Louisiana State University LSU Digital Commons
LSU Historical Dissertations and Theses Graduate School
1969 The aE rthworm Genus Pheretima, Kinberg, 1866, in Louisiana (Oligochaeta: Megascolecidae). Richard Elvin Tandy Louisiana State University and Agricultural & Mechanical College
Follow this and additional works at: https://digitalcommons.lsu.edu/gradschool_disstheses
Recommended Citation Tandy, Richard Elvin, "The Earthworm Genus Pheretima, Kinberg, 1866, in Louisiana (Oligochaeta: Megascolecidae)." (1969). LSU Historical Dissertations and Theses. 1566. https://digitalcommons.lsu.edu/gradschool_disstheses/1566
This Dissertation is brought to you for free and open access by the Graduate School at LSU Digital Commons. It has been accepted for inclusion in LSU Historical Dissertations and Theses by an authorized administrator of LSU Digital Commons. For more information, please contact [email protected]. This dissertation has been microfilmed exactly as received 69-17,130
TANDY, Richard Elvin, 1937- THE EARTHWORM GENUS PHERETIMA KINBERG, 1866, IN LOUISIANA (OLIGOCHAETA: MEGASCOLECIDAE).
Louisiana State University and Agricultural and Mechanical College, Ph.D., 1969 Zoology University Microfilms, Inc., Ann Arbor, Michigan
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. The Earthworm Genus Pheretima Kinberg, 1856, In Louisiana
(01igochaeta:M egascolecidae)
A D issertation
Submitted to the Graduate Faculty of the Louisiana State University and Agricultural and Mechanical College in partial fulfillm ent of the requirements for the degree of Doctor of Philosophy
in
The Department of Zoology and Physiology
by Richard Elvin Tandy B.A., Anderson College, 1960 M.S., Louisiana Polytechnic Institute, 1963 January, 1969
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. ACKNOWLEDGMENT
Sincere appreciation is expressed to Dr. Walter J. Harman for
suggesting that this study be made and for his continuous counsel and
guidance. Special thanlcs are expressed to Dr. Gordon E. Gates, Bangor,
Maine, for the use of an unpublished manuscript on earthworms of
Burma and for the confirmation of identifications; and to Dr. Clair A.
Brown, Department of Botany, Louisiana State University, for his
comments on imported plant species in Louisiana. The author wishes to
express his gratitude to all of. the persons who assisted in any way
during this study, especially those who brought the author collections
of earthworms.
l i
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. TABLE OF CONTENTS
Page
T i t l e P a g e ...... 1
Acknowledgments ...... 11
T a b le o f C o n t e n t s ...... I l l
List of Tables ...... Iv
List of Maps...... V
List of Illustrations...... v i
A b s t r a c t ...... v l l l
I. Introduction...... 1
II. Materials and Methods ...... 3
III. Taxonomy and Conventions ...... 6
A. Generic Account...... 11
B. Species Accounts...... 16.
Pheretima agrestls ...... 16 Pheretima callfornlca ...... 25 Pheretima dlffrlngens ...... 32 Pheretima hawayana...... 58 Pheretima hllgendorf1 ...... 69 Pheretima hupelensls...... 76 Pheretima minima...... 81 Pheretima m o rrlsl ...... 86
C. Key to the Louisiana Species of Pheretim a...... 93
IV. Ecology ...... - 94
V. Summary ...... 98
Literature Cited...... 101-
A p p en d ix ...... 154
V i t a ...... 155
1 1 1
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. LIST OF TABLES
T able Number Page
1. Number of Sites at which Species of Pheretima Occurred Singly and Together in Louisiana...... 108
2. Total Number of Associations of Pheretima Species at 448 S ite s...... 109
3. Total Number of Preclitellar Genital Markings in Pheretima diffringens ...... 110
4. Location of Preclitellar Genital Markings in Pheretima diffringens ...... I l l
5. Frequency of Preclitellar Genital Markings in Pheretima diffringens ...... 112
6. Total Number of Preclitellar Genital Markings in Pheretima hawayana...... 113
7. Location of Preclitellar Genital Markings in Pheretima hawayana ...... 114
8. Total Number of Postclitellar Genital Markings in Pheretima hawayana...... r . 115
9. Location of Postclitellar Genital Markings in Pheretima hawayana ...... 116
10. Location of Midventral Genital Markings in Pheretima m orrisi ...... 117
11. Location of Lateral Preclitellar Genital Markings in Pheretima m orrisi ...... 118
12. Location of Postclitellar Genital Markings in Pheretima m orrisi ...... 119
13. Climatological Data for New Orleans, Louisiana .... 120
14. Climatological Data for Shreveport, Louisiana...... 121
IV
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. LIST OF MAPS
Map Number Page
1 . Distribution of Pheretima agrestis ...... 122
2 . Distribution of Pheretima Californiea ...... 123
3. Distribution of Pheretima diffringens ...... 124
4 . Distribution o f Pheretima hawayana...... 125
5. Distribution of Pheretima hilgendorfi ...... 126
6. Distribution of Pheretima hupeiensis ...... 127
7 . Distribution o f Pheretima minima...... 128
8 . Distribution of Pheretima m orrisi...... 129
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. LIST OF ILLUSTRATIONS
F ig u re Number Page
1. Spermatheca of Pheretima agrestis...... 131
2. Longitudinal Section of the Male Porophore o f P h e re tim a a g r e s t i s ...... 131
3. Modified Epidermis of Pheretima agrestis ...... 133
4. Male Pore Region of Pheretima ag restis ...... 133
5. Manicate Intestinal Caecum of Pheretima agrestis . . . 135
6. Internal Male Terminalia of Pheretima agrestis .... 135
7. Retracted and Partially Everted Male Porophore of Pheretima californica ...... 137
8. Everted Male Porophores of Pheretima californica . . . 137
9. Simple Intestinal Caecum of Pheretima californica. . . 137
10. Hypothetical Total Battery of Genital Markings in P h e re tim a d i f f r i n g e n s ...... 139
11. Genital Markings of Pheretima diffringens ...... 139
12. Genital Markings of Pheretima diffringens ...... 139
13. Histology of Genital Marking Gland of Pheretima diffringens ...... 141
1 4 . Preclitellar Genital Markings of Pheretima hawayana ...... 143
15. Male Pore Region of Pheretima hawayana ...... 143
16. Longitudinal Section of the Male Porophore of Phererima hilgendorfi ...... 145
17. Male Pore Region of Pheretima hilgendorfi ...... 145
VI
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. LIST OF ILLUSTRATIONS (Continued)
F ig u re Number Page
1 8 . Internal Male Terminalia o f Pheretima hilgendorfi. . . 147
1 9 . Male Pore Region of Pheretima hupeiensis...... 149
2 0 . Spermatheca o f Pheretima minima...... 151
2 1 . Internal Male Terminalia o f Pheretima m inim a...... 151
22. Preclitellar Genital Markings of Pheretima morrisi . . 153
2 3 . Male Pore Region of Pheretima m orrisi...... 153
V X i
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. ABSTRACT
The present study, which is the first thorough survey of any taxon
of Oligochaeta in Louisiana, reports the distribution of 8 species of
the introduced earthworm genus Pheretima in the state. A total of 3756
specimens was collected over a three and one-half year period at 448
sites in all 64 parishes of Louisiana. All of the species of Pheretima
previously reported from the state were collected in this study and
were found more widely distributed than the scattered reports in the
literature would indicate. In addition, Pheretima minima is reported
for the first time in the continental United States, from Louisiana.
Four species (P. californica, P. diffringens, _P. hawayana, JP.
m orrisi) are widely distributed in the state having dispersed from
areas of introduction. The remaining 4 species (P. agrestis, P.
hilgendorfi, P. hupeiensis, P. minima) are largely limited to areas
where they were introduced.
A diagnosis of the genus and each of the species is included
together with information regarding genital markings, autotomy,
regeneration, and life history. A key to the species is also provided.
The male pores of P. agrestis and P. hilgendorfi are described from
the first United States specimens which possess male term inalia. These
two species are found to have an annual life cycle in Louisiana. The
histology of the genital marking glands of P. diffringens is described.
The types of habitats in which each of the species occurs are
discussed. Each species is found in a variety of ecological habitats
and soil types,
v i i i
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. I . INTRODUCTION
The A siatic earthworm genus Pheretima was declared by Stephenson
(1930, p. 838) to be " . . . by far the largest genus of Oligochaeta,"
and was estimated to contain some 293 species. Michaelsen (1934)
listed the area in which species were known to be endemic as " . . .
the whole Malay Archipelago and the south-eastern part of Asia with a
part of Burma, China, Korea, and Japan, and embraces to the south-east
the Solomon Islands, the New Hebrides, and the Loyalty Islands." A
number of species of Pheretima have been carried, presumably by man, to
other parts of the world, including the Neotropical and Neartic regions.
Michaelsen (1903) coined the temn "peregrine" to designate those species
of earthworms, which by means of their own, or through transportation
by man, are widespread in their distribution. In that work Michaelsen
listed 22 of 119 species of Pheretima as peregrine.
The first North American report of Pheretima was made by Kinberg
in 1866. That publication was however, buried in the literature until
Michaelsen (1899a) reexamined Kinberg's collection and found P.
californica and P. diffringens in the collections from California.
The first Louisiana records of Pheretima are those of Gates
(1937c) based on specimens sent to museums for deposit. In that paper
Gates lamented that it is "... unfortunate that earthworms have
never been systematically collected in the warmer portions of the
continent in which Pheretima is likely to be found, for we do not yet
know how many of the imported species have become definitely
established within our region, how widely they have been able to
1
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. spread, what ecological niches they have been able to occupy and what
their role may be in supplementing or supplanting the native species."
Since 1937 only scattered records of the Pheretima of Louisiana
have appeared in the literature, including the report of Harman (1952),
which briefly listed several Pheretima species from Lincoln Parish in
the northern part of the state.
This study was undertaken to determine how many species of
Pheretima occur in Louisiana, how widely they ate distributed, something
of their biology in the United States, and what ecological habitats
they occupy.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. I I . MATERIALS AND METHODS
A total of 478 collections was made in Louisiana in the period
from November, 1963 to June, 1967. Nine collections that had been
secured by Harman before 1960 were also used. The following list shows
the number of collections secured in each month of the year; January,
23; February, 22; March, 72; April, 106; May, 21; June, 10; July, 23;
August, 73; September, 33; October, 27; November, 26; and December,
5 1 .
At least five collections were made in each of the 64 parishes of
the state, and all types of available habitats were examined in each of
the parishes. Attempts were made to secure specimens in remote areas
far from the activities of man, as well as in those areas where
unreported species might have been introduced into the state, such as
botanical gardens, nurseries, yards and plantation sites.
At each collecting site all available habitat areas were examined.
For instance, on the floodplain of a bayou, the woods or swamp on
either side of the bayou was also examined. At first, worms from
separate areas at a site or under separate logs at a site were kept
distinct with "a," "b," and "c" designations, but it was found that
several species of Pheretima would occur in the same shovel of dirt or
under the same. log. Thus, with negative evidence of microhabitat
specificity, the complex method of separate subcollections was abandoned.
All of the specimens in this study were collected with a shovel
or a potato fork. The latter has the advantage of not cutting the
worms in two, as frequently occurs with a shovel. In those areas
3
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. where worms were abundant, "fiddling" was employed. This method
involves inserting the shovel or fork into the ground and rapidly
wiggling it back and forth. The vibrations so produced cause the worms
to come to the surface to escape the disturbance. When fiddling was
employed it was also accompanied by digging.
The specimens in this study were killed in 7-10% ethyl alcohol.
When the specimens were killed in weak alcohol, the spermathecal pores
and dorsal pores are more evident because of the muscular, relaxation.
The worms were left in the alcohol until all movement ceased. They
were then straightened in a flat tray and 10% formalin was dripped over
them with a pipette. After about 10 minutes the worms, together with
a number corresponding to the number of the collection in a field
notebook, were placed in fresh 10% formalin in a test tube inside an
o l iv e j a r .
During warm weather (above 70 F) the worms were killed and fixed
at the collecting site, while in cooler weather a whole day's collec
tion was preserved that night. In the latter case a number cor
responding to the collection site in a field notebook was placed in the
can containing the specimens. When kept for an entire day, the
specimens were placed in dirt from the collecting site at air
temperature.
The pH of the soil was measured with a Hellige-Truog No. 693 soil
t e s t k i t .
Serial sections were cut in order to study the preclitellar
genital marking glands of P. diffringens. Specimens to be sectioned
must have the intestinal tract emptied of soil, which would interfere
with the sectioning. This evacuation was accomplished by keeping the
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. specimens on wet, shredded filter paper in finger bowls until they ate
the paper in quantities sufficient to push the soil and grit out of :
intestine. The finger bowls were cleaned and the filter paper changed
daily until there was no longer any soil in the castings. The
intestine was usually cleaned of soil in 3 days' at 70 F.
These specimens were then killed in 10% ethyl alcohol and fixed
in 10% formalin for about 24 hours. They were then cut into pieces
which possessed the genital marking glands, washed in several changes
of water for one hour and dehydrated in a series of 30%, 50%, 70%, and
95% ethyl alcohol, followed by two changes of 100% ethyl alcohol. The
tissue was left in each of the alcohols for one hour. The tissue was
next placed in xylene for about 15 minutes or until cleared. When
cleared, the tissue was placed in a half and half xylene-paraffin
mixture at 55 C for two hours, followed by paraffin alone for an addi
tional two hours at the same temperature. The tissue was then embedded
in paraffin.
