. and K.W. Reynolds. 1992. Les vers de terre (Oligochaeta: Lumbricidae et Sparganophilidae) sur la rive nord du Saint-Laurent (Québec). Megadrilogica 4(9): 145-161. Reynolds, J.W. and H.I. Savard. 1996. The earthworms of New Bruinswick/Les Lombricides du NouveauBrunswick. N.B. Mus., Publ. Nat. Sci., No. 12 (in prep.). Reynolds, J.W. and M.J. Wetzel. 1994. [1996] North American Megadriles (continental North America north of Mexico). In Coates, K.A., M.J. Wetzel, J. Madill, S.R. Gelder, J.W. Reynolds and R.O. Brinkhurst (eds.) Common and scientific names of aquatic invertebrates from the United States and Canada. Annelida. Amer. Fish. Soc. Spec. Publ. No. 17 (in press). Reynolds, J.W., E.E.C. Clebsch and W.M. Reynolds. 1974. The earthworms of Tennessee (Oligochaeta). I. Lumbricidae. Bull. Tall Timbers Res. Stn., No. 17: 1-133. Rosa, D. 1893. Revisione dei Lumbricidi. Mem. Acc. Torino, ser. 2, 43: 399-476. Savigny, J.C. 1826. Analyse d'un Mémoire sur les Lombrics par Cuvier. Mém. Acad. Sci. Inst. France 5: 176184. Scheu, S. and M.A. McLean. 1993. The earthworm (Lumbricidae) distribution in Alberta (Canada). Megadrilogica 4(11): 175-180. Sims, R.W. 1983. The scientific names of earthworms. pp. 467-474. In: Earthworm ecology from Darwin to vermiculture (Ed. J.E. Satchell). London: Chapman and Hall, 495 p. Smith, F. 1917. North American earthworms of the family Lumbricidae in the collections of the United States National Museum. Proc. U.S. Natl. Mus. 52: 157-182. Snider, R.M. 1991. Checklist and distribution of Michigan earthworms. Mich. Academician 24: 105-114. Stephenson, J. 1930. The Oligochaeta. Oxford: Clarendon Press, 978 p. Svetlov, P. 1924. Beobachtungen über die Oligochäten des Gouv. Perm. I. Zur Systematik, Fauna und Ökologie der Regenwürmer. Bull. Inst. Rech. Biol. Perm 2:(8): 315-328. Tandy, R.E. 1969. The earthworm genus Pheretima Kinberg, 1866 in Louisiana. Baton Rouge: Louisiana State Univ., 155 p. (Ph.D. dissertation) Zicsi, A. 1978. Revision der Art Dendrobaena platyura (Fitzinger, 1899) (Oligochaeta: Lumbricidae). Acta Zool.


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Hung. 24(3-4): 439-449. Zicsi, A. 1981. Probleme der Lumbriciden-Systematik sowie die Revision zweier Gattungen. Acta Zool Hung. 27(5-6): 431-442. Zicsi, A. 1982. Verzeichnis der bis 1971 beschriebenen und revidierten Taxa der Familie Lumbricidae (Oligochaeta). Acya Zool Hung. 28(3-4):421-454. Zicsi, A. 1985. Über die Gattungen Helodrilus Hoffmeister, 1845 und Proctodrilus gen. n. (Oligochaeta:Lumbricidae). Acta Zool Hung. 31(1-3): 275-289. List of Figures Figure 1.

The North American distribution of Aporrectodea trapezoides, Ap. tuberculata and Ap. turgida.

Figure 2.

The North American distribution of Dendrobaena octaedra and Dendrodrilus rubidus.

Figure 3.

The North American distribution of Lumbricus rubellus and L. terrestris.

Figure 4.

The North American distribution of Octolasion tyrtaeum.

Figure 5.

The limited North American distribution of some miscellaneous lumbricids (Aporrectodea icterica, Ap. limicola, Bimastos palustris, B. welchi, Dendrobaena attemsi and Eisenia zebra). (The numbers in parentheses represent the number of collection sites.)

Figure 6.

The limited North American distribution of some miscellaneous lumbricids (Allolobophoridella eiseni, Dendrobaena pygmaea, Lumbricus festivus, Murchieona muldali and Satchellius mammalis). (The numbers in parentheses represent the number of collection sites.)

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NOMENCLATURE (NOMENCLATURA) We cannot dispense with names. they are an essential part of the equipment with which we must work . . . anyone who is properly to be called a systematist must consider it a part of his/her equipment . . . . (Ferris, G.F. 1928. The principles of systematic entomology. Stanford, CA: Stanford Univ. Press)

Zoological nomenclature is a complex and specialized subject. Names are given to animal groups, and used to refer to them, under rules which are neither simple nor unchanging. The present rules are the result of a long history or evolution, during which there was much discussion of the various solutions and special cases. History (Historia) !

1901, 5th Intl. Zool. Congress (Berlin) adopted International Rules of Zoological Nomenclature (Règles Internationales de la Nomenclature zoologique); few subsequent changes until 1948; problems taken care of by interpretative opinions

!

1935, 6th Intl. Congress of Entomology (Madrid); emergence of Francis Hemming

!

1948, 13th Intl. Congress of Zoology (Paris); proceedings published in Bull. Zool. Nomencl. vol. 4; amended rules effective as soon as published; spent much time in discussing a series of changes in the rules, amounting to a revision of the entire code; Hemming now Secretary of Commission

!

1953, 14th Intl. Zool. Congress; conclusions published as Copenhagen Decisions on Zoological Nomenclature; first to be sold at a price most could afford (135 p. on 8.5" x 11"); strong admonition (warning) to follow the rules; Rudolf Richter not invited for political reasons; Hemming refused to have any Americans appointed to any committees;

!

1958, 15th Intl. Congress of Zoology (London); made decision on preparation of the new code; included a long list of particular decisions on individual problems

!

1961, International Code of Zoological Nomenclature adopted by the XV International Congress of Zoology; published in English and French; Code editors' statement: "the majority views of interested zoologists throughout the world." [Hemming retired in 1960.]

!

1964, first revision to Code; Commission has exaggerated its authority from advising to ruling.

!

1985, 3rd revision to Code; International Code of Zoological Nomenclature adopted by the XX General Assembly of the International Union of Biological Sciences (Helsenki, 1979)


75

The three major proposals for the 3rd (current) edition were: 1. 2. 3.

to provide a single word for the different categories of types that could be applied equally to all name bearers, i.e. "name-bearing type" was accepted "onomato-phore" was rejected. to provide a single word to mean the second word of a binomen and the second and third word of a trinomen, "epithet" as used in the ICBN was rejected. to dispense with the concept "nominal taxon" and treat name-bearing types as types like ICBN, but this was rejected.

International Commission on Zoological Nomenclature (Comisión Internacional de Nomenclatura Zoologica) Duties: 1. 2. 3. 4. 5. 6. 7.

to consider for a period of at least one year in advance of a Congress (or for such less time as the Commission may agree) any proposal for a change in the Code; to submit to the Congresses recommendations for the clarification or modification of the Code; to render between successive Congresses Declarations (i.e., provisional amendments to the Code) embodying such recommendations; to render Opinions and Directions on questions of zoological nomenclature that do not involve changes in the Code; to compile the Official Lists of accepted, and the Official Indexes of rejected, names and works in zoology; to submit reports to the Congresses on its work; and to discharge such other duties as the Congresses may determine.

Complexity of the Rules (Complejidad de las Reglas) ! ! ! !

many felt the rules were unnecessarily detailed the accumulation of interpretative Opinions made it more difficult by 1948 there were Opinions on Opinions often contradicting one another the following is generally recognized by zoologists:

1. 2. 3. 4.

After 1960 all decisions must be based on the new Code. Decisions between 1901-1953 under the Règles cannot all be upset by the new Code. Decisions between 1953-1961 under the Copenhagen Decisions cannot be upset by the new Code. No decision under any of these Codes is of any value unless it is determined that there has been no contrary action by the Commission under its Plenary Powers. the 1964 revision of the 1961 Code incorporated changes in at least 4 rules. None of the 12 works is adequately indexed for an inexperienced nomenclaturalist. Further editions will continually appear from time to time. In at least 30 places, individual rules apply only between certain dates, making it necessary to follow the history of a name through successive forms of the rule. The extent of mixture of taxonomy into supposedly strictly nomenclatural rules is much more than the amount necessary for regulation of names. This does not interfere with taxonomic work in many respects. Any decision taken in good faith by any taxonomist now stands in greater danger of being reversed by the Commission, because of the established practice of acting in response to individual applications.

5. 6. 7. 8. 9. 10.

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Valid and Available Names (Nombres válidos y Disponibles) valid name - is defined in the code as "The correct name for a given taxon; a taxon may have several available names, but only one of those names . . . is the valid name." available name - is "One that satisfies the provisions of the Code as to publication, formation, and description." Hierarchy of name categories All Latin names Published

Not published

Occupied

Nomina nuda

Available

Pre-occupied (junior homonyms)

Valid (correct)

Invalid (rejected)

Nomina dubia

nomen dubium. A name not certainly applicable to any known taxon. nomen novum. A new name published to replace an earlier name and valid only if the latter is preoccupied. nomen nudum. A name that fails to satisfy certain provisions/conditions of the Code. nomen oblitum. A name that remained unused as a senior synonym in the primary zoological literature for more than50 years is to be considered a forgotten name.

The system used for the naming of animals in zoology is generally described as a binomial syste (taxa) are of several sorts, which can be listed thus: 1. Names of major groups, e.g. Annelida, Oligochaeta 2. Names of family groups, e.g. Glossoscolecidae 3. Names of genera, e.g. Dichogaster 4. Names of species, e.g. Microscolex dubius 5. Names of species with subgenus, e.g. Belladrilus (Santomesia) emilianii 6. Names of subspecies, e.g. Bimastos beddardi sophiae 7. Names of subspecies with subgenus, e.g. Eutrigaster (Graffia) keiteli keiteli The use of varieties, races or forms is no longer recognized by the Code. [1-3 are uninominal; 4 is binominal; 5-6 are trinominal; 7 is quadrinominal.] The suggested endings for certain groups of taxa were covered on pages 14 and 42. ! ! !

homonyms - identical names for different things (taxa) synonyms - two names for the same things (taxa) emendations - intentional changes in spelling of a name


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Synonymy (Sinonimia) - is the relationship between different names designating the same taxon. ! ! ! ! !

applies to species and genus categories most often we will use the species category as an example but these rules, etc. apply to all categories a synonymy list will be a logical way to examine these the list is chronological beginning with the citation of the original citation; I will number the steps for reference only, they would not appear in a list a complete synonymy list would include every reference to the species; this is rare and most taxonomists would include only the major references and/or those with taxonomic and nomenclatural significance

example:

Allolobophora chlorotica (Savigny, 1826)

1

1826

2 3 4 5 6 7

1828 1837 1843 1845 1865 1873

8 9 10 11 12

1882 1884 1885 1892 1896

13 14

1900 1956

Enterion chloroticum + E. virescens Savigny, Mém. Acad. Sci. Inst. Fr. 5: 183. Types, MNHN, Paris, No. xxxx! Lumbricus anatomicus Dugès, Ann. Sci. Nat. 15(1): 289. Typus amissus. Lumbricus chloroticus-Dugès, Ann. Sci. Nat. (2), 8: 17, 19. Lumbricus riparius Hoffmeister, Arch. Naturg. 9(1): 189. Types unknown. Lumbricus communis luteus Hoffmeister, Regenwürmer, p. 29. Typus amissus. Lumbricus viridis Johnston, Cat. British non-paras. worms., p. 60. Types unknown. Allolobophora riparia-Eisen, + A. mucosa Eisen, Öfv. Vet.-Akad. Förh. Stockholm 30(8): 46, 47. Typus amissus. Allolobophora neglecta Rosa, Atti Acc. Torino 18: 170. Types unknown. Allolobophora chlorotica-Vejdovský, Syst. Morph. Oligo., p. 60. Aporrectodea chlorotica-Örley, Ertek. Term. Magyar Akad. 15(18): 22. Allolobophora cambrica Friend, Essex Nat. 6: 31. Typus amissus. Allolobophora curiosa Ribaucourt + A. Waldensis Ribaucourt + A. morgensis Ribaucourt + A. cambria (laps.)-Ribaucourt, Rev. Suisse Zool. 4: 46, 47, 94. Types unknown. Helodrilus (Allolobophora) chloroticus-Michaelsen, as Tierreich, Oligochaeta 10: 486. Allolobophora chlorotica-Omodeo, Arch. Zool. It. 41: 180.

Explanation: 1

after the location of the types; "!" means "I have seen the types", "." means "I know the types are there but I have not seen them." Savigny describes two different species but really only has one species. The one described first gets to carry the name.

2

Dugès describes a new species which later is determined to be a synonym of A. chlorotica.

3

Dugès reports chlorotica but places it in a different genus. The use of "-" between the species epithet (name) and the author denotes that this is not a new species. An alternative method is to use a "," in place of the "-".

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4

Hoffmeister describes a new species which later is determined to be a synonym of A. chlorotica.

5

Hoffmeister describes a new subspecies which later is determined to be a synonym of A. chlorotica.

6

Johnson describes a new species which later is determined to be a synonym of A. chlorotica.

7

Eisen reports the species as Hoffmeister's riparia but places it in another genus. He also describes a new species which turns out to be a synonym for chlorotica.

8

Rosa describes a new species which later is determined to be a synonym of A. chlorotica.

9

Vejdovský is the first to place the species in the genus where it now resides.

10

Örley moves chlorotica to his new genus Aporrectodea.

11

Friend describes a new species which later is determined to be a synonym of A. chlorotica.

12

Ribaucourt describes three new species which all later turn out to be A. chlorotica. He also identifies Friend's species cambrica but spells the name incorrectly.

13

Michaelsen in his reorganization of the Lumbricidae creates Allolobophora as a subgenus of Helodrilus and places chlorotica there.

14

Omodeo designates chlorotica the type species for the genus Allolobophora which later causes Örley's Aporrectodea to be resurrected.

All taxa must have a type designated, e.g. species, genus, family. Discussion Draft of the Fourth Edition of the International Code of Zoological Nomenclature (Discusión somera de la 4 Edición (to be published in 1996; effective 1 January 1997) Explanatory Notes The Current (Third) edition of the Code was published in February 1985. It was inevitable that some constructive suggestions made before that time could not be incorporated, and many others were prompted by the appearance of the new edition. In 1988 the International Commission on Zoological Nomenclature set up an Editorial Committee to work towards an eventual Fourth Edition and published an invitation to zoologists to submit further recommendations. Many have been received, and the Commission is grateful to all those who have assisted it. The Commission held open meetings in 1988 (Canberra), 1990 (Maryland) and 1991 (Amsterdam) for preliminary discussion of proposed improvements to the Code, many of them intended to meet changing needs and rapidly evolving communication techniques.


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The Editorial Committee met in Hamburg in October 1993 and reviewed each Article of the existing Code in the light of the above discussions and all the suggestions which have been made. The Committee now offers to all zoologists and other users of scientific names, and to the full Commission itself, a Discussion Draft of the Fourth Edition. Under Article 16 of its Constitution the Commission will take into full consideration all comments made on the Draft within one year of its original distribution (that is, by the end of May 1996). It is hoped that it will be possible to publish the new Edition, with the approval of the International Union of Biological Sciences, in 1996 and that its provisions will take formal effect (superseding the current Third Edition) on 1 January 1997. To achieve this timetable, and more importantly to ensure that the Code will meet the needs of its users, zoologists and others are now invited to submit comments on the Discussion Draft. The Editorial Committee has been guided by the principle that scientific names are labels for taxa and provide the only universal means of accessing zoological information. Stability in their application and form, consistent with taxonomy, is therefore of paramount importance irrespective of any priority or linguistic consideration. This aim to maintain stability must take precedence over the tools that the Code uses to promote it. Thus, while priority remains the basis for determining validity, and linguistics the basis for the formation of names, neither is an end in itself. Under the changing circumstances of science these and other means of promoting stability must be reviewed for each new Edition. This has been done and the major changes proposed reflect that view. Like all zoologists, members of the Editorial Committee recognize that many names in current use are in breach of the existing Code and that no scientific purpose would be served by continuing to make them vulnerable to change for purely formal reasons. In the proposal for the Fourth Edition every effort has been made to ensure that names in present use will remain valid when the new Code comes into effect, or that they can be easily validated. The proposed Edition differs from earlier ones in enabling zoologists, wherever possible, to adopt automatic solutions directly, rather than requiring them to defer decisions until the Commission has determined the outcome in response to formal applications. It is emphasized that in every such case the proposed "do-it-yourself' solution involves consultation with workers interested in the field. Recourse to the Commission remains in the event of disagreement and as a safeguard against abuse. The notes below are intended to draw attention to the main proposals which distinguish the Discussion draft from the current Code, and to indicate some of the reasons for making changes. The Draft itself must be consulted for details of the Articles and Recommendations. For the sake of continuity the order of Articles in the existing Code has been retained. The Draft does not contain the indexes, appendices and glossary which will be in the definitive Code, nor proposed administrative amendments to the Constitution of the Commission which have been published recently (Bulletin of Zoological Nomenclature 52: 6-10). I. Additional requirements for the availability of names first published after 1996. Several proposals are made to improve the unambiguous definition and typification of new nominal taxa and the recognition and accessibility of their names. In practice the great majority of names published in recent decades meet the suggested requirements, but these will not apply retrospectively. (a) Species-group taxa. (i) The new nominal species or subspecies must be explicitly indicated as being new [Art. 16e]. (ii) The name-bearing type (a holotype, or syntype series) of the new nominal taxon must be unambiguously designated [Arts. 16e, 72c]. (iii) The diagnosis of the taxon must include a summary of characters which are considered to differentiate it from at least one other of the same rank (i.e. related species or subspecies) which must be cited by name [Art. 16a].