Sections were cut on a rotary microtome at 10 micra and secured
on slides with Mayer's albumen. The sections were stained in Harris'
haematoxalin with eosin as a counterstain. Other stains were tried,
but haematoxalin and eosin gave the best results. Slides were mounted
with Harleco synthetic resin.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. I I I . TAXONOMY AND CONVENTIONS
In numerous papers describing the pheretimas of Burma, China, and
India, Gates (1929, 1935a, 1935b, 1937a, 1939, MS) has worked toward a
more nearly complete knowledge and description of the species occurring
in the Orient. This Oriental fauna includes those peregrine species
that have been introduced into the United States. In a paper on the
species of Pheretima occurring in North America (1937c) he stated that
"the types, and (or) metatypes, or metatopotypes of nearly all the
American species have now been studied." This was necessary because
the early descriptions were very inadequate when judged by modern
standards. In addition, the peregrine species were often described as
new when they were found on a continent or island where they had not
been previously known. Because of these thorough studies, the system
of taxonomy herein followed is that of Gates (MS), which does not
recognize subgenera in the genus Pheretima.
The synonymy of the genus as used in this work is restricted to
those works in which nomenc 1 atural changes have been made and to those
publications that include species of the genus from the United States.
The synonymy included with each of the species is also restricted
to those synonyms that have been used for the species in the United
States. The work or works that contain the most nearly complete account
of the species are also listed.
The conventions used in the generic and specific diagnoses are
those of Gates (1937c), which were taken in part from Stephenson
(1923). The segments are represented by lower case Roman numerals:
6
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. i, ii‘, iii, iv, v, etc. The intersegmental furrows (the thin white
line where the epidermis is thinnest between two segments) and the
septa are characterized by fractions; 10/11 refers to the furrow
between segments x and xi, or to the septum between the two coelomic
cavities of those segments, depending on the context of the sentence.
Fractions are also used with the abbreviation for circumference (C) to
show the distance between male or spermathecal pores: 1/2 C, 1/3 C,
e t c .
The setae in the genus Pheretima are numerous in an equatorial
circle around each segment posteriorly from ii. The number of setae in
the circle is expressed as a range; 20-30/ii, which shows that there
are 20 to 30 setae on segment ii. The convention vi/17-20 shows that
there are 17 to 20 setae ventrally on segment vi between two longi
tudinal lines drawn through the spermathecal pores. These are called
spermathecal setae. Those between the male pores are called male setae
and are expressed as: xviii/11-15. The setae are lettered with a, b,
£, d, designations starting from midventral on each side of the body.
The dorsal setae are lettered £, x, w, starting from middorsal on
each side of the body. The space between two setae is referred to as
ab, be, cd, zy, yx, xw. Thus "a genital marking in £d" means that the
marking is located in the space between lines drawn through setae c
and d parallel to the midventral line. A genital marking may be
anterior to the setal circle if characterized as "presetal" or posterior
to the setal circle in characterized as "postsetal."
Bithecal, quadrithecal, sexthecal, and octothecal refer to one,
two, three, and four pairs of spermathecae respectively. The term
"minute" referring to male or spermathecal pores indicates that they
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 8
are the size of the female pore.
Male pores in Pheretima may be invaginate or superficial. I-n
species with invaginate male pores there may be a copulatory chamber
which extends conspicously into the coelom; or a parietal invagination
which is confined to the body wall (as in P. californica) and does not
extend into the coelom. Superficial male pores are located more or
less level with the surface of the body wall. None of the species in
Louisiana has copulatory chambers at the male pores.
Porophore refers to the area surrounding a male pore and may
include an eversible papilla, which bears wrinkles and genital markings;
or may include only the wrinkles around a superficial male pore.
The clitellum encircles the body (annular) in Pheretima and the
setae are usually dehisced in the clitellar segments before the
formation of the clitellum . In some species a few setae may be retained
ventrally in the clitellar segments.
The prostomium of Pheretima is epilobic like the figure of
Michaelsen (1900a, p. 4) with the posterior border of the prostomium
not divided from the peristomium (segment i) by a groove (tongue open).
Dorsal pores in Pheretima are located in the intersegmental
furrows at the middorsal line. The first dorsal pore is the most
anterior one which is functional. There may be a marking one segment
anterior to this functional pore, but it w ill have no connection with
the coelom. The first functional pore can be determined by pressing
on the body wall and observing whether or not coelomic fluid and
preservative escape from the marking.
One pair of intestinal caeca arise in xxvii as evaginatio-i of
the gut in all Louisiana Pheretima, but may arise more anteriorly or be
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. absent in other species. The term "manicate" refers to a caecum that
is compound or glove-shaped and composed of a number of secondary caeca
as in P. agrestis and P. hilgendorfi (Figure 5). The term "simple"
refers to a caecum with a single pouch (Figure 9).
The testis sacs of Pheretima are either unpaired or paired. An
unpaired testis sac is a somewhat dumbbell-shaped subesophageal chamber
that contains a pair of testes and the associated pair of male funnels.
In some species the unpaired testis sac may be annular, forming a
circular chamber in the segment, as in JP. minima. In the paired
condition, a single testis and its associated funnel on one side of a
segment are enclosed in a sac that has no connection with the sac on
the other side of the segment.
The term "parietes" refers to the internal surface of the body
wall. A structure that is confined within the body wall or located on
the internal surface of the body wall is called "parietal."
Each spermatheca of Pheretima is composed of a large ampulla
connected to the exterior by a large, usually muscular duct. A smaller
diverticulum arises from the duct near the parietes and is composed of
a terminal seminal chamber and a stalk. The sperm received during
copulation are stored in the seminal chamber of the diverticulum
(Figure 1) .
The term "trunk" when used in reference to the vascular system
refers to a longitudinal vessel. Hearts are described as "lateral"
when their dorsal connection is with the dorsal trunk and their ventral
connection is with the ventral trunk. Hearts are described as
"esophageal" when they connect dorsally with the supra-esophageal trunk
and ventrally with the ventral trunk. A "latero-esophageal" heart
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 10
bifurcates dorsally to connect with both the dorsal and supra-
esophageal trunks and connects ventrally with the ventral trunk.
The excretory system of Pheretima is composed of numerous
nephridia that are formed by fragmentation of the ectodermal embryonic
rudiments in each segment (Bahl, 1922). There are also three types of
nephridia in any Pheretima; pharyngeal, septal, and parietal. The
numerous paired tufts, of pharyngeal nephridia in iv-vi are joined
together by their terminal ducts, lack a nephrostome (astomate) and
discharge waste into the pharynx (enteronephric). The septal nephridia
located on both the anterior and posterior face of each septum
posterior to 16/17 have a nephrostome (stomate) and discharge waste
into the intestine (enteronephric). The nephrostome, body, and terminal
duct of these septal nephridia are all located in the same segment.
The parietal nephridia covering the parietes in every segment posteri
orly from iii are astomate and discharge waste to the exterior through
nephridiopores (exonephric). All three types of nephridia lack a
bladder (avesiculate). Thus in every segment except vii-xv two types
of nephridia are found. In addition, each of the types is abundant in
a segment, unlike the condition found in lumbricus terrestris.
In the species localities that follow, the terminal three figures
(1-2-3, etc.) represent the number of specimens secured at the site.
The first figure represents the number of immature specimens. These
have no genital markings, no male or female pores and can be identified
only when one species of Pheretima is present in the collection and
then only tentatively. The second figure represents the number of
aclitellate specimens, which do possess genital markings and genital
pores. The third figure represents the number of clitellate specimens
in the collection.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 11
A. Generic Account
Pheretima Kinberg, 1866
1866 Amyntas + N itrocris + Pheretima + Rhodopis + Perichaeta; Kinb,,
Ofv. Kongl. Vet. Akad. Forhandl, Stockholm, 23:101-102.
(Amyntas, N itocris, Rhodopis and Perichaeta were preoccupied.
Type-species of Pheretima not designated.)
1895 Perichaeta: Beddard, A Monograph of the Order of Oligochaeta,
Oxford, p. 388.
1899 Amyntas: Michaelsen, M itt. N aturhist. Mus. Hamburg, 16:3.
1900 Pheretima: Michaelsen, Zool. Anz., 23:567.
1900 Pheretima; (part) Michaelsen, Das Tierreich, 10:234. (Excluding
burliarensis and P. lawsoni.)
1907 Pheretima: Michaelsen, In Michaelsen, W. and Hartmeyer, H.,
Fauna Sudwestaustraliens, 1:164. (Pheretima montana Kinberg,
1866 designated type species of the genus.)
1934 Pheretima: Michaelsen, Quart. J. Micros. Sci. 77:12. (P.
montana withdrawn as type species of genus. P. californica
Kinberg, 1866 designated type species of genus.)
1937 Pheretima: Gates, Bull. Mus. Comp. Zool. 80:340.
1958 Pheretima: Gates, Am. Mus. Novitates 1888:1
1960 Pheretima: Gates, Bull. Mus. Comp. Zool. 123:246.
MS Pheretima: Gates, "Burmese Earthworms" p. 513.
Diagnosis: Digestive system, without supra-intestinal and
calciferous glands (calciferous tissues in low, nonlamelliform ridges
in region of x iii ?) but with a gizzard that develops in v iii.
Vascular system, with unpaired dorsal, ventral and supra-esophageal
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 12
trunks, a complete subneural trunk adherent to the parietes, paired
extra-esophageal trunks (median to the hearts) that are on the gut
through x-xlll passing off to the subneural, latero-esophageal hearts
In some of x-xlll. Excretory system of multiple nephridia formed by
fragmentation of embryonic rudiments In each segment (merolc); paired
clusters of small astomate nephridia In Iv-vl with ducts opening Into
the pharynx; astomate, exonephric, very small v-shaped parietal
nephridia numerous In each segment from 111 posteriorly; larger stomate
nephridia on both sides of the septa from 16/17 posteriorly, joining
postseptal canals that pass to a longitudinal, supra-Intestinal
excretory duct which opens at frequent Intervals Into the gut. All
nephridia avesiculate. Setae numerous. In a circle at the equator of
each segment from 11 posteriorly. Dorsal pores present.
Blprostatlc, male pores (apertures of united vasa deferentla and
prostatlc ducts) postclitellar. Female pores always minute. In xlv.
Clitellum annular, Intersegmental furrows and dorsal pores obliterated.
Seminal vesicles postseptal. Spermathecae dlvertlculate and In front
o f X . Ovaries restricted to x lll, fan-shaped and with several egg
strings. Type-specles: Pheretima montana Kinberg, 1866.
The above diagnosis Is that of Gates (1 9 6 0 ), revised In a
manuscript on Burmese earthworms (Gates, MS). This diagnosis Is based
on somatic anatomy and follows the classification established by Gates
In a redefinition of the family Megascolecldae to Include only those
forms with racemose prostates of mesodermal origin (Gates, 1 9 5 9 ).
The history and synonomy of the genus Pheretima are quite complex
due to the Incomplete early descriptions that were obtained from an
external examination of the worms with a hand lens. When It was later
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 13
discovered that the internal anatomy also possessed useful taxonomic
characters, the older descriptions were useless and the types, when
available, had to be re-examined.
Schmarda (1861) established the genus Perichaeta for four species
of earthworms from Ceylon with 40-50 setae per segment, having an equal
number of setae on both anterior and posterior segments. Three of
these are now known to belong to the genus Megascolex and the fourth
possibly does (Michaelsen, 1900a).
The genus Pheretima was established by Kinberg (1865) to include
P. montana and P. californica collected in Tahiti and California, which
were described as follows: "Terminal cephalic lobe fixed transversely to
the anterio-dorsal margin of the buccal segment. Setae are disposed
both radially and serially, there being a greater number of setae on
the posterior segments than are found on the anterior ones. Foramen
dorsal; cingulum clitellum ; two ventral tubercules." This is a trans
lation of the Latin description of the genus. In the same paper Kinberg
also erected the genera Amyntas, N itocris and Rhodopis for specimens
that are now known to be Pheretima (Michaelsen, 1899a).
Eighteen years before Schmarda, Templeton (1844) had established
the genus Megascolex for M. caeruleus from Ceylon with a continous band
of setae, excepting a narrow asetigerous gap at mid-dorsal. Schmarda
(1861) misunderstood the description of Templeton and thought Megascolex
to include specimens with mid-dorsal setae only. Baird (1 8 6 9 ), in
describing Megascolex diffringens, pointed out this misconception and
used the older name Megascolex, considering Megascolex and Perichaeta
to be identical. .
By 1895 the distinctions between Megascolex and Perichaeta had
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 14
been established on the basis of the gizzard position, among other
things, and they were treated as separate genera by Beddard (1895).
Michaelsen (1899b) changed the generic name to Amyntas Kinberg,
1866, after showing that Perichaeta was not available, having been used
as a d ip ter o n g e n e r ic name by Rodani in 18 5 9 . •
Michaelsen (1900a, 1900b) again changed the generic name to
Pheretima Kinberg, 1866, after showing that both Amyntas and Nitocris
were not available. Michaelsen (1907) designated Pheretima montana
Kinberg, 1866, as the type-species of the genus Pheretima.
C o g n etti (1914) separated the many Pheretima species into the sub
genera Parapheretima and Pheretima. The subgenus Parapheretima included
those forms with copulatory chambers at the male pores into which open
glands with muscular w alls. The subgenus Pheretima lacked the muscular
walled glands at the male pores. Michaelsen (1928) d e sig n a te d
Pheretima montana Kinberg, 1866, the type-species of the genus to also
serve as the type-species of the subgenus Pheretima.
Ude (1932) showed that P. montana does in fact have glands with a
muscular wall at the male pores and was therefore a member of the sub
genus Parapheretima. According to strict application of taxonomic
rules the members of the subgenus Parapheretima should have been placed
in the subgenus Pheretima and a new subgeneric name coined for the
previous subgenus Pheretima. Michaelsen (1934) responded by stripping
P. montana of its position as type-species and designating Pheretima
(Pheretima) californica Kinberg, 1866 as type-species of the genus and
subgenus Pheretima.
Since the concept of geographic ranges as the basis for recogni
tion of subgenera has become obsolete and the distinguishing characters
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 15
or combinations of characters of the subgenera have been found in
diverse parts of the range, the subgenera of Cognetti (1914) and
Michaelsen (1928) are now obsolete (Gates, MS). In view of these
facts, the type-species of the genus Pheretima should be recognized as
Pheretima montana Kinberg, 1866, by the subsequent designation of
Michaelsen (1907).
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 16
B. Species Accounts
Pheretima agrestis (Goto and Hatal, 1899)
1899 Perichaeta agrestis Goto and Hatal, Annotatlones Zoologieae
Japonenses 3(1);17.
1937 Pheretima hatall Ohfuchl, Research Bull. Salto Ho-on Kal Mus.
Sendai, Japan 1 2 :1 3 -1 8 F ig . 1 .
1958 Pheretima agrestis: Gates, Am. Mus. Novitates 1888:1.
Diagnosis: Sexthecal, spermathecal pores minute and superficial
In 5/Ô-7/8, about 1/3 of circumference (C) apart. Female pore single,
midventral on xlv. Male pores usually absent; when present minute and
located on an eversible papilla. Inside a transverse sllt-llke parietal
Invagination on an elevated porophore. In the setal circle of xviil
about 1/3 C apart. Genital markings usually lacking, but on an
occasional specimen a pair of large presetal papillae In xvlll
(0.8-1.8 mm In diameter), median to the male porophores. If the latter
are present. With transversely ellptlcal, synmetrlcally placed areas
of thin, finely wrinkled epidermis having a light brown color ventrally
In the setal circle of some of Iv-lx, most commonly In v ll-v lll. Setae
absent In these patches at the height of development, but with 1-5
setae mldventrally between the patches. Clitellum annular 13/14-16/17,
dorsal pores, setae and Intersegmental furrows obliterated. Setae,
2 4 -3 7 /1 1 , 32-44/lir, 36-50/lv, 46-6 . 52-76/xll, 51-72/xx, vl/16-24,
vll/1-26, vlll/1-5. First dorsal p-:e usually at 12/13. Prostomium
epilobic with open tongue. Length, 70-200 mm. Diameter, 5-8 mm.
Segments, 67-110.
Septa 5/6-6/7 and 10/11-14/15 thickened, 8/9-9/10 absent.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 17
Intestine beginning in xv. Intestinal caeca manicate, arising in
xxvii and extending forward to about xxiv, with 6-9 seconds y caeca,
the dorsalmost being the longest. Typhlosole low and lamelliform,
arising in xxvii and ending in region of Ixiv-lxxvii. Hearts of x
lacking, those of xi-xiii latero-esophageal. Testis sacs unpaired and
ventral in x and xi. Seminal vesicles in xi and xii, filling coelomic
cavities of those segments. Prostates in xvi-xxii only when male
term inalia are present, large and meeting mid-dorsally, with muscular
U-shaped duct about 5 mm long, thinning at ectal end. In the absence
of male term inalia, the vasa deferentia ending blindly with or without
a swelling in xvii-xxiv without passing into the parietes. Spermathecae
in vi, vii, and v iii. Spermathecal duct with thick wall, shorter than
ampulla. Ampulla dorsoventrally flattened with thin wall and numerous
wrinkles. Diverticulum longer than combined lengths of duct and ampulla,
with sausage-shaped seminal chamber. Genital marking glands low and
confined to or extending only slightly above the parietes.
The above diagnosis is a composite t^en from the original
description of Goto and Hatai (1899) and from other references covering
this species.in Korea, Japan and the United States (Gates, 1953a, 1954,
1958; Hatai, 1930; Kobayashi, 1938; Yamaguchi, 1962).
Discussion: The 146 specimens of P. agrestis examined in this
study agree with the above diagnosis except for the number of sperma
thecal setae, v/12-22 (literature: no record), vi/14-23 (vi/16-24),
vii/11-24 (vii/1-26), viii/8-25 (viii/1-5); and in a more posterior
extent of the typhlosole back to Ixxxi (Ixxvii).
The clitellate, nonautotomized specimens in this study ranged
from 74-174 mm long and from 4.5-7 mm in diameter; the number of
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 18
segments ranged from 95-111. The average length of 103 of these
specimens was 100 mm, the number of segments averaged 104.
Pheretima agrestis is a very active species that crawls rapidly,
but it does not jump and thrash about on the surface of the ground when
disturbed as does P. diffringens. It may autotomize a posterior portion
of the body when disturbed, as did 35 of the 146 specimens collected in
this study. The level of autotomy ranged from 56/57-97/98, and was
always in the intersegmental furrow.
The recognition of posterior regenerates in P. agrestis is
impossible on the basis of criteria employed for other species of
earthworms (Gates, 1953a). In most species of earthworms, a posterior
regenerate can be recognized by the shorter length and smaller diameter
of the individual regenerated segments at the posterior end of the body.
In freshly preserved m aterial, the regenerated segments also have a
lighter color, but on longer preservation the older segments fades, and
the regenerated portion can only be recognized by the shorter length
and smaller diameter of the individual segments. Gates (1953a) has
observed that the regenerated segments in P. agrestis are produced in
a slow, one-at-a-time manner and that the regenerated segments attain
a size comparable to that of the older segments, before another new
segment is completed by the growth zone. He also found evidence that
adult worms produce segments in the same manner as in regeneration.
The specimens of JP. agrestis in Louisiana tend to support his
conclusions. In a specimen of 65 segments (counting the growth zone as
a segment), which had previously autotomized at 64/65, Ixiv s till had
some setae ventrally, while the setal follicles were visible dorsally
where the former setae had fallen out. The growth zone in this
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 19
specimen was 0.64 mm long and was divided by an intersegraental furrow
at 64/65. There was no dorsal pore at 64/65. In a second specimen of
66 segments (counting the growth zone as a segment), which had
previously autotomized at 65/66, the setae had all fallen out of Ixv
and a growth zone 0.80 mm long was formed behind Ixv. There was a
dorsal pore and intersegmental furrow at 65/66, but the growth zone
bore no setae nor was there any indication of an intersegmental furrow
at a future site of 66/67. In a third specimen of 94 segments (counting
the growth zone as a segment), which had previously autotomized at
93/94, there was a growth zone 1.36 mm long, while x ciii was 1.12 mm
long. The dorsal pore and intersegmental furrow at 93/94 were present,
but the growth zone (xciv) had no setae, nor was there any indication
of a future intersegmental furrow at 94/95.
À sim ilar growth zone was found in clitellate P. agrestis of 96,
97, 98, 100, 106, 107, 108, and 111 segments. Unless every specimen of
P. agrestis in this study was a posterior amputee in the past, segments
are added in mature adults by the same method employed in regeneration.
In other species in this study (except _P« hilgendorfi) . growth
zones of these lengths would be divided by intersegmental furrows into
a number of smaller segments, rather than one long segment. In
P. diffringens regenerated segments measured 0.25-0.44 mm long, while
normal old posterior segments measured 0.98-1.28 mm long.
The following lis t shows the number of specimens (in parentheses)
that possess the characteristic areas of thin, brown, wrinkled epidermis
on the segments indicated by Roman numerals; v ii-v iii (50), v ii (19),
vi-viii (6), vii-ix (3), v-viii (2), vi-vii (1), viii-ix (1). While all
of the specimens of JP. agrestis in this study possessed these areas of
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 20
modified epidermis, records were kept only for the 82 specimens referred
to above. These areas are more evident in preserved specimens that have
been allowed,to dry somewhat and after the removal of the cuticle
(Figure 3).
Two of the 146 specimens examined in this study had a single male
pore and the associated genital marking in xviii, on the right side of
one specimen from the Shreveport site in Caddo Parish and on the left in
the specimen from the Bayou Macon site in Franklin Parish. A third
specimen possesses a pair of genital markings in xviii but has no male
term inalia, while a fourth specimen has a single genital marking on the
left in xviii and also lacks male term inalia. These specimens
represent the first United States records of P. agrestis possessing
either male term inalia or the associated genital markings in xviii.
In the original description. Goto and Haiti (1899) found a few
specimens with the genital markings in xviii but none with male
terminalia in a series of more than a hundred specimens. In a later
addition to the distribution of P. agrestis in Japan (Hatai, 1930),
mention is made of specimens with male term inalia but no description
is given. Yamaguchi (1930b) found male term inalia and genital markings
in all of the.specimens of P. agrestis found at Sapporo, Hokkaido,
Japan but did not describe or figure them adequately. In P. agrestis
from Korea, Kobayashi (1938) found male pores in 2 specimens of 248,
but did not adequately describe them.
In both specimens with male term inalia in the present study, the
male porophore is an elevated mound about 1.3 mm in diameter located in
the setal circle of xviii, with a central transverse aperture. This
aperture leads to a shallow parietal invagination and the margins of
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 21
the aperture are surrounded by numerous wrinkles of the epidermis.
Inside this shallow parietal invagination, there is a mound-shaped
papilla on which the minute male pore is located. Judging from the
spongy internal nature of this papilla, it is eversible through the
opening of the male porophore. However, in both of the specimens the
papillae are contained within the aperture of the male porophore
(Figure 2 and Figure 4).
In the specimen with the male porophore on the right side, there
were 6 setae between the male porophore and the nerve cord, while in
the other specimen the genital marking extends into the setal circle of
xviii; so there were only 2 setae in tnis location. On xix there were
10-12 setae between the middle of the aperture of the male porophore
and the nerve cord.
Internally, the prostates are present only on the side of the
body that bears the male porophore and extend from 15/16-21/22,
reaching the level of the dorsal blood vessel. They are divided into
3 or 4 main lobes and are segmentally constricted by the septa. The
vas deferens passes into the prostatic duct close to the connection of
the latter with the prostate (Figure 6). The muscular prostatic duct
is about 5 mm long and at its junction with the papilla of the male
pore, it loses its muscular nature, becomes spongy, and fuses with the
tissue of the papilla (Figure 2).
Pheretima hataii Ohfuchi, 1937, was established for a collection
of specimens that differ from P. agrestis only in the absence of the
areas of thin, brown, wrinkled epidermis on v iii. They possessed these
characteristic areas only in v ii. This difference is not sufficient to
warrant the erection of a separate species, because 19 of the specimens
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 22
collected in the present study possess these areas only in v ii, with
otherwise indistinguishable morphology.
D istribution; Pheretima agrestis is widely distributed in Japan,
its presumed original home, having been reported from the four principal
Japanese islands (Goto and Hatai, 1899; Hatai, 1930; Yamaguchi, 1930b).
It has also been reported from Korea (Kobayashi, 1938). In the
United States the distribution is widely scattered, presumably the
result of a number of introductions. It has been found in greenhouses
or sim ilar inside habitats in New York (Gates, 1954) and Maine (Gates,
1965) . In natural outside environments it has been found in Maryland
(Howell, 1939); Massachusetts (Gates, 1953a); New Jersey (Davies,
1954); New York, Connecticut, Louisiana (Gates, 1958); Missouri
(Stebbings, 1962); and Pennsylvania (Bhatti, 1965).
In Louisiana, P . agrestis was found in 13 parishes at 17 s i t e s
limited to the upland pine area approximately northeast of the Red
River drainage (Map 1 ) . Eight of the sites are the margins of lakes
or bayous where it may have been introduced by fishermen. Four of the
sites are in flower beds where it may have been introduced with exotic
plants. Three of the sites are dumps and one site each is from a
sawmill and a pasture. The worms in the Jackson Parish record were
stated to have been brought from Caddo Parish for the purpose of
establishing a bait population.
Unlike P. diffringens, this species has not as yet dispersed
over large areas following introduction but is limited to areas
influenced by the activities of man. In the East Carroll Parish site
from the shore of Lake Providence, this species could be collected
near a boat landing for a distance of about 200 yards, but 2 .5 m ile s
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 23
north of this site no specimens could be secured in the same type of
habitat. This lack of rapid dispersal may be due to the annual life
cycle in which cocoons are produced in the fall and hatch late in the
spring, as evidenced by the fact that immatures were found only in
April, May and June, while clitellate specimens were found only from
May to Septemoer. Attempts to recollect sites known to contain P.
agrestis were futile after September. This species apparently has a
life history sim ilar to that of _P. hilgendorfi in which the adults die
in the fall and the species overwinters in the cocoon.
In the northeastern United States, clitellate specimens have been
found as early as June in New Jersey (Davies, 1954) and as late as
November in Pennsylvania (Bhatti, 1965) . Only immatures were collected
in June in Boston (Gates, 1953b), clitellate specimens being found in
September (no collections in the interim) (Gates, 1953a).
Localities;
Bossier Parish
3.1 air mi N Lake Bistineau Dam, near Cryer's Camp, 24 Aug. 1966,
0- 0- 2 .
Caddo Parish
Shreveport, 0.1 mi S Cross Bayou on N Common S t., 25 Aug. 1966,
0- 0- 11 .
Spring Ridge, pasture, 10 June 1967, 0-0-4.
1 mi. W US Hwy 171 on La. Hwy 511, 10 June 1967, 0-1-29.
Claiborne Parish
Homer, K eller S t., 11 Aug. 1966. 0-0-8.
East Carroll Parish
3.1 mi NW La. Hwy 596 on US Hwy 65, 11 July 1966, 0-0-8.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 24
Franklin Parish
Winnsboro city dump on Eighth St., 9 July 1966, 0-0-5.
1.3 mi S Lcrelein along Bayou Macon, 18 May 1967, 0-0-8.
Jackson Parish
Jonesboro, Old Gansville Rd., 31 Aug. 1966, 0-0-5.
Lincoln Parish
Ruston, W M ississippi S t., May 1959, 0-4-2.
Ruston, La. Polytech. Inst. Farm Pond, 28 May 1966, 0-1-17.
Morehouse Parish
Near Arkansas line on Bayou Bartholomew, 31 July 1966, 0-0-1.
Natchitoches Parish
Natchitoches, Cane River at Robbins Dr., 26 May 1966, 0-2-17.
Ouachita Parish
Sterlington, La. Hwy 553 a t US Hwy 1 6 5 , 9 Aug. 1966, 0-0-16,
Richland Parish
Clear Lake on La. Hwy 133, 11 Sept. 1965, 0-0-1.