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(iv) The diagnosis must be given in a language which uses the Latin alphabet; it is recommended that the diagnosis should be given in a language which is widely used, and also in those of the regions relevant to the taxon [Art. 16b]. (v) A new name must be recorded as such in the Zoological Record within five years of its first publication; if it is not, it is deemed not available from that publication [Arts. 8e, 11b]. Procedures and safeguards are recommended. Mandatory listing in the Zoological Record (which is accessible on paper, compact disk, and electronically) means that only this single source will need to be searched for the existence of new names. (b) Genus-group taxa. The new requirements are the same as those given above, with the necessary changes of wording [Arts. 11b, 16] is already required [Art. 13c], and this is now extended to new ichnogenera (genera of trace fossils) [Arts. 16d, 66]. (c) Family-group taxa. The additional requirements are entirely analogous to those mentioned above. It is proposed that the name of the type genus must be explicitly cited [Art. 16c]. II. Use of the term "epithet" in species-group names Because there has been some confusion between the "name of a species" (which is a binomen) and the "species name" (the word in the binomen which denotes the species), it is proposed that "specific name" be replaced by "specific epithet". The same applies to "subspecific -epithet". This change is in harmony with the International Code of Botanical Nomenclature. III. New provisions relating to the application of the Principle of Priority Although priority is the main criterion in determining the validity of competing names the Draft makes provisions enabling zoologists to depart from it in some situations, without the recourse to the Commission required by the present Code. (a) Conservation of junior synonyms. When a senior synonym has not been used as valid in the previous fifty years and, a junior name has been universally and widely used in this period, then the junior name is to be given precedence [Art. 23j]. (b) Conservation of subsequent spellings. Providing the same criteria as mentioned for junior synonyms are met, a subsequent spelling of a name which is different from that first published is to be accepted as the correct original spelling [Art. 33d]. (c) Usage of family-group names contrary to priority. If two family-group names are in general current use such that the taxon denoted by the senior name (e.g. a subfamily) is included within that (e.g. a family) denoted by the junior name, such usage is to continue even though it is contrary to priority [Art. 35e]. V. New provisions relating to the typification of nominal taxa The additional criteria for a name published after 1996 to be available include some relating to type fixation, and these have been mentioned under I above. The draft contains other provisions which relate to typification but do not affect the availability of names, and which propose solutions independent of reference of cases to the Commission. (a) Acceptance of name-bearing types found to have been misidentified.


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(i) family-group taxa: if a zoologist discovers that the type genus of a family-group nominal taxon has been misidentified when the taxon was established, or that there were errors or overlooked fixations in the typification of the type genus itself, the erroneous nominal fixations actually made should be accepted unless stability is threatened [Art. 41a]. (ii) genus-group taxa; analogous provisions apply [Art. 70b]. (c) Status of neotypes following rediscovery of original type material. A validly designated neotype is to be retained as the name-bearing type of a species-group taxon if rediscovery of original type material causes no instability [Art. 75j]. V. New provisions concerning the grammar and spelling of names Zoologists have spent much time debating matters which are purely of grammar or spelling, and many destabilizing name changes have been caused as a result. Very few modern zoologists are at ease with Latin, although this was the language of international communication to Linnaeus and his successors (who, even so, were not always rigorous in their grammatical practices). Even fewer have any knowledge of classical Greek. The Discussion Draft attempts (i) to respect the names of the past but to preserve them in the forms in which they have been used in modem times, (ii) to avoid name changes, i.e. obstacles to information retrieval, made for non-taxonomic reasons, and (iii) not to regulate or "correct" the spelling of new names. The new provisions will no doubt be controversial, but the Editorial Committee hopes that criticism of them will be for zoological or practical reasons and not simply on linguistic or historical grounds. The following changes are proposed. (a) Gender of generic names. It is proposed that after 1996 generic names should be treated as words having no gender and therefore not affecting the spelling of adjectival specific epithets (e.g. albus, -a, -um) combined with them [Art. 30]. Consequent on this, the Editorial Committee offers two alternatives [see Arts. 31b, 32c and 48] for discussion, as follows. Either: (i) After 1996 the original ending of such an epithet is to be used in all combinations, whether or not the combination is new and whether or not a change in an existing binomen results; Or: (ii) the ending of such an epithet is to remain as it is in an existing combination (so a binomen already in use for a species remains unchanged), but in new combinations first published after 1996 the original ending of the epithet is to be used. (b) Acceptance of incorrect spellings. (i) The original spelling of an adjectival species-group epithet first published after 1996 should be accepted as correct, even if its gender ending is grammatically improper in the original combination [Art 31b(iii)]. (ii) Certain endings of species-group epithets that are formed from personal names are to be treated as identical; thus spellings such as smithi and smithii are permissible variants [Art. 31b(v)]. (iii) If an incorrect spelling of a name has been generally accepted that spelling is not to be changed [Arts. 29d, 33d]. (iv) A new family-group name may be formed by adding the appropriate ending (e.g. -IDAE, -INAE) to the entire name of the type genus, rather than only to its stem [Art. 29a, c]. This may be necessary to avoid the new familygroup name being a junior homonym of one based on another generic name which has the same stem. The spelling of a new familygroup name should not be amended by reason of having a grammatically incorrect stem [Art. 29c].

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If a disused family-group name is a senior homonym of one in use its stem may be emended so as to conserve the junior name [Art. 55c]. VI. Adoption of Lists of Available and Potentially Valid Names In some taxonomic fields workers may wish to establish Lists of names at particular ranks (e.g. the family-, genus- or species-groups) which will automatically take precedence over any relevant names not listed [Art. 78j], so that the listed names and name-bearing types may be used with confidence (taxonomic validity being left, as always, to individual judgement). Procedures are proposed [Art. 77] for the adoption by the Commission of such Lists, it being made clear that adoption will be only in response to initiatives by international bodies of zoologists interested in the relevant taxonomic fields and that extensive consultations will always be needed before the adoption of a List. The Preface to the current (1985) edition points out (p. xii): "No Code is perfect. None will please everyone. Indeed, it is unlikely that any Code would be completely satisfactory to any individual". Some of the innovations proposed for the Fourth Edition will be controversial, but the Editorial Committee believes that the Discussion Draft provides a significant advance for the future while respecting and preserving the names of the past. We urge zoologists to test the changes proposed by the Editorial Committee constructively and without prejudice, with the needs of the wider biological community in mind, and with awareness of the changed circumstances of taxonomists today. We ask those who are not specialists but who use scientific names, in whatever way, to make suggestions so that the Code will meet their needs. We are aware that we are asking those who will comment on the proposals and those who will vote on their adoption, who are mostly specialists located in long-established centers, to review proposals sympathetically even though many will consider them unnecessary in their own situations. The most important of these changes, if adopted, will widen the medium of publication beyond print media into an electronic age, will reduce dependence upon expensive and comprehensive library holdings and ancient works for nomenclatural searches, and will make familiarity with classical Latin and Greek grammar unnecessary. We look forward to receiving suggestions that will improve the proposal, and for support that we are confident will result in the Commission and the International Union of Biological Sciences adopting on behalf of zoologists and users of scientific names a Fourth Edition that will more effectively meet the needs of the 21st century than could its predecessor. 0. Kraus President The International Commission on Zoological Nomenclature


W. D. L. Ride Chairman, Editorial Committee

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EARTHWORM ECOLOGY (ECOLOGIA DE LAS LOMBRICES DE TIERRA)

Earthworm ecology is a very broad topic and is frequently a course in itself. During our treatment of this topic, we will concentrate on only four aspects: ! ! ! !

Habitat requirements, Barriers to migration, Functions of earthworms in the soil, and Earthworm ecological classification systems.

Everyone thinks of earthworms as living in the soil but it is more specific than that. There is a well known concept called the niche utilization theory: no two species can live in the same place at the same time doing the same thing. One eventually adapts better to the exclusion of the other. So although it may seem there are a number of species in our backyard, garden, compost, forest, etc., on closer examination it becomes apparent that each species is exploiting a different part of the habitat. Two good examples of this from North America, are: 1) the removal of Dendrobaena octaedra and Dendrodrilus rubidus from the under log and under bark habitats above the southern limit of the glacial boundary and their replacement by species of the nearctic Bimastos south of this boundary; 2) in the native forests of western United States where several species of endemic Acanthodrilidae (e.g. Arctiostrotus spp.) are found, once clear-cutting takes place and cultivation (agriculture) takes place, common European Lumbricidae, e.g. Aporrectodea trapezoides take over these distinctive habitats. As we will see in the section on Earthworm Biogeography, earthworms are distributed around the world in their native habitats and, in the case of some families, to new areas where they have been transported by humans and other animals. Earthworms' ability to migrate any distance by their own means is limited by several environmental requirements for their successful habitats. These are animals which have no limbs, no eyes, and require a moist skin in order to breathe (Reynolds, 1973, 1994). Habitat Requirements (Requerimientos habitacionales) The following conditions are required in a habitat in order for earthworms to survive:

1. Adequate and suitable food supplies As feeders of dead or decaying plant remains, and sometimes dead animal remains, earthworms remain close to their food source.

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Figure 2.

Earthworm species' habitats and their availability to the American woodcock (Philohela minor) in northern North America. (1 - Lumbricus terrestris noctural habitat; 2 - L. terrestris diurnal habitat; 3 - surface soil species which could be available to woodcock; 4 - corticole species - Dendrodrilus rubidus - generally unavailable to woodcock and; 5 - limicolous species - Eiseniella tetraedra - possibly available to woodcock, if present). [From Reynolds, J.W. 1977. Earthworms utilized by American woodcock. Proc. Woodcock Symp. 6: 161-169.]


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2. Adequate moisture Because earthworms breathe through their outer skin layer, a moist habitat is essential to allow respiration to occur. 3. Adequate dissolved oxygen Earthworms are air-breathing animals, and therefore, dissolved oxygen in their habitat is essential for respiration. Earthworms have lived submerged for up to a year in oxygen-rich environments. Death eventually occurred from starvation and not suffocation. 4. Protection from sunlight Ultra-violet radiation is lethal to earthworms in a very short time period. 5. Suitable pH Some acidity is not a problem, but it is difficult for the earthworms to extract nutrients from a food source under very acidic (high pH) or very basic (low pH) conditions. In these two situations, the necessary chemical reactions that permit digestion to occur are inhibited. The presence of calciferous glands help buffer reasonable acidic conditions so that some species can live under conditions of pH 2.8-4.5. Most species are found in habitats where the pH is 4.5-7.5 and none have been found where the pH exceeds 9. 6. Absence of toxic substances Habitats with high concentrations of various salts do not contain earthworms. Many pesticides have little or no chemical effect on earthworms. In fact, they can survive under conditions where the concentrations are 20 times the lethal limits for other species. There has been much debate between scientists and organic gardeners about the effects of pesticides on earthworms. A recent general summary of some common pesticides follows (Reynolds, 1992):

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1)

Azinphosmethyl, diazinon, malathion, and menazon appear to have no effect on earthworms.

2)

In normal doses, Aldrin, BHC, carbofuran, chlorfenvinphos, DDT, Dieldrin, disulfoton, Dyfonate, Endrin, fenitrothion, Teladrin, and trichlorophon have very little effect on earthworms.

3)

In large doses, Aldicarb, carbofuran, and Parathion are moderately toxic.

4)

Carbaryl, Chlorodane, Heptichlor, and Phorate are toxic to earthworms.

5)

There are also some substances, causing osmotic or water balance responses, that are lethal to earthworms even in small doses.

J.W. Reynolds

Figure 1.

Vertical distribution of earthworms in the soil profile. [From Reynolds, J.W. 1973. Proc. 1st Soil Microcommunities Conf., p. 103.]


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7.

Suitable soil temperatures The optimal soil temperature range for the development of most earthworm species is 1220 C. Certain species, such as Eisenia foetida, have a much higher optimal developmental temperature, and hence are very successful when used for vermi-composting in North America (Reynolds and Eggen, 1993). In Africa, Eudrilus eugeniae is used for this purpose, while in India and its neighbours Dichogaster bolaui and Perionyx excavatus are employed in vermi-composting (Dash and Senapati, 1986). The major advantage of worm composting over conventional methods is speed or turnover rate. When the proper earthworm species are used, the time required for decomposition can be reduced to one third.

Barriers to Migration (Barreras de Migración) The barriers to migration are tied closely to the habitat requirements discussed above: 1.

Mountain ranges The shallow soil or bare rock of mountain habitats means earthworms are exposed to unfavourable conditions of drying, ultra-violet radiation, reduced oxygen, and limited suitable food sources.

2.

Deserts Limited food sources and a severe reduction in available moisture in the desert environment mean earthworms may become inactive or estivate.

3.

Salt water Most earthworms can't survive in salt water or soils saturated with salt water. Pontoscolex corethrurus, a tropical species, which has invaded the southern tip of Florida (USA) from the Neotropical Region, seems to be able to tolerate a certain level of brackish conditions.

4.

Climate Extremely hot dry areas will restrict earthworms; under favourable climatic conditions earthworms go into quiescent periods, see Deserts and Ice/snow.

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5.

Ice or snow areas Prolonged periods of ice or snow may restrict available food sources, available moisture, and the earthworms may become inactive or hibernate.

6.

Competition New species invading an area may not be able to adapt to native species already in place, or conversely may adapt better to the habitat. An example is found in North America, where native species of Bimastos utilize the under bark habitat of decaying trees below the glacial boundary. North of the glacial boundary, these habitats are exploited by Dendrodrilus rubidus in the absence of Bimastos spp. South of the glacial boundary, D. rubidus, when present, is forced out into the soil surrounding these decaying trees.

7.

Parasites or predators There are a large number of organisms that prey on earthworms and restrict their movement and/or establishment in a habitat. These include various insects (Pollenia rudis), mites (Uropodans agitans), certain centipedes, snakes, salamanders, frogs, certain birds (robins, woodcock), and small mammals (moles, voles, and shrews).

Functions of Earthworms in the Soil (Funciones de las Lombrices en el Suelo) Although earthworms may be found in the soil and adjacent habitats, there is considerable diversity in their activities and functions in these habitats. The following list of annotated functions are the main ones: 1.

Organic matter decomposition Some species (e.g. Lumbricus terrestris) actually take the leaves and surface debris and physically or mechanically break them apart, which enables micro-organisms to complete the decomposition. Other species, which are "soil eaters" (e.g. Aporrectodea turgida), chemically alter the organic matter as it passes through their intestines. The breakdown and incorporation of organic matter into the soil increases its water-holding capacity for use by other plants and animals.

2.

Soil neutralization As soil is passed through the digestive tract of the earthworms, their calciferous glands secrete calcium compounds that raise the pH of the material nearer to the neutral state. The less acidic the soil, the more easily chemical reactions take place. These reactions are necessary for most aspects of plant and animal life in the soil, e.g. incorporation of nutrients into plant roots.

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3.

Soil aggregation As the small soil particles pass through the earthworms' intestines, they are bound together with a mucus substance into larger masses of particles. This improves the structure of the soil, as well as incorporating the decomposed organic matter into these masses. These larger masses or aggregates (fecal material) are frequently referred to as "castings".

4.

Soil aeration By eating their way through the soils and producing larger aggregates, channels develop in the upper soil horizons, which permit the increased movement of air, specifically oxygen, into the soil. The presence of adequate oxygen in the upper soil layers is essential for the growth and development of all aerobic plants and animals.

5.

Infiltration of water into the soil The channelling and aggregate production by earthworms increase the ability of water to penetrate the soil surface into the upper layer. Infiltration is one of two important aspects of soil water movement.

6.

Percolation of water within the soil The second aspect is percolation, the movement of water within the soil. The activity of earthworms increases the ability of soil water to move down and throughout the soil horizons. The availability of soil water at the level of the plant roots is important for plant growth and development.

7.

Soil turnover Soil turnover is the amount of soil that passes through the body of an earthworm in a certain time period. Experiments in North America and Europe have measured this activity and found it to be 85-120 mg of soil per day per worm. All habitat factors being equal, the range is primarily due to the varying sizes of different earthworm species, in that the amount of soil passing through an earthworm's digestive system per day represents approximately 20-30% of its live body weight.

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Oh, I thought about getting a dog but an earthworm is so much easier to feed and look after. Oh, yo pensé conseguir un perro pero una lombríz de tierra es más fácil de alimentar y cuidar.

Fuere!! Yo soy importante para el ambiente.

Worms and the Environment Lombrices de tierra y Ambiente


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Earthworm Ecological Classification Systems (Sistema de Clasificación Ecológica de las Lombrices de Tierra) Little work has been done on the ecology and distribution of Canadian earthworms and it remains an open field of investigation. From a study of 14 species in Maine (USA), Gates (1961) concluded that dietary preferences might be an important factor influencing distribution. He divided these species into three groups. The first group comprised those species that pass much soil through the intestine and hence are termed geophagous, viz. Allolobophora chlorotica, Aporrectodea longa, Ap. rosea*, Ap. tuberculata, Ap. turgida*, Lumbricus terrestris and Octolasion cyaneum* (* present in Argentina). These species are known from soils of pH as low as 4.5-5.5 and the kinds of soil seem to be of little significance. Within this group only L. terrestris normally and regularly copulates at the surface. Because these are tolerant of different soil types and occur in a wide range of habitats they are likely to be widely distributed, aided especially by the activities of man. The second group, containing only Eiseniella tetraedra* and possibly Al. chlorotica (in Maine), is limiphagous (mud-eating) or limicolous (mud-inhabiting). Another species, Sparganophilus eiseni, has been reported from Ontario (Reynolds, 1977) and Quebec (Reynolds and Reynolds, 1992). The members of this group thrive in mud well under water or in saturated soils. Their typholosoles are greatly reduced as a modification to this type of habitat. The third group consists of the litter feeders: Eisenia foetida*, Dendrobaena octaedra, Dendrodrilus rubidus*, Lumbricus castaneus, and L. rubellus. Litter includes all types of accumulation of organic matter such as leaves, manure, compost, etc. The activities of farmers and horticulturalists greatly affect the distribution of these forms, and also, of course, of geophagous species that may stray into organic matter. In contrast, prior to hibernation, litter feeders become geophagous when they burrow down to the levels where they hibernate. The extent to which Julin's (1949) ecological classification of the Lumbricidae can be applied to North America is limited. Only Reynolds et al. (1974), Reynolds (1977) and Reynolds and Savard (1996) have attempted to apply the system to North American regions, viz. Tennessee, Ontario and New Brunswick, respectively. Considering only those species in common with these three regions, they placed Al. chlorotica, Ap. rosea*, Dd. octaedra, Dd. rubidus*, E. foetida*, El. tetraedra*, L. rubellus and O. cyaneum* among the hermophiles (occurrence appears to be strongly tied to human activity). The hemerodiaphores (widespread occurrence of these species indicates their indifference to human activity) include Ap. trapezoides*, Ap. tuberculata, Ap. turgida*, and O. tyrtaeum*. Information was lacking for Ap. longa, L. castaneus and L. terrestris. L. festivus was not found in Tennessee and only one specimen in New Brunswick.

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95

There were no species which appear to be true hermophobes (those species favoured by culture) or hemerobiont (those species entirely dependent on culture). In New Brunswick, E. foetida might be considered a hemerobiont (Reynolds and Savard, 1996). Five species (36%) were found so rarely that they really can't be classified in New Brunswick: Al. chlorotica, Ap. longa, Ap. trapezoides, L. castaneus and L. festivus. Bouché (1972) developed another ecological classification which is based on habitat and morphological features common within the groups: Épigé (Epigeics) forms Morphological and physiological characters 1. 2. 3. 4. 5. 6. 7. 8. 9. 10.