Union Parish
Marion, old sawmill on La. Hwy 551, 1 Aug. 1966, 0-0-1.
West Carroll Parish
4.5 mi N Oak Grove on La. Hwy 1 7 , 12 J u ly 1966, 0-0-3.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 25
Pheretima californica Kinberg, 1866
1866 Pheretima californica Kinberg, (part) Ofv. Kongl. Vet. Akad.
Forhandl. Stockholm 23(4):102. (Type from Sausolita Bay,
California and two paratypes from San Francisco. Excluding
octothecal specimens)
1937 Pheretima californica; Gates, Bull. Mus. Comp, Zool. 80:348-
3 50.
Diagnosis : Quadrithecal, spermathecal pores minute and
superficial in 7/8-8/9, about 1/3 G apart. Female pore single mid-
ventral on xiv. Male pores minute, on a very small tubercule at the
end of a columnar protuberance from the roof of an eversible, deep,
transversely slitlike parietal invagination in the setal circle of
xviii, about 1/3 C apart. No genital markings. Clitellum annular
13/14-16/17, dorsal pores, setae and intersegmental furrows obliterated.
Setae slightly enlarged ventrally in some or all of iii-ix , 20-30/iii,
28-40/vi, 35-48/viii, 49-60/xii, 43-61/xx, viii/12-20, xviii/9-20.
First dorsal pore at 1 1 /1 2 . Prostomium epilobic with open tongue.
L ength, 50-156 mm. Diameter, 3-5 mm. Segments, 8 5 -1 1 5 .
Septa 5/6-7/8 and 10/11-13/14 muscular, 8/9-9/10 absent.
Intestine beginning in xv. Intestinal caeca simple, arising in xxvii
and extending forward to about xxii, with or without very short dorsal
and/or ventral lobes. Typhlosole low and lamelliform, arising in xvi
and extending back to the region of Ixiv-lxxiv. Hearts of x
esophageal, those of xi-xiii latero-esophageal. Testis sacs unpaired
and ventral in x and xi. Seminal vesicles in xi and xii. Prostates
in xvi-xxii with muscular ducts 1.5-3 mm long, narrow and looped just
before passing into male invagination, which extends slightly above
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 26
the parietes. Spermathecae in v iii and ix, anterior pair sometimes in
front of septum 7/8. Spermathecal duct shorter than globular ampulla
into which it is invaginated. Diverticulum arising from anterior face
of muscular spermathecal duct near parietes. Diverticulum longer than
combined lengths of duct and ampulla, coiled and looped. Stalk of
diverticulum shorter than elongated seminal chamber.
Since the original description is very brief, the above
diagnosis is taken from several publications (Gates, 1937b, 1937c,
1939; Tsai, 1964) and a manuscript by Gates entitled "Burmese
Earthworms."
Discussion; The 399 specimens of P. californica collected in the
present study conform well with the above diagnosis, differing only in
a greater length, 177 mm (Literature:156 mm); in a more posterior extent
of the typhlosole back to Ixxvi (Ixxiv); and in numbers of setae on
various segments, 23-31/iii (20-30/iii), 17-31/iv (no record),
27-37/vi (28-40/vi), 38-50/viii (35-48/viii), 50-63/xx (43-61/xx),
vii/11-18 (no record), viii/14-22 (viii/12-20), ix/16-22 (no record),
xviii/10-21 (xviii/9-20).
The clitellate, nonautotomized specimens in this study ranged
from 54-177 mm long and from 2.5-5 mm in diameter, the number of
segments ranged from 93-115. The average length of 198 of these
specimens was 98 mm, the number of segments averaged 106. There is no
correlation between the number of segments and the length of the worm,
because the shortest worm possessed 108 segments, while the longest
had 105 segments.
Pheretima californica is an active species that jumps and
thrashes about on the surface of the ground when disturbed. It may
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. . 27
autotomize a posterior portion of its body as did 34 of the 399
specimens collected in this study. The level of autotomy ranged from
57/58-94/95, and was always in the intersegmental furrow,
.Tail regeneratates, which can be recognized by the lighter color,
shorter length and smaller diameter of the new segments, were observed
at various levels between 75/76 and 93/94. At the anterior end of this
range 6 segments were added, while at the posterior end of the range
only 3 segments were regenerated.
D istribution; Pheretima californica is widely distributed in
China, its presumed original home, and has also been reported from
A ustralia, Egypt, Madeira, Central and South America, Easter Island,
the Hawaiian Islands, Marquesas Islands (Gates, 1937c), and Taiwan
(Tsai, 1964), In the United States it has been reported from
California (Kinberg, 1866); an Arkansas worm farm (Causey, 1952);
Louisiana (Harman, 1952); outside a greenhouse in New York, and
Louisiana (Gates, 1954). The Lincoln Parish record of _P«
californica (Harman, 1952) is approximately 150 miles north of any
locality in the present study. No information is given as to whether
this record is based on worms from a greenhouse or an outside habitat.
The specimens are not at present in the Harman collection; so perhaps
that record is best considered doubtful.
In the present study, P, californica has been found in 20
parishes at 70 sites, widely distributed on the lower floodplains of
the M ississippi and Atchafalaya Rivers (Map 2), Thirty-three of the
sites are the margins of lakes and swamps or the floodplains of bayous
or rivers, 19 of the sites are in flower beds or similar sites in
yards, 7 sites are wooded areas, 5 sites are in cemeteries, 2 sites
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 28
each are in dumps and cultivated farm land, and one site each in the
dried sludge from a sewage plant and a sawmill. While P. californica
has been found in a number of areas where it may have been introduced
by man, the number of sites in which it was found widely removed from
the activities of man suggest that it has been able to colonize large
areas by its own activities. This species is very common in the black,
heavy clay of the southeastern portion of the state and at 10 sites was
found living in clay soil that was devoid of any vegetation or humus
c o v e r in g .
Immature specimens were found in April and May, indicating that
in Louisiana this species produces cocoons that hatch in the spring.
The adults apparently survive the entire year, as clitellate
specimens were collected in every month of the year except January and
September. C litellate specimens have been reported from Louisiana in
December and January and from New York in August and November (Gates,
1954) .
Localities;
Ascension Parish
Donaldsonville, Charles St. at Lafourche St., 28 A p ril 1965, 1-5-4.
Donaldsonville Fairgrounds, 28 A p ril 1965, 2-2-7.
E b an k M iss. River at La. Hwy 22, 28 April 1965, 0-0-3.
Assumption Parish
Bellerose, on Bayou Lafourche, 14 A p ril 1967, 0-0-9.
5.5 mi N Pierre Pass on La. Hwy 70, 14 April 1967, 0-0-3.
Near Lafourche Par. line on La. Hwy 1, 14 April 1967, 0-2-6.
East Baton Rouge Parish
L. S. U. Campus, b e s id e Dodson Pond, 3 May 1964, 0 - 0 -3 .
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 29
W Lakeshore Dr. near Corporation Canal, 3 Nov. 1964, 0 - 1 - 0 .
0.7 mi E Corporation Canal on W Lakeshore Dr., 3 Nov. 1964, 0-2-7.
Stanford Ave., 28 Nov. 1964, 0-0-1.
0.2 mi S Highland Rd. on Ben Hur Rd., 7 Feb, 1965, 0-1-3.
0 .2 mi N Highland Rd. on Nelson D r ., 7 F eb . 1965, 0-1-8,
Gardens near State Capital, 1 A p r il 1965, 0-0-1.
Highland Rd. at Dentation Dr., 8 April 1965, 0-0-1.
W. Parker Blvd. at Dodson Ave., 8 April1965, 0-0-9.
L. S. U. Campus, Audubon Hall, 29 May 1965, 0-0-1.
Iberia Parish
Avery Island, Jungle Garden, Timber bamboo, 27 A p r il 1965, 0-2-14.
Avery Island, Jungle Garden, Camélia garden,27 A p r il 1965, 0-1-2.
Avery Island, Jungle Garden, near Buddha, 27 A p r il 1965, 0-0-1.
New Iberia sewage plant on La. Hwy 83, 28 April 1965, 0-3-6.
Jeanerette, E Main S t., 5 March 1966, 0-0-2.
Iberville Parish
Plaquemine, Desorby St. at Calvin St., 17 May 1964, 0-1-2.
1.6 mi NW Sunshine on La. Hwy 337, 12 Dec. 1964, 0-1-4.
Spanish Lake, 4 mi S L a. Hwy 427 on La. Hwy 9 2 8 , 1 A p ril 1965, 0-0-5.
Jefferson Parish
1 .4 mi NW Bayou Barataria on La. Hwy 2 3 , 23 A p r il 1967, 0-0-15.
Lafayette Parish
Youngsville, cemetery on La. Hwy8 9 , 1 A p r il 1967, 0-1-1.
Broussard, R. U. Bernard Park, 1 A p r il 1967, 0-0-1.
0.3 mi S Vermillion River on La. Hwy 182, 1 April 1967, 0-0-1.
Lafourche Parish
8.5 mi W Larose on La. Hwy 24, 8 April 1967, 0-0-3.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 30
0.7 mi S Lockport on La. Hwy 1, 9 April 1967, 0-0-13.
2 .7 mi SW Allemands on old US Hwy 90, 9 A p r il 1967, 0-0-9.
Thibodaux, La. Hwy 1 at Cherokee Ave., 9 A p r il 1967, 0-0-1.
2 mi W Thibodaux on La. Hwy 1 , 9 A p r il 1967, 0-0-7.
Orleans Parish
Canal Blvd. at Chappelle S t., 26 June 1965, 0-0-2.
Audubon Park, 26 June 1965, 0-0-4.
0.7 mi N Intracoastal Canal on La. Hw^r 47, 23 April 1967, 0-0-10.
Chartres St. at Ursulines Ave., 28 April 1967, 0-3-1.
L ittle Woods on Hayne Dr., 28 April 1967, 0-0-8.
Plaquemines Parish
Near St. Bernard Par. line on La. Hwy 39, 22 April 1967, 0-0-1,
Knobloch Plantation near Dalcour, 22 April 1967, 0-0-1.
3 mi S Pointe a la Hache on La. Hwy 39, 22 April 1967, 0-0-1.
1 mi S Venice on La. Hwy 23, 22 April 1967, 0-0-1.
Pointe Coupee Parish
3.2 mi N Oscar on La. Hwy 1, 14 March 1965, 0-0-1.
Saint Bernard Parish
4.6 mi E Poydras on La. H\fy 46, 22 April 1967, 0-0-5.
2 .1 mi S Orleans Par. line on La. Hwy 4 7 , 23 A p r il 1967, 0-0-4.
Chalmette, De la Ronde House, 23 A p r il 1967, 0-1-8.
Saint Charles Parish
2.4 mi NW Norco on US Hwy 61, 21 Feb. 1965, 0-0-7.
1 mi NE US Hwy 61 on W Bonnet Carre Spillway levee, 13 March 1965,
0 - 0 - 5 .
3 mi NE Norco on E Bonnet Carre Spillway levee, 20 Dec. 1965, 0-0-1.
2 mi NE Norco on E Bonnet Carre Spillway levee, 20 Dec. 1965, Ot O -2.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 31
Luling on La. Hwy 5 2 , 29 A p r il 1967, 0-0-6.
Saint James Parish
5.5 mi NW Gramercy on US Hwy 61, 13 Nov. 1965, 0-0-3.
4 mi S A scen sio n P a r . li n e on La. Hwy 44, 5 D ec. 1965, 0-0-1,
1 mi S Convent on L a . Hwy 4 4 , 5 D ec. 1965, 0-0-2.
5 m i S V a ch erie on L a. Hwy 2 0 , 5 D ec. 1965, 0-0-8.
Saint John The Baptist Parish
7.2 mi N. Laplace on US Hwy 51, 13 Nov. 1965, Ot O -7.
2.3 mi S Laplace on La. Hwy 628, 30 April 1967, 0-0-5.
2 .5 mi N Killona on La. Hwy 18, 30 A p r il 1967, 0-0-3.
3.6 mi S Edgard on La. Hwy 640, 30 April 1967, 0-0-5.
Saint Landry Parish
1.5 mi S Cortableau on levee rd., 20 Feb. 1965, 0-0-1.
Bayou Cortableau at US Hwy 190, 15 Oct. 1966, 0-0-1.
Saint Martin Parish
Lake Martin at La. Hwy 3 5 3 , 20 F eb . 1965, 0-0-43.
Saint Mary Parish
1 mi E Franklin on US Hwy 90, 28 April 1965, 0-0-16.
Morgan City, cemetery on Young Dr., 2 April 1967, 0-0-5.
Terrebonne Parish
Gray on La. Hwy 24, 22 Aug. 1966, 0-2-1.
Crozier, cemetery on La. Hwy31 5 , 7 A p ril 1967, 0-0-11.
2 .3 mi SE G ibson on US Hwy 90, 8 April 1967, 0-3-3.
West Baton Rouge Parish
5 mi E Rosedale on La. Hwy 76, 31 Dec. 1964, 0-0-1.
4.3 mi W Port Allen on La. Hwy 76, 17 April 1965, 0-1-3.
4.2 mi W Brusly on La. Hwy 989-1, 23 Oct. 1965, 0-0-23.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 32
Pheretima diffringens (Baird, 1869)
1869 Megascolex diffringens Baird, Froc. Zool. Soc. London 1 8 6 9 :4 0 -
4 3 .
1866 Pheretima californica Kinberg (part), Ofv. Kong. Vet. Akad.
Forhandl. Stockholm 23:102. (Two paratypes from Sausolita Bay,
California. Excluding quadrithecal specimens.)
1894 Perichaeta indica: Michaelsen, Z o ü l. Jahrb. S y s t. 8 :1 9 1 -1 9 4 .
1895 Perichaeta indica: Ude, Z. Wiss. Zoül. 61:129.
1915 Pheretima heterochaeta: Smith, Bull. Illinois State Lab. Nat.
H is t . 1 0 :5 5 7 .