Colour normally red or pinkish (litter adapation). Reduced "dissepimentaire" musculature (no pedogenetic activity). Partial musculature generally pennate (feather-shaped). Mobility often important (adaptation to flight before predators). Respiration rate relatively elevated. Ability to regenerate generally reduced (predation total). Nephropores arranged in solfeggi (evaporative cooling?). No diapause, only quiescence. Normal tail. Small body.

These are worms that normally live outside the mineral substrate of the soil, but are found in the litter (straminicoles), in decaying logs (corticoles), in crevices, burrows, galleries (pholeophiles) and in some compost and dung piles. Most often they are eliminated by deforestation. The characteristics are all the more prominent since the species develop freely above the level of the mineral horizons. The corticoles present at times a fasculate musculature and often a flattened body. Selective predator pressure may have induced the development of characters 1, 3, 4, 6, 8 and 10; and characters 2, 3, 4, 5, 7, 9 and 10 are evolutionary adaptations to terrestrial soil surface micro-habitats with temporarily favourable climates. Chief among the straminicoles Lumbricus castaneus, L. rubellus, Dendrobaena octaedra; amongst the corticoles Eisenia foetida, Dendrodrilus rubidus. Pholeophiles are not known to inhabit Canada.

Anécique (Anecics) forms Morphological and physiological characters 1. 2. 3. 4. 5. 6. 7. 8.

96

Colour normally brown to blackish brown sometimes iridescent. Very developed "dissepimented" musculature, infundibuliform (intensive predogenetic activity, grasping food). Radiating parietal musculature, often pennate. Contractability well developed (retreats rapidly into burrows). Ability to regenerate developed (predation partial). Nephropores arranged on "solfeggi" (evaporative cooling). Diapause. Lanceolate tail.

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97

9.

Body medium to gigantic.

These worms live in the soil but "hang-out" in the litter at night which they pull into their galleries (often vertical) by grasping the burrow opening with their prehensile (capable of grasping) tail. These animals will remain following deforestation and ploughing. They characterize intense pedogenetic activity by their ability to dig deep and eject the burrowed soils at the surface (castings). Predator pressure may have induced characters 1, 4, 5 and 8; characters 1, 2, 3, 6, 8 and 9 from surface activities; the characters 2 and 7 from their partially endogee habits. Most of the Aporrectodea belong to this category. Éndogé (Endogeics) forms Morphological and physiological characters 1. 2. 3. 4. 5. 6. 7. 8. 9.

No cutaneous pigmentation (albinoism). Well developed "disspimentaire" musculature, infundibuliform (rhizophilus) or not (geophilus). Variable parietal musculature. Cell-wall varying in thickness from normal to mediocre (épiendogés) or very fine (hypoendogés). Ability to regenerate variable (often unknown!). Nephropores normally aligned. Diapause (?) [in response to drought] Tail normal. Body small to medium.

These worms live constantly in mineral rich organic soils from which foods, such as dead roots (rhysophilus) or rich organic matter integrated within the substrate (geophilus), are extracted. Their galleries are essentially horizontal except for the smaller geophilus types which follow dead roots. This is not a homogeneous category. The "epiendoges" live in the A horizon of the soil, while the geophile "hypoendoges" live below this horizon in poorly aerated compact soils. Because this group is functionally heterogeneous, interpretation of the morphological characters is difficult. Endogenous lifestyle may have induced characters 1, 4, 6, 8 and 9; the lack of pigmentation is a general feature of edaphic fauna living deep in the soil, e.g. springtails (Collembola). These are just three examples of attempts to ecologically classify earthworms. They have centred on the Lumbricidae because they were developed by Europeans (Julin, Bouché) based on earthworms in their areas (Sweden, France) or an American (Gates) in a state (Maine) which are areas only inhabited by this family of earthworms.

Camarero, yo dijo bien hechol! * re !

98

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99

EARTHWORM BIOGEOGRAPHY (BIOGEOGRAFIA DE LAS LOMBRICES DE TIERRA) Earthworms are no different than plants and other animals, in that each continent (except Antarctica) has its own distinct or indigenous species, plus others which have been introduced by humans deliberately or by accident. Many of the species and even some of the families of earthworms have been known from only a very few individuals and species. There are currently 18 families of terrestrial earthworms; of these only about a dozen exhibit any sort of widespread distribution (Reynolds, 1995a). In the map of the world (see opposite), the distribution of the families is illustrated. The numbers in parentheses in the list below (infra) following their name represent those families in the figure. The zoogeographical regions where earthworm families are believed to have originated are listed below, as well as other continents to which they have been transported (Reynolds, 1995a; Sims, 1980). Australian = Australia, Tasmania, New Guinea and some smaller islands of the Malayan Archipelago. Native: Exotic: Ethiopian =

Native: Exotic: Nearctic = Native: Exotic:

Acanthodrilidae (1), Megascolecidae (3), Octochaetidae (6) Lumbricidae (2), Megascolecidae (3) Africa south of the Sahara Desert and Atlas Mountains, and the southern corner of Arabia. Eudrilidae (4), Microchaetidae (5) Lumbricidae (2), Megascolecidae (3), Moniligasteridae (7) Canada, United States, Greenland, and northern Mexico. Acanthodrilidae (1), Komarekionidae (8), Lumbricidae (2), Lutodrilidae (9), Sparganophilidae (10) Eudrilidae (4), Glossoscolecidae (11), Lumbricidae (2), Megascolecidae (3), Octochaetidae (6), Ocnerodrilidae (12)

Neotropical= South America, Central America, most of Mexico, the West Indies and New Zealand. Native: Exotic:

100

Acanthodrilidae (1), Almidae (13), Glossoscolecidae (11), Ocnerodrilidae (12), Sparganophilidae (10) Eudrilidae (4), Lumbricidae (2), Megascolecidae (3)

J.W. Reynolds

Figure 1.

The origin and dispersal of the major terrestrial earthworm families. [From Reynolds, J.W. 1994. Earthworms of the world. Global Biodiversity 4(1): 11-16.]


101

Oriental = Native: Exotic: Palearctic = Native: Exotic:

India, Indochina, south China, Malaya including the westerly islands of the Malayan Archipelago. Megascolecidae (3), Moniligasteridae (7), Octochaetidae (6) Almidae (13), Eudrilidae (4), Lumbricidae (2) Europe to the Pacific Ocean, Africa north of the Sahara Desert, and Asia north of the Himalaya Mountains. Almidae (13), Diporochaetidae (14), Hormogastridae (15), Lumbricidae (2) Megascolecidae (3), Sparganophilidae (10)

An examination of the distribution of various earthworm families throughout North America is illustrated in a paper found in the end packet of this book (Reynolds, 1995a). This paper will form the basis of our discussion on this topic.

"Nosotros entendemos su preocupación, señora, pero esto no es suficiente para que nosotros continuemos. Ahora encuentre Usted la otra mitad de su esposo y entonces tendremos un caso."

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103

IDENTIFICATION - The Earthworms of Argentina (Lombrices de Tierra de la Argentina) This section was prepared by John W. Reynolds Oligochaetology Laboratory, Lindsay, Ontario, Canada

and Catalina C. de Mischis Facultad de Ciencias Exactas, Físicas y Naturales, Universidad Nacional de Córdoba, Argentina

The following 66 species are those reported so far from Argentina. The identification of some of the following species can be done quite easily, with the aid of several simple external taxonomic characters. The determination of others may require detailed internal dissection. Abbreviations 1st dp cl fp GM GT mf mp np

= first dorsal pore (1 poro dorsale) = clitellum (clitelo) = female pore (poro femenino) = Genital markings (Signos genitales, Señales de pubertad) = Genital tumescences (Tumescencias genitales) = genital male field (campo genital masculino) = male pore (poro masculino) = nephropore (nefridioporo)

pp pr ps sg sp TP

= prostatic pore (poro prostático) = prostomium (prostomio) = peristomium (peristomio) = seminal groove (surcos seminales) = spermathecal pores (poros espermatecales, Tecóporos) = Tubercula pubertatis (Tubérculos pubertales)

"Yo estoy seguro, Usted encontrará la disección de hoy como un desafío y además muy instructivo."

ACANTHODRILIDAE

104

J.W. Reynolds

Acanthodrilus doello-junadoi Cordero, 1942 1942

Acanthodrilus doello-junadoi Ann. Mus. Argent. 40: 272. Types, MACN 5782.

Length: (Largo) 61-93 mm Diameter: (Diámetro) 6-7 mm No. segments: (No. segmentos) 112-123 Colour: (Color) dorsum brown Prostomium: (Prostomio) epilobic, tongue open b-¾ First dorsal pore: (1o poro dorsale) lacking Clitellum: (Clitelo) annular, xiii-xvii (13-17) Tubercula pubertatis: (Tubérculos puberales) lacking Setae: (Quates\Sedas) closely paired, lumbricin aa = 4ab, ab = cd, bc = 3.5ab, dd = 8ab (=½C) Male pores: (Poros masculinos) xviii (17) near b Female pores: (Poros femeninos) xiv (14) Spermathecae: (Espermatecas) 2 pair in 8, 9 pores opening near B Seminal vesicles: (Vesículas seminales) Prostates: (Próstatas) 2 pairs in xvii and xix (17, 19) pores opening near B


105

Microscolex collislupi Michaelsen, 1910 1910

Microscolex collislupi Mitt. Mus. Hamb. 27(2): 63. MNHU 4781-2.

Length: (Largo) 32-45 mm Diameter: (Diámetro) 1.5-1.75 mm No. segments: (No. segmentos) 76-88 Colour: (Color) unpigmented Prostomium: (Prostomio) epilobic, tongue ¾ open to tanylobic First dorsal pore: (1o poro dorsale) lacking Clitellum: (Clitelo) annular, xiii¾-½xvii (13¾-½17) Tubercula pubertatis: (Tubérculos puberales) lacking Setae: (Quates\Sedas) widely paired, lumbricin aa:ab:bc:cd:dd = 3:2:3:2:4 Male pores: (Poros masculinos) xviii (18) near b Female pores: (Poros femeninos) xiv (14) near a Spermathecae: (Espermatecas) 2 pair in viii and ix (8, 9) with pores opening near a Seminal vesicles: (Vesículas seminales) Prostates: (Próstatas) 1 pair in xvii (17) pores opening near a

106

J.W. Reynolds

Microscolex dubius (Fletcher, 1887) 1887

Eudrilus dubius Proc. Linn. Soc. N.S. Wales 2: 378. Typus amissus.

Length: (Largo) 30-102 mm Diameter: (Diámetro) 2-5 mm No. segments: (No. segmentos) 110-120 Colour: (Color) pigment lacking Prostomium: (Prostomio) epilobic, tongue open First dorsal pore: (1o poro dorsale) lacking Clitellum: (Clitelo) annular, xiii-xvi, xvii (13-16, 17) Tubercula pubertatis: (Tubérculos puberales) lacking Setae: (Quates\Sedas) widely paired, lumbricin dd > aa > bc > cd > ab Male pores: (Poros masculinos) xvii (17) Female pores: (Poros femeninos) xiv (14) Spermathecae: (Espermatecas) lacking Seminal vesicles: (Vesículas seminales) in xi, xii (11, 12) Prostates: (Próstatas) 1 pair tubular in xvii (17)

ventral view - genital field [From Righi, 1979]


[From Jamieson, 1974]

107

Microscolex michaelseni Beddard, 1895 1895

Microscolex michaelseni Proc. Zool. Soc. Lond., 1985, p. 231. MNHU 2858, 7418.

Length: (Largo) 85 mm Diameter: (Diámetro) 3 mm No. segments: (No. segmentos) 92 Colour: (Color) unpigmented Prostomium: (Prostomio) epilobic First dorsal pore: (1o poro dorsale) Clitellum: (Clitelo) annular, xiii-xvi (13-16) Tubercula pubertatis: (Tubérculos puberales) lacking Setae: (Quates\Sedas) widely paired, lumbricin Male pores:(Poros masculinos) xvii (17) Female pores: (Poros femeninos) xiv (14) Spermathecae: (Espermatecas) Seminal vesicles: (Vesículas seminales) 1 pair in xi (11) Prostates: (Próstatas) 2 pair in xvii and xviii (17, 18)

108

J.W. Reynolds

Microscolex phosphoreus (Dugès, 1837) 1837

Lumbricus phosphoreus Ann. Sci. Nat. (2), 8: 17. Typus amissus.

Length: (Largo) 10-55 mm Diameter: (Diámetro) 1-2 mm No. segments: (No. segmentos) 73-90 Colour: (Color) pigment lacking, phosphorescent when alive Prostomium: (Prostomio) epilobic, tongue open First dorsal pore: (1o poro dorsale) lacking Clitellum: (Clitelo) annular, xiii, xiv-xvi, xvii (13, 14-16, 17) Tubercula pubertatis: (Tubérculos puberales) lacking Setae: (Quates\Sedas) widely paired, lumbricin aa = bc; dd > aa > cd > ab Male pores: (Poros masculinos) xvii (17) Female pores: (Poros femeninos) xiv (14) Spermathecae: (Espermatecas) 1 pair in viii (8) opening at 8/9 lateral to A Seminal vesicles: (Vesículas seminales) in xi and xii (11, 12) Prostates: (Próstatas) 1 pair tubular in xvii (17)

[From Bouché, 1972]


109

Notiodrilus bovei (Rosa, 1889) 1889

Mandane bovei Ann. Mus. Genova (2), 27: 143. Types, MGDG 44036!

Length: (Largo) 35-90 mm Diameter: (Diámetro) 3 mm No. segments: (No. segmentos) ± 100 Colour: (Color) brown Prostomium: (Prostomio) epilobic First dorsal pore: (1o poro dorsale) lacking Clitellum: (Clitelo) annular, xiii-xvi (13-16) Tubercula pubertatis: (Tubérculos puberales) lacking Setae: (Quates\Sedas) Male pores: (Poros masculinos) xvii (17) Female pores: (Poros femeninos) xiv (14) Spermathecae: (Espermatecas) 2 pair in viii and ix (8, 9) Seminal vesicles: (Vesículas seminales) in xi and xii (11, 12) Prostates: (Próstatas)

110

J.W. Reynolds

Notiodrilus silvestrii Rosa, 1901 1901

Notiodrilus silvestrii Atti Soc. Ligustica (4), 4(35): 7. Types unknown.

Length: (Largo) mm Diameter: (Diámetro) mm No. segments: (No. segmentos) Colour: (Color) Prostomium: (Prostomio) First dorsal pore: (1o poro dorsale) lacking Clitellum: (Clitelo) annular, Tubercula pubertatis: (Tubérculos puberales) lacking Setae: (Quates\Sedas) Male pores: (Poros masculinos) xvii (17) Female pores: (Poros femeninos) xiv (14) Spermathecae: (Espermatecas) Seminal vesicles: (Vesículas seminales) Prostates: (Próstatas)


111

GLOSSOSCOLECIDAE Anteoides rosae Cognetti, 1902 1902

Anteoides rosae Boll. Mus. Torino 17(420): 4. Types, MZUT 446!

Length: (Largo) 45-70 mm Diameter: (Diámetro) 2-3 mm No. segments: (No. segmentos) 132-187 Colour: (Color) whitish Prostomium: (Prostomio) prolobic First dorsal pore: (1o poro dorsale) lacking Clitellum: (Clitelo) annular, xiv-xxi (14-21) Tubercula pubertatis: (Tubérculos puberales) xvii½-xviii, ½xix (17½-18, ½19) Setae: (Quates\Sedas) closely paired, lumbricin aa:ab:bc:cd:dd = 25:5:42:2:122 Male pores: (Poros masculinos) xviii (18) Female pores: (Poros femeninos) xiv (14) near b Spermathecae: (Espermatecas) Seminal vesicles: (Vesículas seminales) Prostates: (Próstatas)

[From Cognetti, 1902]

112

J.W. Reynolds

Diaguita michaelseni Cordero, 1942 1942

Diaguita michaelseni Ann. Mus. Argent. 40: 286. Types unknown.

Length: (Largo) 50-60 mm Diameter: (Diámetro) mm No. segments: (No. segmentos) 80-100 Colour: (Color) Prostomium: (Prostomio) prolobic First dorsal pore: (1o poro dorsale) Clitellum: (Clitelo) annular, xv-xxi (15-21) Tubercula pubertatis: (Tubérculos puberales) Setae: (Quates\Sedas) widely paired, lumbricin aa = 3ab, dd = ½C Male pores: (Poros masculinos) xviii (18) Female pores: (Poros femeninos) xiv (14) Spermathecae: (Espermatecas) 2 pair in viii and ix (8, 9) opening in 7/8 and 8/9 near b Seminal vesicles: (Vesículas seminales) Prostates: (Próstatas)


113

Enantiodrilus borelli Cognetti, 1902 1902

Enantiodrilus borelli Boll. Mus. Torino 17(420): 9. Types, MZUT 277-9!

Length: (Largo) 60-150 mm Diameter: (Diámetro) 5 mm No. segments: (No. segmentos) 200-250 Colour: (Color) brownish Prostomium: (Prostomio) "breve, retracttile, disfinto" First dorsal pore: (1o poro dorsale) lacking "assenti" Clitellum: (Clitelo) annular, xiv-xxii (14-22) Tubercula pubertatis: (Tubérculos puberales) lacking Setae: (Quates\Sedas) aa = 2bc, dd = 2aa, dd >½C aa:ab:bc:cd:dd = 30:1:10:1:60 Male pores: (Poros masculinos) xix (19) near b Female pores: (Poros femeninos) xiii or xiv (13 or 14) near a Spermathecae: (Espermatecas) 2 pair in viii and ix (8 and 9) Seminal vesicles: (Vesículas seminales) lacking "assenti" Prostates: (Próstatas) ?

[From Cognetti, 1902]

114

J.W. Reynolds

Glossoscolex bergi (Rosa, 1900) 1900

Geoscolex bergi Comun. Zool. Mus. Hist. Nat. 1: 209. Types, MACN 4797.