1928 Pheretima heterochaeta: Smith, Illinois Nat. Hist. Survey Bull.
1 7 :3 5 9 .
1929 Pheretima indica: Kindred, J . Morph. 4 7 :4 4 1 .
1933 Pheretima heterochaeta: Stephenson, Proc. Zool. Soc. London.
1 9 3 2 :9 1 6 .
1937 Pheretim a d if f r in g e n s : G a tes, B u ll. Mus. Comp. Z o o l. 80:350-352.
1942 Pheretima (indica?): Liebmann, J. Exptl. Zool. 91:379.
Diagnosis: Octothecal, spermathecal pores minute and superficial
in 5 / 6 - 8 / 9 , about 1 /3 C or a little more apart. Female pore single,
midventral on xiv. Male pores minute and superficial on circular to
transversely elliptical disc-shaped porophores in the setal circle of
xviii, about 1 /3 C apart. Genital markings small ( 0 .3 - 0 .5 mm in
diameter), circular to elliptical discs with a central translucence
located in six longitudinal rows in some or all of v-ix. Two rows of
postsetal markings in line with the spermathecal pores, each marking
located just anterior to the spermathecal pore in v -v iii. Four rows
of presetal markings in vi-ix, the two lateral rows with each marking
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 33
positioned just median and posterior to a spermathecal pore. The two
median presetal rows located in ^ or cd, each marking just median and
posterior to the lateral presetal marking. One or more small genital
markings in each male porophore in xviii. A complete set of such
markings probably never present and on an occasional specimen lacking
entirely. Clitellum annular 13/14-16/17, occasionally extending
slightly into x iii and (or) xvii, occasionally with a few setae
ventrally in xvi, dorsal pores and intersegmental furrows obliterated.
Setae, enlarged ventrally in iii-x iii, size decreased laterally from a
to d in iii-xiii, in iii-vi a, b, or c follicles protrude conspicuously
into the coelom, more widely separated ventrally in iv-vii, enlarged
setae ornamented ectally, 21-28/iii, 26-36/vi, 26-46/viii, 39-46/xii,
39-54/xx, vi/6-11, vii/8-14, viii/10-15, xviii/8-16. First dorsal pore
usually at 11/12, occasionally at 10/11 or 12/13. Prostomium epilobic
with open tongue. Length, 45-170 mm. Diameter, 3-6 mm. Segments,
79-121, but usually 105-118.
S ep ta 5 /6 - 7 /8 and 10/11-12/13 slightly muscular, 8 /9 -9 /1 0 a b se n t.
Intestine beginning in xvi. Intestinal caeca simple, arising in
xxvii and extending forward to about xxii. Typhlosole simply lam elli
form, arising in xxvii and extending back to the region of Ixxi-lxxxvi.
Hearts of x esophageal when present but usually absent, those of x i-x iii
latero-esophageal. Testis sacs unpaired and ventral in x and xi.
Seminal vesicles in xi and x ii. Prostates in xvi-xxii with muscular
ducts up to 6 mm long, looped. Spermathecae in vi, v ii, v iii, and ix.
Muscular spermathecal duct slender, shorter than pear-shaped ampulla.
Diverticulum shorter than combined length of duct and ampulla, arising
from anterior face of duct at the parietes. Small, spherical to
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 34
ellipsoidal seminal chamber well differentiated from long, slender
stalk of the diverticulum. Genital marking glands, stalked, coelomic,
bound down to parietes or retained within body wall, composite.
Since the original description is very inadequate, the above
description is taken from the publications of Gates, who has examined
the type m aterial (1937a, 1937c, MS).
Discussion; The 2240 specimens of _P. diffringens collected in
Louisiana conform to the lim its of the above diagnosis with the excep
tion of a greater total length, 203 mm (170 mm) and in the number of
setae on various segments, 22-36/vi (26-36/vi), 38-52/xx (39-54/xx),
v/6-10 (no record), vi/8-12 (vi/6-11), viii/12-17 (viii/10-16), ix/12-17
(no record).
The c lite lla te, nonautotomized specimens ranged from 61-203 mm
long and from 2.5-5.5 mm in diameter, the number of segments ranged
from 81-119. The average length of 1159 of these specimens was 109 mm.
The average number of segments in 200 of these specimens was 108. There
is no correlation between the length of the worm and the number of
segments as the shortest worm had 108 segments, while the longest worm
possessed 111 segments. The maximum number of segments was recorded
from worms 132, 135 and 150 mm long.
Pheretima diffringens was so named by Baird (1869) fo r i t s "...
habit of breaking into pieces" or autotomizing. It is a very active
species which jumps and thrashes around by violent contractions of the
body when it is disturbed. In such circumstances it may also auto
tomize a posterior portion of the body as did410 o f th e 2240 specim ens
collected in this study. The level of autotomy was always at the
intersegmental furrow and occurred from 50/51-110/111. In 103 specim ens
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 35
the level of autotomy was as follows; 50/51-59/60 (10 specimens),
60/61-69-70 (21), 70/71-79/80 (22), 80/81-89/90 (22), 90/91-99/100 (20),
100/101-109/110 (7), 110/111 (1). These data show that autotomy is
more frequent in the range 60/61-99/100, tapering off at either end of
th e r a n g e .
Tail regenerates, which can be recognized by the lighter color,
shorter length and smaller diameter of the regenerated segments, were
observed at various levels between 52/53 and 107/108. At the anterior
lim it of this range 11 segments were added, while at the posterior
lim it 4 segments were added. The maximal number of 20 regenerated
segments was observed twice, at 66/67 and 86/87. The minimal number of
3 regenerated segments was observed at various levels in those
specimens that had just begun the process of regeneration.
Liebmann (1942) reported from a study done in New Orleans, that
diffringens operated on in December and January did not begin to
regenerate until the end of February, while those operated on in
February began to regenerate immediately. He concluded that the
inhibition of regeneration in December and January might be caused by a
fall or winter diapause. The data in this study, however, do not support
such a conclusion since regenerates and active specimens were collected
in every month of the year.
The preclitellar genital markings (GMs) found in _P. diffringens
are subject to a great range of variation, both in numbers found on a
specimen and in possible combinations. From the diagnosis and
Figure 10, it can be seen that there is a possibility of 24 pre
clitellar GMs occurring on a specimen. The highest number of
preclitellar GMs on a specimen in this study was 16 in one instance.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 36
while 56 of the specimens lacked GMs entirely. Table 3 shows the total
number of preclitellar GMs per specimen in 1087 specimens. The peak in
Table 3 at 6 GMs is due to the most common pattern of GMs, paired
median presetal in vii-ix found in 306 specimens (Figure 12). The next
most frequent patterns were paired median presetal GMs in v iii-ix on 91
specimens, and in v ii-v iii on 78 specimens producing the peak at 4 GMs
in Table 3. In 45 specimens the only GMs present were the paired
median presetal GMs in v iii. In 19 specimens only paired median
presetal GMs were present in vi-ix.
As can be seen in columns B and E of Table 5, the lateral pre
setal GMs were less common than the median presetal GMs. In 44
specimens both paired lateral and median presetal GMs were present in
vii-ix (Figure 11). In 33 specimens paired lateral presetal GMs were
found in vii-ix, together with paired median presetal GMs in v iii-ix .
The postsetal preclitellar GMs are noticeably lacking in Louisiana
specimens of P. diffringens. Only 4 specimens in a series of 1087
possessed these GMs as can be seen in Table 4«
The remainder of the 1087 specimens had various combinations of
paired or unpaired, lateral and (or) median presetal preclitellar GMs.
Table 4 shows the number of specimens which had at least one GM on the
segments indicated.
The postclitellar GMs of P. diffringens are much smaller than
those found in P. hawayana and are found in the series of concentric
wrinkles of the male porophores, either pre- or postsetal, either
median or lateral to the male pores. At least one of these GMs was
present in 51 o f th e 1087 specimens for which records of the GMs were
k e p t.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 37
The function of the internal genital marking glands, located at
the site of the external GM, is at present unknown, but Beddard (1890)
proposed several possible functions. He considered that because of
their location in areas around the spermathecae and male pores, they
were of some special significance in reporduction. He supposed that
they might be glands that secrete albumin for the cocoon or that they
might secrete mucus used to attach two worms together during copulation.
I would like to suggest that these might be pheromone glands.
Microtome sections of these glands were made and are illustrated
in Figure 13. The figure is a composite drawing since the stalked
portion of the gland curves anteriorly in passing to the secretory end
piece, thus no one section contained an entire glandular apparatus and
stalk. In the sections at hand it looks as though the cells of the
secretory end piece have individual elongate tubes extending to the
exterior, but this cannot be definitely traced.
Gates (1937c, 1939) has mentioned that prostates may be rudi
mentary or entirely lacking in P. diffringens. In a total of 111
specimens in this study, prostates were absent in 58 specimens,
rudimentary and confined to xviii in 22 specimens, well developed on
one side and absent on the other side in 20 specimens, well developed
on both sides in 7 specimens, rudimentary on one side and absent on the
other side in 4 specimens. The prostatic duct is usually present even
though the prostate is lacking.
D istribution; P. diffringens has been reported from India, Ceylon,
Burma, China, Taiwan, Japan, Sumatra, Java, the Philippines, New
Caledonia, A ustralia, New Zealand, Madagascar, South Africa, Egypt,
Europe, the B ritish Isles, Russia, St. Helena, Cape Verde, Hawaiian and
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 38
F iji Islands, Central and South America (Gates, MS). In the colder
portions of some of the above areas it is found only in greenhouses.
_P. diffringens is the most widely distributed species of the
genus in the United States. It has been reported from natural outdoor
sites in Florida, Georgia (Michaelsen, 1894; Ude, 1 8 9 5 ); V ir g in ia
(K indred, 1929) ; North Carolina (Stephenson, 1933) ; Alabama, California,
Florida, Georgia, Louisiana, M ississippi (Gates, 1 9 3 7 c ); L o u isia n a
(Harman, 1952)'; Arkansas (Causey, 1952, 1953); Alabama, Arkansas,
Connecticut, Florida, Louisiana, North Carolina, Nebraska, Texas,
Virginia (Gates, 1 9 5 4 ); Pennsylvania (Bhatti, 1 9 6 5 ); New Jersey,
South Carolina, and Tennessee (Gates, 1 9 6 6 ). It has also been reported
from greenhouses in Illin ois (Smith, 1 9 1 5 ); Oregon (Gates, 1 9 5 4 );
Maine (Gates, 1963); and New Jersey (Gates, 1966).
In the present study, P. diffringens was collected at 376 sites
in all 64 parishes of Louisiana (Map 3). It is a common inhabitant of
the floodplains of bayous, creeks, rivers and the margins of lakes or
swamps where it was found at 195 sites. It was found in deep woods at
76 sites. At 125 of the above sites P. diffringens was found in or
under logs. Seven of these logs had been carried by flood waters and
recently deposited in new areas presumably carrying the worms with them.
This method of dispersal has probably accounted for a great deal of
downstream distribution of this species in Louisiana.
At 26 sites P. diffringens was collected from flower beds or
compost piles in yards. P. diffringens is also a common inhabitant of
bark and sawdust piles around sawmills where it was found in 23
instances. At several of the sawmills it was being used as a source of
bait by local inhabitants, further increasing the chances of wider
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 39
distribution. At 21 sites it was found under grassland or pasture in
all types of soil, from the red dirt of North Louisiana to the heavy
black clays of South Louisiana. Nine of the collections came from city
dumps. Five collections were obtained from botanical gardens where the
worms may have been introduced with plant m aterial. P. diffringens was
collected in 5 cemeteries where it may have been introduced with potted
plants. At 4 sites P. diffringens was found in cultivated soil. In 4
instances it was found in barnyard situations in soil mixed with cow
manure. Two collections were obtained from the soil under plant benches
in nurseries. Three collections of jP. diffringens came from areas on
the edge of salt marshes that were under salt water during the hurri
canes of 1957 and 1965. Two collections were obtained from rotting
bagasse at sugar m ills. One collection is from the sediment of a
sewage plant.
As can be seen from the above list of the types of situations
from which it was collected, P. diffringens may be found in almost any
area suitable for an earthworm. Of the species of Pheretima in
Louisiana, _P" diffringens is the one most frequently encountered in
areas widely removed from the habitations and activities of man. While
_P. diffringens is distributed all over the state, more effort was
required to collect it from natural habitats in the extreme northern
and western parishes. In those upland pine areas it is largely limited
to floodplain situations.
_P. diffringens was found in all stages of maturity in every month
of the year except October. In October immature specimens were not
collected. The absence of immature specimens in October is probably
because of a decrease in, or absence of, cocoon production during the
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 40
driest portion of the year (Tables 13, 14). This year-round activity,
together with the earthworm's extreme adaptability, have allowed P.
diffringens to colonize the entire state of Louisiana in a period of
somewhat less than 250 years. According to Davis (1955) only small
farms that raised crops that could be planted from seed were prevalent
in Louisiana before 1720. The supposed method of introduction of
Pheretima in Louisiana is in dirt around the roots of imported plant
materials (see p, 96 for a further discussion).
Localities;
Acadia Parish
Mermentau River at Interstate Hwy 10, 26 Sept. 1965, 1-0-6.
Bayou Nezpique at La. Hwy97 , 26 March 1967, 1-0-10.
Bayou Des Cannes at La. Hwy368, 26 March 1967, 0-3-2.
1 .5 mi S Church Point on La. Hwy 3 5 , 26 March 1967, 1-2-6.
Bayou Mallet at La. Hwy1 3 , 31 March 1967, 1-0-4.
Bayou Queue Tortue at La. Hwy1 3 , 31 March 1967, 1-5-4.
Allen Parish
O akdale, 25 March 1965, 0-0-1.
West Bay Game Management Area, 2 O ct. 1965, 0-1-0.
6.3 mi W Kinder on US Hwy 190, 20 Feb. 1966, 0-0-7.
K in d er, 19 S e p t. 1966, 0-3-7.
Calcasieu River at La. Hwy2 6 , 25 March 1967, 1-3-2.
Whiskey Chitto Creek at La. Hwy 26, 25 March 1967, 0-0-3.
Ascension Parish
Bayou Manchac at River Rd.,21 F eb . 1964, 0-0-1.
2 mi W Dutchtown on La. Hwy 74, 12 Dec. 1964, 0-0-15.
3.9 mi S Sorrento on US Hwy 61, 30 Jan. 1965, 0-0-2.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 41
Donaldsonville Fairgrounds, 28 April 1965, 0-0-1.
5.5 mi W Port Vincent on La. Hwy 42, 3 June 1965, 4-9-3.
Assumption Parish
0.8 mi N US Hwy 90 on La. Hwy 662, 7 April 1967, 0-1-3.
Near Lafourche Par. line on La. Hwy 1 , 14 A p r il 1967, 0-0-1.
Avoyelles Parish
1 mi N Bunkie on US Hwy 71, 29 March 1965, 0-0-1.
Across Red River from end of La. Hwy 910, 20 March 1966, 0-3-3.
2.7 mi E Bordelonville on La. Hwy 451, 20 March 1966, 0-0-6.
Marksville Prehistoric Indian State Park, 20 March 1966, 0-0-2.
Simsport on Atchafalaya River, 20 March 1966, 0-0-7.
Beauregard Parish
1 mi S Sugartown on La. Hwy 1 1 3 , 20 F eb . 1966, 0-2-5.
Bayou Anacoco at La. Hwy11 1 , 30 A p ril 1966, 0-2-4.
Sabine River at US Hwy 190, 30 April 1966, 0-0-2.
Longville Lake on La. Hwy 1 1 0 , 1 May 1966, 0-1-8.
Hickory Creek at La. Hwy1 2 , 1 May 1966, 0-4-2.
DeRidder, 8 April 1967, 0-0-1.
Bienville Parish
Friendship, pasture, 27 May 1966, 0-1-8.
Kepler Lake on La. Hwy 153, 12 Aug. 1966, 0-5-3.
2.1 mi SW Castor on La. Hwy 153, 12 Aug. 1966, 0-4-9.
Gibsiand city dump on La. Hwy794, 13 Aug. 1966, 0-1-5.
Bossier Parish
Plain Dealing city dump on La. Hwy 1 5 7 , 23 Aug. 1966, 0-0-1.
Plain Dealing, sawmill, 23 Aug. 1966, 1-2-8.
Cypress Lake, 2.8 mi NW Benton, 23 Aug. 1966, 0-1-2.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 42
Bayou Bodcau at La. Hwy 157, 23 Aug. 1955, 0-1-5.
Caddo Parish
1 .8 a ir mi SE Rodessa on Black Bayou, 11 March 1955, 0-0-1.
0.3 mi S Myrtis on La, Hwy 1, 12 March 1955, 2-2-10.
1.9 mi W Mira on La. Hwy 759, 12 March 1955, 0-0-1.
Boggy Bayou at US Hwy 171, 12 March 1955, 0-0-10.
Spring Ridge, pasture, 10 June 1957, 0-0-1.
Calcasieu Parish
1 mi E Sabine River on US Hwy 90, 23 Aug. 1965, 2-1-3.
0.3 mi W La. Hwy 109 on N ibletts Bluff Rd., 23 Aug. 1955, 0-2-7.
5 mi NE Starks on La. Hwy 12, sawmill, 1 May 1955, 0-5-38.
3 .9 m i W C a rly ss on L a . Hwy 108, 1 May 1955, 2-2-1.
Choupique Bayou at La. Hwy1 0 8 , 1 May 1956, 3-5-2.
5 .3 mi N Cameron Par. line on La. Hwy 27, 21 S e p t. 1956, 0-0-9.
Moss Bluff on La. Hwy 378, 25 March 1957, 0-0-5.
Caldwell Parish
3.2 air mi SSE Hebert on Lafourche Lake, 28 Nov. 1955, 0-0-2.
Kelly on La. Hwy 505, 3 April 1955, 0-1-0.
7 .2 mi S Grayson on US Hwy 1 6 5 , 3 A p r il 1955, 1-1-5.
Columbia Lock and Dam on Ouachita River, 3 April 1955, 0-1-2.
5 .4 mi NE Catahoula Par. line on La. Hwy 559, 3 A p r il 1955, 3-4-1.
Cameron Parish
1 5 .4 mi W Holly Beach on La. Hwy8 2 , 20 S e p t. 1955, 0-0-7.
3 .1 mi W Hackberry on L a. Hwy 3 90, 20 S e p t. 1955, 0-0-52.
Cameron, o ld Henry Home, 20 S e p t. 1955, 0-0-5.
Cameron, o ld Cameron H otel, 21 S e p t. 1955, 0-0-1.
5.5 mi W Grand Chenier on La. Hwy 82, 21 Sept. 1955, 0-0-23.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 43
Grand Chenier, near post office, 21 S e p t. 1966, 0-0-4.
Catahoula Parish
2 .5 mi S Eva, then across Black River, 19 March 1966, 0-2-4.
2 .4 mi ME LaSalle Par. line on La. Hwy 28, 26 March 1966, 0-2-3.
Wallace Lake, 1.7 mi N Jonesville, 26 March Ï966, 1-0-3.
Sicily Island, old sawmill, 26 March 1966, 0-2-10.
0.4 mi N Enterprize on La. Hwy 124, 27 March 1966, 0-9-4.
Claiborne Parish
2.6 mi MW Corney Lalce Dam, 11 Aug. 1966, 0-3-0.
2.4 mi N Haynesville on US H(fy 79, 11 Aug. 1966, 0-0-14.
Black Bayou at La. Hwy 2, 14 Aug. 1966, 0-0-1.
2 .6 mi NW Corney Lake Dam, 19 May1967, 0-0-1.
Concordia Parish
1 mi S Clayton on US Hwy 65, 2 March 1965, 0-0-5.
1 .5 mi S Clayton on US Hwy6 5 , 8 S e p t. 1965, 0-0-2.
2 .8 mi S Clayton on US Hwy6 5 , 8 S e p t. 1965, 0-1-1.
1.4 mi N Old River Locks on La. Hwy 15, 19 March 1966, 1-1-2.
1 .5 mi N Old River Locks on La. Hwy1 5 , 19 March 1966, 0-1-3.
1 mi N Shaw on La. Hwy 15, 19 March 1966, 0-1-2.
3.3 mi S Deer Park on La. Hwy 15, 19 March 1966, 0-0-1.
2.5 mi S Eva on La. Hwy 129, 19 March 1966, 0-0-5.
DeSoto Parish
Clear Lake at La. Hwy 509, 13 March 1966, 0-1-0.
Logansport, La. Hwy 765 a t US Hwy 8 4 , 25 Aug. 1966, 0-1-4.
Stanley, sawmill, 25 Aug. 1966, 2-1-1.
0 .5 mi N Mansfield on US Hwy 17 1 , 25 Aug. 1966, 0-0-4.
1 .9 mi S Frierson on La. Hwy 175, 25 Aug. 1966, 0-0-2.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 44
Clear Lake at La. Hwy 509, 27 Aug. 1966, 0-0-3.
East Baton Rouge Parish
Highland Rd. at Dentation Dr., 11 Nov. 1963, 0-0-2.
Highland Rd. near Highland Park Dr., 11 A p r il 1964, 0-2-2.
River Rd. at Brightside Dr., 31 Oct. 1964, 0-0-4.
Highlandia D r. at Highland Rd., 7 Nov. 1964, 0 - 0 - 5 .
River Rd. at Ben Hur Rd., 6 Dec. 1964, 0-0-9.
0.3 mi N Kleinpeter on old Perkins Rd., 6 Feb. 1965, 0-0-1.
0 .2 mi S Highland Rd. on Ben Hur Rd., 7 F eb. 1965, 0-0-5.
Highlandia Dr. at Highland Rd., 11 F eb . 1965, 0-0-4.
0.3 mi N Kleinpeter on old Perkins Rd., 3 March 1965, 0-1-2.
0.3 mi N Kleinpeter on old Perkins Rd., 21 March 1965, 0-0-2.
N end Profit Island in Miss. River, 3 April 1965, 0-1-6.
S end Profit Island in Miss. River, 3 A p r il 1965, 0-3-3.
Highland Rd. at Dentation Dr., 8 A p r il 1965, 0-0-2.
W. Parker Blvd. at Dodson Ave., 8 April1965, 0-0-23.
1.5 mi SE Pride on Carson Rd., 31 May 1965, 1-0-1.
Comite River at Greenwell Springs Rd., 23 S e p t. 1965, 0-0-11.
5 mi S Pride on La. Hwy 409, 23 Sept. 1965, 0-3-9.
0 .3 mi N Kleinpeter on old Perkins Rd., 2 O ct. 1965, 0-0-2.
Highlandia Dr. at Highland Rd., 6 Nov. 1965, 0-0-1.
East Carroll Parish
Sondheimer, sawmill on US Hwy 6 5 , 10 J u ly 1966, 0-2-7.
2.7 mi S Transylvania, then to Miss. River, 11 July 1966, 1-11-3.
3.1 mi NW La. Hwy 596 on US Hwy 65, 11 July 1966, 0-3-2.
Brunett on L a . Hwy 5 9 6 , 11 J u ly 1966, 4-1-1.
0.6 mi N La. Hwy 2 on Russell Bayou, 11 July 1966, 0-1-3.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 45
East Feliciana Parish
1.7 mi S Blairstovm on La. Hwy 409, 10 Aug. 1964, 0-0-1.
1 .8 mi W Bluff Creek, La. on La. Hwy 959, 23 S e p t. 1965, 0-2-19.
Olive Branch Creek at La. Hwy 67, 16 O ct. 1965, 0-0-4.
0 .5 mi NW Clinton at Pretty Creek, 16 O ct. 1965, 0-5-12.
2.2 mi E Norwood on La. Hwy 422, 16 Oct. 1965, 0-3-14,
Beaver Creek, 2 .4 mi NE Felps on gravel rd., 16 Oct. 1965, 0-0-2.
Evangeline Parish
Chicot State Park, 9 Jan . 1966, 1-2-1.
3 mi S Turkey Creek, La. on La. Hwy13, 9 J an . 1966, 0-0-1.
Boggy Bayou at La. Hwy10 6 , 9 Jan . 1966, 0-2-2.
Bayou Des Cannes at La. Hwy1 3 , 9 Jan . 1966, 2-11-3.
9 mi N Ville Platte on La. Hwy 3042, 9 April 1967, 1-0-1.
Franklin Parish
1 .7 mi S Lorelein along Bayou Macon, 17 A p ril 1966, 0-0-4.
2.5 mi N Crowville on La. Hwy 17, 17 April 1966, 0-0-1.
2 mi N Gilbert on La. Hwy 15, 17 April 1966, 0-0-2.
Winnsboro city dump on Eighth S t., 9 J u ly 1966, 1-5-4.
1 mi E Mangham on La. Hwy 132 at Big Creek, 9 July 1966, 0-9-2.
Grant Parish
3.6 mi N Pollock on US Hwy 165, 8 Aug. 1965, 1-0-3.
0.8 mi N Pollock on US Hwy 165, 8 Aug. 1965, 0-1-1.
1 .3 m i W Bentley on L a. Hwy 8, 15 Aug. 1965, 0-3-3.
0.6 mi NW Rapides Par. line on US Hwy 71, 11 March 1966, 0-0-3,
2 .6 mi S Georgetown on old US Hwy 16 5 , 27 March 1966, 0-0-2.
Iberia Parish
Avery Island, Jungle Garden, Timber Bamboo, 27 A p r il 1965, 0-1-0.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 46
Avery Island, Jungle Garden, near Buddha, 27 A p ril 1965, 0-0-1.
Avery Island, Jungle Garden, Cactus garden, 27 A p r il 1965, 0-1-0.
New Iberia, 28 April 1965, 0-1-1.
4 mi S New I b e r ia on L a. Hwy 8 3 , 28 A p ril 1965, 0-2-5.
1 mi N Weeks on La. Hwy 83, 28 April 1965, 2-1-1.
3 .3 mi S P ig eo n on La. Hwy 997, 30 Oct. 1965, 0-0-18.
Iberville Parish
1 mi NW Sunshine on La. Hwy 337, 18 Oct. 1964, 0-0-11.
2 .7 mi Nlf Sunshine on La. Hwy337, 6 D ec. 1964, 0-0-6.
1 .6 mi NW Sunshine on La. Hwy337, 12 D ec. 1964, 0-0-32.
4 mi W Dutchtown on La. Hwy 74, 12 Dec. 1964, 0-0-1.
1 mi E Rosedale on La. Hwy 76, 31 Dec. 1964, 0-0-2.
Spanish Lake, 4 mi S La. Hwy 427 on La. Hwy 928, 1 A p r il 1965, 0-0-4.
1 mi S Pigeon on La. Hwy 997, 30 O ct. 1965, 0-3-22.
6 .8 mi S Ramah on Bayou M aringouin le v e e , 30 O ct. 1965, 0-0-8.
Jackson Parish
2 mi N Vernon on La. Hwy 14 5 , 28 May 1966, 0-0-2.
0.9 mi E Vernon on gravel rd ., 31 Aug. 1966, 0-0-1.
Chatham Lake on La. Hwy 34, 31 Aug. 1966, 0-0-7.
4 .8 mi E Jonesboro on La. Hwy 4, 31 Aug. 1966, 0-0-13.
Jonesboro, Tenth St., 31 Aug. 1966, 0-0-3.
Jonesboro, Old Gansville Rd., 31 Aug. 1966, 0-0-17.
Jefferson Parish
Grand Isle, near church on Ludwig Lane, 9 A p r il 1967, 0-1-1.
Grand Isle, Chighizola Lane at Melon Lane, 9 A p r il 1967, 0-0-2.