Length: (Largo) 290-305 mm Diameter: (Diámetro) 10-12 mm No. segments: (No. segmentos) 310-380 Colour: (Color) unpigmented (?) Prostomium: (Prostomio) epilobic (?) First dorsal pore: (1o poro dorsale) 13/14 Clitellum: (Clitelo) annular, xv-xxiv (15-24) Tubercula pubertatis: (Tubérculos puberales) lacking Setae: (Quates\Sedas) closely paired, lumbricin, ab = cd, bc = ± 3ab, dd > ½C Male pores: (Poros masculinos) xx/xxi (20/21) near a Female pores: (Poros femeninos) xiv (14) near b Spermathecae: (Espermatecas) Seminal vesicles: (Vesículas seminales) 1 pair in xii (12) Prostates: (Próstatas)


115

Glossoscolex bonariensis Cordero, 1942 1942

Glossoscolex bonariensis Ann. Mus. Argent. 40: 284. Types, MACN 19390.

Length: (Largo) 57-65 mm Diameter: (Diámetro) mm No. segments: (No. segmentos) 130-145 Colour: (Color) unpigmented Prostomium: (Prostomio) prolobic First dorsal pore: (1o poro dorsale) lacking Clitellum: (Clitelo) saddle, xv-xxv (15-25) Tubercula pubertatis: (Tubérculos puberales) lacking Setae: (Quates\Sedas) closely paired, lumbricin, aa:ab:bc:cd:dd = 40:1:9.7:1:51.4 Male pores: (Poros masculinos) xvii (17) Female pores: (Poros femeninos) xiv (14) Spermathecae: (Espermatecas) lacking (?) Seminal vesicles: (Vesículas seminales) 1 pair in xii (12) Prostates: (Próstatas)

[From Righi, 1978]

116

J.W. Reynolds

Glossoscolex corrientus Righi, 1984 1984

Glossoscolex corrientus Stud. Neotrop. Fauna Environ. 19(2): 116. Types, DZSP 537-8.

Length: (Largo) 100 mm Diameter: (Diámetro) 3-3.7 mm No. segments: (No. segmentos) Colour: (Color) unpigmented Prostomium: (Prostomio) First dorsal pore: (1o poro dorsale) lacking Clitellum: (Clitelo) annular, ½xiii or xiii-xxvi (½13 or 13-26) Tubercula pubertatis: (Tubérculos puberales) lacking Setae: (Quates\Sedas) closely paired, lumbricin aa:ab:bc:cd:dd = 43:1:5.5:1:39 Male pores: (Poros masculinos) xxvii (17) Female pores: (Poros femeninos) xiv (14) Spermathecae: (Espermatecas) lacking Seminal vesicles: (Vesículas seminales) 1 pair in xii (12) Prostates: (Próstatas)

[From Righi, 1984]


117

Glossoscolex forguesi (Perrier, 1881) 1881

Titanus forguesi Arch. Zool. Exp. Gén. 9: 217. Types, unknown.

Length: (Largo) 100 mm Diameter: (Diámetro) mm No. segments: (No. segmentos) Colour: (Color) unpigmented Prostomium: (Prostomio) First dorsal pore: (1o poro dorsale) lacking Clitellum: (Clitelo) saddle, xvi-xxii (16-22) Tubercula pubertatis: (Tubérculos puberales) Setae: (Quates\Sedas) lumbricin Male pores: (Poros masculinos) xvii (17) Female pores: (Poros femeninos) xiv (14) Spermathecae: (Espermatecas) lacking Seminal vesicles: (Vesículas seminales) Prostates: (Próstatas)

118

J.W. Reynolds

Glossoscolex uruguayensis uruguayensis Cordero, 1943 1943

Glossoscolex uruguayensis Comun. Zool. Mus. Hist. Nat. Montevideo 1(2): 4. Types, unknown.

Length: (Largo) 111-315 mm Diameter: (Diámetro) 4-8 mm No. segments: (No. segmentos) 230-354 Colour: (Color) unpigmented Prostomium: (Prostomio) epilobic First dorsal pore: (1o poro dorsale) lacking (?) Clitellum: (Clitelo) saddle, xv-xxii (15-22) Tubercula pubertatis: (Tubérculos puberales) lacking Setae: (Quates\Sedas) widely paired, lumbricin aa = 4bc, dd = 5bc, aa < ½C, dd >aC Male pores: (Poros masculinos) xvii (17) Female pores: (Poros femeninos) xiv (14) Spermathecae: (Espermatecas) lacking Seminal vesicles: (Vesículas seminales) in xii-xiii, xiv (12-13, 14) Prostates: (Próstatas) ?


119

Glossoscolex uruguayensis ljungstromi Righi, 1978 1978

Glossoscolex uruguayensis ljungstromi Fave 1(3): 172. Types, DZSP 402-3.

Length: (Largo) mm Diameter: (Diámetro) mm No. segments: (No. segmentos) Colour: (Color) unpigmented Prostomium: (Prostomio) epilobic First dorsal pore: (1o poro dorsale) lacking (?) Clitellum: (Clitelo) saddle, xi-xxii (11-22) Tubercula pubertatis: (Tubérculos puberales) xvi-xviii (16-18) Setae: (Quates\Sedas) closely paired, lumbricin aa:ab:bc:cd:dd = 16.4:1:5.4:1:12 Male pores: (Poros masculinos) xvii (17) Female pores: (Poros femeninos) xiv (14) Spermathecae: (Espermatecas) lacking Seminal vesicles: (Vesículas seminales) lacking Prostates: (Próstatas)

[From Righi, 1978]

120

J.W. Reynolds

Opistodrilus borelli Rosa, 1895 1895

Opistodrilus borelli Boll. Mus. Torino 10(204): 3. Types, unknown.

Length: (Largo) 70-75 mm Diameter: (Diámetro) 2.5-4.5 mm No. segments: (No. segmentos) 159-230 (x 181) Colour: (Color) dark grey dorsally, light grey-whitish ventrally Prostomium: (Prostomio) First dorsal pore: (1o poro dorsale) lacking Clitellum: (Clitelo) saddle, ½xiii, xiv, xv-xxiv (½13, 14, 15-24) Tubercula pubertatis: (Tubérculos puberales) present far behind clitellum xxxii-xxxvi (32-36) Setae: (Quates\Sedas) closely paired, lumbricin aa:ab:bc:cd:dd = 6.1:1:7.8:1:23.3 Male pores: (Poros masculinos) xxxiv (34) Female pores: (Poros femeninos) xiv (14) Spermathecae: (Espermatecas) 3 pair 7, 8 and 9 Seminal vesicles: (Vesículas seminales) Prostates: (Próstatas)


121

[From Righi, 1984] Pontoscolex corethrurus (Müller, 1847) 1856

Lumbricus corethrurus Abh. Naturg. Ges. Halle 4: 26. Typus amissus.

Length: (Largo) 60-120 mm Diameter: (Diámetro) 4-6 mm No. segments: (No. segmentos) 167-222 Colour: (Color) unpigmented Prostomium: (Prostomio) lacking First dorsal pore: (1o poro dorsale) lacking Clitellum: (Clitelo) saddle, xv, xvi-xxii, xxiii (15, 16-22, 23) Tubercula pubertatis: (Tubérculos puberales) xix-xxi, xxii (19-21, 22) Setae: (Quates\Sedas) closely paired, "quincunx" arrangement towards posterior end Male pores: (Poros masculinos) 20/21 lateral to B Female pore: (Poro femenino) xiv on left side Spermathecae: (Espermatecas) 3 pairs opening 6/7-8/9 Seminal vesicles: (Vesículas seminales) 1 pair extending through viii-x Prostates: (Próstatas)

122

J.W. Reynolds

Rhinodrilus parvus (Rosa, 1895) 1895

Anteus parvus Boll. Mus. Torino 10(204): 2.. Types unknown.

Length: (Largo) mm Diameter: (Diámetro) mm No. segments: (No. segmentos) Colour: (Color) Prostomium: (Prostomio) First dorsal pore: (1o poro dorsale) Clitellum: (Clitelo) Tubercula pubertatis: (Tubérculos puberales) Setae: (Quates\Sedas) Male pores: (Poros masculinos) Female pores: (Poros femeninos) Spermathecae: (Espermatecas) Seminal vesicles: (Vesículas seminales) Prostates: (Próstatas)


123

LUMBRICIDAE Allolobophora georgii Michaelsen, 1890 1890

Allolobophora georgii Mitt. Mus. Hamb. 7: 3. Types, ZMUH 13000!

Length: (Largo) 38-80 mm Diameter: (Diámetro) mm No. segments: (No. segmentos) 98-115 Colour: (Color) unpigmented Prostomium: (Prostomio) epilobic First dorsal pore: (1o poro dorsale) 4/5 Clitellum: (Clitelo) saddle, xxviii, xxix-xxxv, ½xxxvi (28, 29-35, ½36) Tubercula pubertatis: (Tubérculos puberales) 2 sucker-like discs on xxxi and xxxiii (31 & 33) Setae: (Quates\Sedas) aa > bc; dd = ½C Male pores: (Poros masculinos) xv (15) Female pores: (Poros femeninos) xiv (14) Spermathecae: (Espermatecas) 2 pairs, in ix and x or x and xi (9 and 10 or 10 and 11) Seminal vesicles: (Vesículas seminales) 4 pairs in 9-12 Prostates: (Próstatas) lacking

[From Bouché, 1972]

124

J.W. Reynolds

Aporrectodea rosea (Savigny, 1826) pink soil worm 1826

Enterion roseum Mém. Acad. Sci. Inst. Fr. 5: 182. Types, unknown.