Grand Isle, Chighizola Lane, 9 A p r il 1967, 0-0-4.
1 .4 mi NW Bayou Barataria on La. Hwy2 3 , 23 A p ril 1967, 0-0-1.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 47
0.4 mi S Lafitte on shell rd., 29 April 1967, 0-0-5.
8 mi N Lafitte on La. Hwy 45, 29 April 1967,0-0-2.
Central Ave. at Miss. River, 29 A p r il 1967, 0-0-2.
1.9 mi SW La. Hwy 19 on US Hwy 90, 20 April 1967, 0-0-4.
Jefferson Davis Parish
5 .1 m i N Fenton on US Hwy 1 6 5 , 19 S e p t. 1966, 0-2-7.
Bayou Lacassine at La. Hwy 14, 22 S e p t. 1966, 0-0-7.
Lake Arthur, 0 .7 mi E La. Hwy 26 on Third St., 22 S e p t. 1966, 0-1-5.
Near Lake Arthur Marina, 22 S e p t. 1966, 0-0-3.
Bayou Serpent at La. Hwy 383, 25 March 1967, 0-2-5.
Bayou Lacassine at US Hwy 9 0 , 26 March 1967, 2-4-5.
Lafayette Parish
Milton, old Picard Cemetery on Vermillion River,1 A p r il 1967, 0-2-3.
0 .3 mi S Vermillion River on La. Hwy18 2 , 1 A p r il 1967, 0-0-1.
0.8 mi W US Hwy 167 on Mouton Switch Rd., 2 April 1967, 0-0-7.
Lafourche Parish
6 .2 mi S Golden Meadow on L a. Hwy 1 , 9 A p ril 1967, 0-0-5.
2 mi NW Thibodaux on La. Hwy 1, 9 April 1967, 0-0-2.
LaSalle Parish
Urania, sawmill, 27 March 1966, 0-0-14.
8 .8 m i SW Trout on La. Hwy 8, 2 April 1966, 3-1-1.
0.2 mi S La. Hwy 127 on US Hwy 165, 2 April 1966, 0-2-6.
1.9 mi NE Jena on La. Hwy 772, 20 June 1966, 1-2-4.
Lincoln Parish
Ruston sewage plant, 14 Nov. 1951, 0-0-1.
Ruston, Woodland Park, 5 Feb. 1952, 0-0-7.
Ruston Cattle Sale Barn, 14 March 1952, 0-0-1.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 48
Ruston, Rigdell's Nursery, 21 Aug. 1954, 1-2-2.
1 mi N Ruston on US Hwy 167, 24 S e p t. 1954, 0-0-1.
Ruston, Drigger's Nursery, 16 D ec. 1954, 0-0-3.
Livingston Parish
2 mi S Walker on Gaylord Rd., 29 Oct. 1964, 0-0-9.
12.5 mi N Holden on La. Hwy 441, 11 Nov. 1964, 0-0-1.
2 mi E Livingston on US Hwy 190, 29 Nov. 1964, 0-1-5.
1.5 mi S Springfield on La. Hwy 22, 19 D ec. 1964, 0-0-2.
0.5 mi N Denson on La. Hwy 1039, 19 Dec. 1964, 0-0-7.
2.5 mi E Maurepas on La. Hwy 22, 19 Dec. 1964, 0-0-5.
Port Vincent, old sawmill, 15 March 1965, 0-2-21.
2 mi S Walker on Gaylord Rd., 21 April 1965, 0-0-2.
Denham Springs, Christmas Bend homestead, 24 A p r il 1965, 1-0-3.
Tickfaw River at La. Hwy42, 2 May 1965, 0-0-2.
0.5 mi S Albany on La. Hwy 43, 10 Oct. 1965, 0-0-6.
5 mi NE Port Vincent on La. Hwy 42, 30 Oct. 1965, 0-0-2.
1.2 mi NE Springfield on La. Hwy 42, 13 Nov. 1965, 0-0-4.
4 mi NE Maurepas on La. Hwy 22, 22 Feb. 1966, 0-0-3,
2 mi NW Bayou Barbary on La. Hwy 446, 2 April 1967, 0-0-4.
Madison Parish
7.5 mi N Somerset on US Hwy 65, 9 S e p t. 1965, 0-0-1.
2.1 mi S Waverly, then W to Cow Bayou, 17 April 1966, 0-1-2.
6.6 mi N Waverly on La. Hwy 579, 10 July 1966, 2-1-1.
Tallulah, sawmill on US Hwy 80, 10 J u ly 1966, 1-8-4.
6.5 mi N Mound on Miss. River levee, 10 July 1966, 2-1-0.
Morehouse Parish
Near Arkansas line on Bayou Bartholomew, 31 July 1966, 0-2-11.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 49
2.2 mi S Bastrop on La. Hwy 139, 31 July 1966, 0-0-4.
Chemin-a-haut State Park, 1 Aug. 1966, 0-4-1.
0.2 mi N Log Cabin on La.Hwy 139, 1 Aug. 1966, 0-4-5.
0.6 mi S Arkansas line on La. Hwy 142, 1 Aug. 1966, 0=2-7.
Natchitoches Parish
Natchitoches, La. Hwy 6 a t S ib le y D r., 27 March 1965, 0-0-1.
1 mi S.Red River Par. line on La. Hwy480, 27 March 1965, 0-0-1,
0.5 mi SW Natchitoches on La. Hwy 6, 26 May 1966, 0-0-1.
Black Lake Bayou at La. Hwy 155, 29 Aug. 1966, 0-3-4.
Saline Bayou at La. Hwy126, 29 Aug. 1966, 0-0-1.
Pardee Camp Rd. at Clear Lake, 30 Aug. 1966, 0-0-4.
Orleans Parish
0.7 mi N Intracoastal Canal on La. Hwy 47, 23 April 1967, 0-0-1.
Ouachita Parish
Wilders Creek at La. Hwy 546, 27 May 1965, 0-0-5.
Chenier Lake Dam at La. Hwy 33, 9 Aug. 1966, 1-2-7.
Monroe. 0.2 mi S Forsythe Ave. on 19th St., 9 Aug. 1966, 0-1-3.
Sterlington, La. Hwy 553 at US Hwy 165, 9 Aug. 1966, 0-1-1.
Bayou De Siard at Midway Dam, 10 Aug. 1966, 0-7-12.
Bayou Choudrant at La. Hwy15, 10 Aug. 1966, 0-0-1.
Plaquemines Parish
Near St. Bernard Par. line on La. Hwy 39, 22 April 1967, 0-0-4.
Knobloch Plantation near Dalcour, 22 April 1967, 0-1-1.
3 mi S Pointe a la Hache on La. Hwy 39, 22 April 1967, 0-0-1.
1 mi S Venice on La. Hwy 23, 22 April 1967, 0-0-1.
Pointe Coupee Parish
1 mi N Jarreau on La. Hwy 413, 25 April 1964, 0-0-9.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 50
3.2 mi N Oscar on La. Hwy 1, 14 March 1955, 0-0-1.
8.2 mi SE Morganza on La. Hwy1, 13 J u ly 1965, 0-0-3.
4.4 mi N Morganza on La. Hwy 419, 13 July 1965, 0-6-2.
Rapides Parish
0.3 mi SE Latanier on La. Hwy 457, 7 Nov. 1965, 0-0-12.
4.2 mi NE Gardner on La. Hwy 28, 7 Nov. 1965, 0-0-5.
2 mi E Union H ill on La. Hwy 113, 20 F eb. 1966, 0-5-1.
2.5 mi N Deville on La. Hwy 455, 26 March 1966, 0-0-1.
Red River Parish
0.4 mi E Red River on US Hwy 84, 26 Aug. 1966, 0-1-7.
Grand Bayou at La. Hwy507, 26 Aug. 1966, 0-0-2.
Wilson Lake at La. Hwy 1, 28 Aug. 1966, 0-0-9.
0.1 mi E Red River on US Hwy 84, 29 Aug. 1966, 0-0-1.
0.3 mi W Black Lake Bayou on La. Hwy155, 29 Aug. 1966, 0-0-1.
Richland Parish
1.3 mi NE Rayville on La. Hwy 852, 11 Sept. 1965, 0-1-0.
Clear Lake on La. Hwy 133, 11 Sept. 1965, 1-0-16.
Woolen Lake at La. Hwy 561, 21 June 1966, 0-1-1.
Boeuf River a t La. Hwy 15, 21 June 1966, 0-2-2.
1.6 mi N, 1.5 mi W Delhi, 21 June 1966, 0-0-11.
Sabine Parish
Hodges Gardens, Rose Arbor, 26 March 1965, 0-0-8.
Hodges Gardens, Guest Lodge, 29 March 1965, 0-0-1.
0.4 mi E Pleasant H ill on La. Hwy 174, 1 S e p t. 1966, 1-9-0.
Pleasant H ill, 1 S e p t. 1966, 0-0-5.
3.4 mi S Converse on US Hwy 171, 1 Sept. 1966, 0-1-1.
1.2 mi SE Zwolle on US Hwy 171, 1 S e p t. 1966, 0-0-5.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 51
Saint Bernard Parish
2 .4 mi SE Yscloskey on La. Hwy624, 21 A p r il 1967, 0-2-4.
Bayou St. Malo near Lake Borgne, 21 A p ril 1967, 0-0-1.
2 .9 mi N Delacroix on La. Hwy300, 22 A p r il 1967, 0-0-5.
4.6 mi E Poydras on La. Hwy 46, 22 April 1967, 0-0-3.
2.1 mi S Orleans Par. line on La. Hwy 47, 23 April 1967, 0-0-1.
Chalmette, De la Ronde Park, 23 April 1967, 0-0-2.
Saint Charles Parish
1 mi NE US Hwy 61 on W Bonnet Carre Spillway levee, 13 March 1965,
0-1 -7 .
2 mi NE US Hwy 61 on W Bonnet Carre Spillway levee, 30 O ct. 1965,
0 -0 -3 .
2 m i SW US Hwy 61 on La. Hwy 628, 20 D ec. 1965, 0-0-5.
3 mi NE Norco on E Bonnet Carre Spillway levee,20 D ec. 1965, 0-0-2.
2.8 mi NE Paradis on US Hwy 90, 29 April 1967, 0-0-10.
Saint Helena Parish
1.5 mi S Orangeville on La. Hwy 63, 26 Aug. 1964, 0-0-1.
8 mi NE Orangeville on La. Hwy 37, 26 Aug. 1964, 1-1-2.
2 mi E Oreensburg on La. Hwy 10, 26 Aug. 1964, 0-0-4.
6.8 mi N Orangeville on La. Hwy 448, 15 F eb. 1965, 0-0-1.
5 mi W Chipola on La. Hwy 432, 18 Sept. 1965, 0-0-1.
6.8 mi N Liverpool on La. Hwy43, 22 F eb . 1966, 0-0-2.
Saint James Parish
5.5 mi NW Oramercy on US Hwy 61, 13 Nov. 1965, 0-0-18.
2 mi NW Oramercy on US Hwy 61, 4 Dec. 1965, 0-0-4.
2 mi S Ascension Par. line on La. Hwy 44, 5 Dec. 1965, 0-1-2.
4 mi S Ascension Par. line on La. Hwy 44, 5 D ec. 1965, 0-0-2.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 52
1 mi S Convent on La. Hwy 44, 5 Dec. 1965, 0-0-2.
5 mi S Vacherie on La. Hwy 20, 5 Dec. 1965, 0-0-4.
Saint John The Baptist Parish
2.5 mi N Killona on La. Hwy 18, 30 A p ril 1967, 0-0-6.
Saint Landry Parish
1.5 mi S Cortableau on levee rd., 20 Feb. 1965, 0-0-2.-
Krotz Springs on La. Hwy 105, 24 March 1965, 0-2-1.
1.3 mi N Krotz Springs on La. Hwy 105, 24 March 1965, 0-0-5,
2 mi W M e lv ille on L a. Hwy 10, 24 March 1965, 0-1-6.
5.8 mi N Beggs on La. Hwy 182, 24 March 1965, 0-0-7.
14.2 mi SE Lebeau on US Hwy 71, 6 Nov. 1965, 0-0-1.
6.9 mi SE Lebeau on US Hwy 71, 6 Nov. 1965, 0-0-4.
5 mi S Cortableau on levee rd., 2 Jan. 1966, 0-0-1.
8.2 mi S Cortableau on levee rd., 2 Jan. 1966, 0-0-2.
Bayou Cortableau at US Hwy 190, 15 Oct. 1966, 0-0-1.
4 .4 mi S Cortableau on levee rd., 2 J an . 1967, 0-0-7.
2.2 mi S Cortableau on levee rd., 24 Jan. 1967, 0-0-1.
3.7 mi NW Washington on La. Hwy 745, 24 Jan. 1967, 0-0-3.
Saint Martin Parish
Lake Martin at La. Hwy 353, 20 F eb . 1965, 0-1-27.
4 mi E Cecilia on La. Hwy 678, 31 Oct. 1965, 0-0-3.
3.4 mi E Coteau Holmes on La. Hwy 3083, 31 Oct. 1965, 0-1-6.
5.9 mi N Morgan City on La. Hwy 70, 7 A p r il 1967, 0-2-12.
Saint Mary Parish
4 mi N Charenton on levee rd., 31 Oct. 1965, 0-1-11.
Intracoastal Canal at La. Hwy 319, 2 April 1967, 2-3-1.
2.2 mi W Paterson on US Hwy 90, 2 April 1967, 0-0-8.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 53
Morgan City, cemetery on Young Dr., 2 A p ril 1967, 0-0-1.
S a in t Tammany P a rish
4 .3 mi W M adisonville on La. Hwy 22, 26 D ec. 1964, 0-2-17. .
M adisonville, 26 D ec. 1964, 0-0-1.
Bayou Chinchuba at US Hwy 190, 26 Dec. 1964, 0-0-4.