Length: (Largo) 25-85 mm Diameter: (Diámetro) 3-5 mm No. segments: (No. segmentos) 120-150 Colour: (Color) unpigmented, but colour appears rosy or greyish when alive, and white when preserved Prostomium: (Prostomio) epilobic First dorsal pore: (1o poro dorsale) 4/5 Clitellum: (Clitelo) xxv, xxvi-xxxii (25, 26-32), somewhat flared ventrally Tubercula pubertatis: (Tubérculos puberales) xxix-xxxi (29-31) Setae: (Quates\Sedas) closely paired, lumbricin, AA > BC < DD; AB > CD; anteriorly DD = ½C; posteriorly DD = aC Male pores: (Poros masculinos) xv (15) Female pores: (Poros femeninos) xiv (14) Spermathecae: (Espermatecas) 2 pairs in ix and xi (9, 11) Seminal vesicles: (Vesículas seminales) 4 pairs, 9-12 Prostates: (Próstatas) lacking

Figure 2.

External longitudinal views of Aporrectodea rosea showing taxonomic characters. A. Lateral view. B. Ventral view [From Reynolds, 1977]


125

Aporrectodea trapezoides (Dugès, 1828) 1828

Lumbricus trapezoides Ann. Sci. Nat. 15(1): 289. Typus amissus.

Length: (Largo) 80-140 mm Diameter: (Diámetro) 3-7 mm No. segments: (No. segmentos) 93-169, generally > 130 Colour: (Color) variable, lighter behind clitellum then deeper slate, brown, brownish, reddish brown, occasionally reddish but not purple Prostomium: (Prostomio) epilobic First dorsal pore: (1o poro dorsale) 12/13 Clitellum: (Clitelo) saddle, xxvii, xxviii-xxxiii, xxxiv (27, 28-33, 34) Tubercula pubertatis: (Tubérculos puberales) xxxi-xxxiii (31-33) Genital Tumescences: (Tumescences genitales) a and b only in ix-xi, xxxii-xxxiv, often in xxviii, ocassionally xxvi-xxix (9-11, 32-34, often 28, occasionally 26-29) Setae: (Quates\Sedas) closely paried, lumbricin, AA > AB, DD < ½C Male pores: (Poros masculinos) xv (15) Female pores: (Poros femeninos) xiv (14) Spermathecae: (Espermatecas) 2 pairs in ix and xi (9, 11) Seminal vesicles: (Vesículas seminales) 4 pairs in 9-12 Prostates: (Próstatas) lacking

Figure 4.

126

External longitudinal views of Aporrectodea trapezoides showing taxonomic characters. A. Lateral view. B. Ventral view [From Reynolds, 1977]

J.W. Reynolds

Aporrectodea turgida (Eisen, 1873) pasture worm 1873

Allolobophora turgida Öfv. Vet.-Akad. Förh. Stockholm 30(8): 46. Types, USNM 1157!

Length: (Largo) 60-85 mm Diameter: (Diámetro) 3.5-5 mm No. segments: (No. segmentos) 130-168 Colour: (Color) unpigmented, with the region anterior to the crop flesh pink and the remaining segments pale grey, or occasionally with light pigmentation on the dorsal surface Prostomium: (Prostomio) epilobic First dorsal pore: (1 poro dorsale) 12/13 or 13/14 Clitellum: (Clitelo) xxvii, xxviii, xxix-xxxiv, xxxv (27, 28, 29-34, 35) Tubercula pubertatis: (Tubérculos puberales) xxix-xxxiii (29-33) Genital tumescences: (Tumescencias genitales) a and b only in xxx, xxxii-xxxiv (30, 32-34) and frequently in xxvii (27) Setae: (Quates/Sedas) closely paired, lumbricin aa:ab:bc:cd:dd = 3:1:2:.6:10 Male pores: (Poros masculinos) xv (15) Female pores: (Poros femeninos) xiv (14) Spermathecae: (Espermatecas) 2 pairs in ix and xi (9, 11) Seminal vesicles: (Vesículas seminales) 4 pairs in 9-12 Prostates: (Próstatas) lacking

Figure 14.

External longitudinal views of Aporrectodea turgida showing taxonomic characters. A. Lateral view. B. Ventral view. [From Reynolds, 1977]


127

Bimastos beddardi sophiae Mercadal de Barrio et Barrio, 1988 1988

Bimastos beddardi sophiae Physis Secc. C Cont. Org. Terr. 46(110): 2. Types, MACN.

Length: (Largo) mm Diameter: (Diámetro) mm No. segments: (No. segmentos) 135-150 Colour: (Color) Prostomium: (Prostomio) epilobic First dorsal pore: (1o poro dorsale) 6/7 Clitellum: (Clitelo) saddle, xxiv-xxxii (24-32) Tubercula pubertatis: (Tubérculos puberales) xxix-xxxi (29-31) Setae: (Quates\Sedas) closely paried, lumbricin aa > bc; ab > cd Male pores: (Poros masculinos) xv Female pores: (Poros femeninos) xiv Spermathecae: (Espermatecas) Seminal vesicles: (Vesículas seminales) Prostates: (Próstatas)

128

J.W. Reynolds

Bimastos parvus (Eisen, 1874) American bark worm 1874

Allolobophora parva Öfv. Vet.-Akad. Förh. Stockholm 31(2): 46. Types, unknown.

Length: (Largo) 17-65 mm Diameter: (Diámetro) 1.5-3.0 mm No. segments: (No. segmentos) 65-97 Colour: (Color) reddish dorsally Prostomium: (Prostomio) epilobic First dorsal pore: (1o poro dorsale) 5/6 Clitellum: (Clitelo) saddle, xxiii, xxiv-xxxi, xxxii (23, 24-31, 32) Tubercula pubertatis: (Tubérculos puberales) lacking Setae: (Quates\Sedas) closely paired, lumbricin, CD = ¾AB; AA slightly > BC, DD = ½C Male pores: (Poros masculinos) xv (15) Female pores: (Poros femeninos) xiv (14) Spermathecae: (Espermatecas) lacking Seminal vesicles: (Vesículas seminales) 2 pair in xi and xii (11, 12) Prostates: (Próstatas) lacking

Figure 16.

External longitudinal views of Bimastos parvus showing taxonomic characters. A. Lateral view. B. Ventrolateral view. [From Reynolds, 1977]


129

Dendrodrilus rubidus (Savigny, 1826) European bark worm 1826

Enterion rubidum Mém. Acad. Sci. Fr. 5: 182. Types, unknown.

Length: (Largo) 20-90 mm Diameter: (Diámetro) 2-5 mm No. segments: (No. segmentos) 50-120 Colour: (Color) red and darker dorsally Prostomium: (Prostomio) epilobic First dorsal pore: (1o poro dorsale) 5/6 Clitellum: (Clitelo) xxvi, xxvii-xxxi, xxxii (26, 27-31, 32) Tubercula pubertatis: (Tubérculos puberales) xxviii, xxix-xxx (28, 29-30); some morphs lacking Setae: (Quates\Sedas) widely paired, lumbricin, AB < CD, BC = 2CD Male pores: (Poros masculinos) xv (15) Female pores: (Poros femeninos) xiv (14) Spermathecae: (Espermatecas) 2 pairs in ix and xi (9, 11) Seminal vesicles: (Vesículas seminales) 3 pairs in 9, 11 and 12 Prostates: (Próstatas) lacking

Figure 6.

130

External longitudinal views of Dendrodrilus rubidus showing taxonomic characters. A. Lateral view. B. Ventral view. [From Reynolds, 1977]

J.W. Reynolds

Eisenia foetida (Savigny, 1826) manure worm, red wiggler, etc. 1826

Enterion fetidum (corr. foetidum) Mém. Acad. Sci. Inst. Fr. 5: 182. Types, MNHN.

Length: (Largo) 35-130 mm (generally < 70 mm) Diameter: (Diámetro) 3-5 mm No. segments: (No. segmentos) 80-110 Colour: (Color) variable, purple, red, dark red, banded with yellow intersegmental furrows Prostomium: (Prostomio) epilobic First dorsal pore: (1 poro dorsale) 4/5 (sometimes 5/6) Clitellum: (Clitelo) xxiv, xxv, xxvi-xxxii (24, 25, 26-32) Tubercula pubertatis: (Tubérculos puberales) xxviii-xxx (28-30) Setae: (Quates/Sedas) closely paired, lumbricin, AB = CD; BC < AA; anteriorly DD = ½C, posteriorly DD ab = bc > cd, dd = 2 aa Male pores: (Poros masculinos) xv (15) Female pores: (Poros femeninos) xiv (14) Spermathecae: (Espermatecas) 5 pairs Seminal vesicles: (Vesículas seminales) 4 pairs in 9-12 Prostates: (Próstatas) lacking

, 1991]


133

Octolasion complanatum (Dugès, 1828) 1828

Lumbricus complanatus Ann. Sci. Nat. 15: 289. Typus amissus.

Length: (Largo) 80-270 mm Diameter: (Diámetro) mm No. segments: (No. segmentos) 150-251 Colour: (Color) dark grey Prostomium: (Prostomio) epilobic First dorsal pore: (1o poro dorsale) 11/12-13/14 Clitellum: (Clitelo) saddle, ½xxvii, xxviii, xxix, xxx-xxxvii, xxxviii (½27, 28, 29, 30-37, 38) Tubercula pubertatis: (Tubérculos puberales) xxviii, xxix, xxx-xxxviii, xxxix, xl, xli (28, 29, 30-38, 39, 40, 41) Setae: (Quates\Sedas) widely paired, lumbricin, aa>ab>bc