6.4 mi N Hickory on La.- Hwy 41, 15 Feb. 1965, 0-0-1.
Near Sun on La. Hwy 21, 10 Oct. 1965, 0-0-1.
1.2 mi W Madisonville on La. Hwy 22, 26 Nov. 1965, 0-0-1.
3.4 mi E Pearl River, La. on old US Hwy11, 12 D ec. 1965, 0-0-1.
Near Hickory on La. Hwy41, 12 D ec. 1965, 0-0-1.
Pearl River Canal Lock No. 1, 24 Jan . 1966, 0-0-2.
3.9 mi E Pearl River, La. on old US Hwy 11, 24 Jan. 1966, 0-0-2.
0.2 mi S Hickory on La. Hwy 41, 5 March 1966, 0-0-6.
0.3 mi W Pearl River Bridge on US Hwy 90, 5 March 1966, 0-0-1.
7 mi N Pearl River, La. on US Hwy 11, 5 March 1966, 0-2-9.
Tangipahoa Parish
Tchefuncte River at La. Hwy16, 26 Aug. 1964, 0-1-5.
3.1 mi S Hammond on US Hwy 51, 5 Dec. 1964, 0-0-1.
Ponchatoula Creek at La. Hwy 22, 19 D ec. 1964, 0-0-9.
2.4 mi N Hammond on La. Hwy 443, 23 Dec, 1964, 0-1-3,
4.2 mi N Hammond on La. Hwy 443, 23 Dec. 1964, 0-0-7.
Tangipahoa River at La. Hwy 443, 23 Dec. 1964, 1-8-11.
Natalbany River at La. Hwy 40, 29 Dec. 1964, 0-0-4.
1 mi E Holten on La. Hwy 16, 29 Dec. 1964, 0-0-8.
Tangipahoa River at La. Hwy 22, 26 Nov. 1965, 0-0-1.
Tangipahoa River at La. Hwy445, 26 Nov. 1965, 1-7-5.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 54
Tensas Parish
6.2 mi N Waterproof on US Hwy 65, 2 March 1965, 0-0-7.
6.1 mi S Waterproof on US Hwy 65, 8 Sept. 1965, 0-0-15.
Waterproof on Miss. River, 9 Sept. 1965, 0-0-2.
5.2 mi N Waterproof on US Hwy 65, 9 Sept. 1965, 0-0-12.
M 0 R Landing on Yucatan Lake, 16 April 1966, 0-0-1.
2 mi N Yucatan Lake on M 0 R Landing Rd., 16 April 1966, 0-0-5.
0.2 mi N Saranac on La. Hwy 128, 16 April 1966, 0-0-3.
Terrebonne Parish
21.3 mi S US Hwy 90 on La. Hwy 315, 7 A p ril 1967, 0-4-0.
Crozier, cemetery on La. Hwy 315, 7 A p ril 1967,'“0-0-1.
2.3 mi SE Gibson on US Hwy 90, 8 April 1967, 0-0-2.
Montegut, Bayou Terrebonne at La. Hwy58, 8 A p ril 1967, 0-0-1.
Union Parish
1 mi N Marion on La. Hwy 33, 1 Aug. 1966, 0-3-5.
Marion, old sawmill on La. Hwy 551, 1 Aug. 1966, 0-0-1.
Farmerville,- M iller St., 2 Aug. 1966, 0-0-1.
Ouachita River at La. Hwy 2, 10 Aug. 1966, 0-4-1.
5.6 mi SW Point on La. Hwy 552, 10 Aug. 1966, 0-1-4.
3.6 mi N Bernice on US Hwy 167, 10 Aug. 1966, 0-2-3.
Vermillion Parish
Pecan Island, 5 D ec. 1965, 0-0-1.
Pecan Island, 25 A p r il 1966, 0-0-1.
5 .4 mi W Gueydan on La. Hwy14, 22 S e p t. 1966, 0-0-7.
Indian Bayou, La., 31 March 1967, 0-1-8.
0.2 mi S Kaplan on La. Hwy 35, 31 March 1967, 0-2-3.
0.8 mi W Esther on La. Hwy 82, 31 March 1967, 0-0-3.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 55
Eratli, sugar mill on W Landry St., 1 April 1957, 0-0-4.
Vernon Parish
Sabine River at La. Hwy 8,25 March 1965, 0-0-4.
0.2 mi E Sabine River on La. Hwy 8, 25 March 1965, 0-0-2.
6.2 air mi W Knight on Sabine River, 19 F eb. 1966, 0-1-0.
12.9 mi SW Anacoco on La. Hwy 111, 19 Feb. 1966, 2-4-7.
3.4.mi E Pitkin on La. Hwy 113, 20 Feb. 1966, 0-0-7.
Washington Parish
Bogue Chitto River in Franklinton, 26 Aug. 1964, 0-1-3.
0.3 mi S Sunny Hill on La. Hwy 440, 29 Dec. 1964, 0-1-3.
3.7 mi W Clifton on La. Hwy 38, 29 Dec. 1964, 2-1-3.
1.2 mi E Thomas on La. Hwy 438, 29 Dec. 1964, 0-0-2.
1.1 mi S State Line, La. on La. Hwy 62, 29 Dec. 1964, 0-2-1.
6 mi S Bogalusa on La. Hwy 21, 21 Jan. 1965, 1-3-6.
Bogalusa, St. Rosa Pearl Lane at Pearl River, 31 May1965, 0-0-1.
8 mi W Bogalusa on La. Hwy 439, 25 J u ly 1965, 0-0-1.
Bogalusa, St. Rosa Pearl Lane at Pearl River, 10 Oct. 1965, 0-4-28.
2 mi N Sheridan on La. Hwy 424, 5 March 1966, 0-0-1.
Webster Parish
Crows Creek at US Hwy 80, 13 Aug. 1966, 0-0-2.
1.2 mi E Minden on US Hwy 80, 13 Aug. 1966, 1-5-2.
S ib le y , 0.2 mi E La. Hwy 7 on Road 19, 13 Aug. 1966, 0-3-3.
4.5 mi N Minden on Pine St., 13 Aug. 1966, 0-0-15.
Camp Yatasi on Upper Caney Lake, 22 Aug. 1966, 4-1-2.
0.5 mi S Leton on La. Hwy 159, 22 Aug. 1966, 1-2-5.
Springhill, sawmill on La. Hwy 7, 22 Aug. 1966, 0-8-8.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 56
West Baton Rouge Parish
6 mi E Rosedale on La. Hwy 76, 31 Dec. 1964, 0-0-1.
5 mi E Rosedale on La. Hwy 76, 31 Dec. 1964, 0-0-1.
1 mi N Addis on River Rd., 23 Oct. 1965, 0-0-3.
4.2 mi W Brusly on La. Hwy 989-1, 23 Oct. 1965, 0-0-4.
7 mi W Port Allen on La. Hwy 76, then 2,8 mi S, 23 O ct. 1965, 0-0-5.
West Carroll Parish
4.5 mi N Oak Grove on La. Hwy 17, 12 July 1966, 0-0-3.
Boeuf River at La. Hwy 2, 12 July 1966, 0-1-3.
2.3 mi E Kilbourne on La. Hwy 585, then 1.1 mi S to Bayou Macon,
29 July 1966, 0-1-4.
0.6 mi S Floyd on La. Hwy 577, 29 July 1966, 1-0-1.
Oak Grove, Johnson St. at E Noble S t.,18 May 1967, 0-0-1.
West Feliciana Parish
Alexander Creek at US Hwy 61, 1 Nov. 1964, 0-0-4.
3.2 air mi SW Baines on Bayou Sara, 20 J a n . 1965, 7-1-21.
2.1 mi SW Plettenberg on dirt rd., 20 Jan. 1965, 0-0-1.
N end Fancy Point Towhead Island in Miss. River, 3 April 1965, 1-0-1.
S end Fancy Point Towhead Island in Miss. River, 3 April 1965, 0-0-5.
2 a ir mi SE Innis on Raccourci Island, 14 J u ly 1965, 1-1-0.
Tunica, sawmill on La. Hwy 66, 30 July 1965, 2-21-21.
Tunica, on Tunica Bayou, 30 J u ly 1965, 0-1-3.
1.5 mi N Laurel Hill on US Hwy 61, 12 Sept. 1965, 0-0-7.
2 mi W L ittle Bayou Sara on La. Hwy 66, 26 March 1966, 0-0-3.
Winn Parish
Saline Bayou a t La. Hwy 126, 29 Aug. 1966, 0-0-2.
Saline Bayou at US Hwy 84, 30 Aug. 1966, 0-0-4.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 57
0.8 mi S Winnfield on US Hwy 167, 30 Aug. 1966, 0-0-3,
Winnfleld, sawmill on Front St., 30 Aug, 1966, 0-6-14,
Saline Bayou at La. Hwy156, 30 Aug. 1966, 0-1-0.
3.4 mi N Dodson on US Hwy 167, 31 Aug. 1966, 0-1-1.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 58
Pheretima hawayana (Rosa, 1891)
1891 Pcrichaeta hawayana Rosa, Annal.K.K. Naturhist. Hofmuseim Wien
6:396-397 F ig s . 7 and 9.
1905 Pcrichaeta burmudensis: Harper, Biol. Bull. 10:18.
1965 Pheretima hawayana: Gates, Proc. Biol. Soc. Washington 78:1-16.
Diagnosis : Sexthecal, spermathecal pores minute and superficial
in 5/6-7/8, about 1/3 C apart. Female pore single, midventral on xiv.
Male pores minute and superficial on transversely elliptical disc
shaped porophores in the setal circle of xviii, about 1/3 C apart.
Genital markings all postsetal, small discs, paired and slightly median
to the spermathecal pores, closer to the intersegmental furrows than to
the setal line in some of vi-ix, in xviii median to male porophores and
in clusters of 1-11, occasionally absent. Clitellum annular
13/14-16/17, occasionally failing to reach 13/14 and (or) 16/17, dorsal
pores and intersegmental furrows obliterated but setae usually present
ventrally in xvi, occasionally also in one or both of xiv-xv. Setae,
size as well as intersetal intervals increasing through ii-vi especially
ventrally, much smaller in vii though somewhat larger than in v iii and
posteriorly, 17-21/iii, 36-40/viii, 44-49/xii, 0-16/xvi, 48-56/xx,
47-56/xxx-xci, vi/4-8, vii/10-15, xviii/10-16. First dorsal pore usually
at 10/11. Prostomium epilobic with open tongue. Length, 56-200 mm.
Diameter, 3-6 mm. Segments, 70-101, but usually 91-98.
Septa 5/6-7/8 and 10/11-13/14 somewhat thickened, 8/9-9/10 absent.
Intestine beginning in xv. Intestinal caeca simple but with several
short lobes on ventral and (or) dorsal sides, arising in xxvii and
extending forward to about xxiv. Typhlosole rudimentary or even
lacking anteriorly, slightly widened but interrupted and with diagonal
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 59
ridges in some part of xli-lxiv. Hearts of x esophageal, those of
xi-xiii latero-esophageal. Testis sacs unpaired and ventral in x and
xi. Seminal vesicles in xi and xii. Prostates through some or all of
xvi-xxiv with ducts 3-6 mm long, often with a considerable portion
rather thickly muscular. Spermathecae in vi, vii, and viii.
Spermathecal duct slender and usually shorter than large globular
ampulla. Diverticulum arising from anterior face of duct close to
parietes, shorter than combined length of duct and ampulla, but
usually longer than duct. Seminal chamber not sharply demarcated from
stalk of diverticulum, but enlarged. Genital marking glands stalked,
coelomic and composite.
The above diagnosis is a composite taken from the original
description of Rosa (1891), a manuscript by Gates and a paper on the
variation in _P. hawayana (Gates, 1965b).
Discussion; The 678 specimens of P. hawayana collected in
Louisiana conform well with the above diagnosis except for the number
of setae on various segments, 15-27/vi (no record), 31-41/viii
(36-40/viii), 0-18/xvi (0-16/xvi), 47-57/xx (48-56/xx), v/4-7 (no
record), vi/4-9 (vi/4-8), vii/9-18 (vii/10-15), viii/10-18 (no record),
xviii/8-17 (xviii/10-16).
The clitellate, nonautotomized specimens in this study ranged
from 60-179 mm long and from 2.5-6 mm in diameter, the number of
segments ranged from 77-97. The average length of 100 of these
specimens was 111 mm, the number of segments averaged 90.
This species is extremely active and when disturbed will jump and
thrash about in a violent manner. In this study, 85 of the 678
specimens autotomized when they were•collected. The level of autotomy
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 60
was always In the intersegmental furrow and occurred from 40/41-89/90.
As Gates (1965b) has noted, the recognition of regenerated specimens
at the more posterior levels is more difficult than at the more anterior
levels. In recently preserved specimens the regenerated portion is of
a distinctly lighter color than the older segments. The regenerated
segments are also smaller in diameter and shorter in length than the
older segments. The greatest number of regenerated segments in this
study was 19 produced at 51/52 in a worm of 70 segments.
Records of the genital markings (GMs) were kept on 436 of the 678
specimens collected in Louisiana. In addition to the 409 specimens in
Tables 6-9, 27 specimens lacked GMs entirely. .As can be seen in Tables
6 and 7, 174 specimens lacked preclitellar GMs. The most common
pattern of preclitellar GMs was a pair in vii, found in 170 specimens
(Figure 14) . The total number of postclitellar GMs on a specimen can
be seen in Table 8. The most common pattern of postclitellar GMs was
one on each side of xviii, found in 13^ specimens (Table 9). The pre-
and postclitellar GMs may be asymmetrical, present on one side and
lacking on the other.
The size of the preclitellar GMs ranged from 0.12-0.2 mm in
diameter, while the postclitellar GMs ranged from 0.2-0.36 mm in
diameter. The size of these GMs has never been recorded in the
literature.
D istribution; _P. hawayana has been reported from Burma, Ceylon,
China, Borneo, Java, Samoa, Tahiti, F iji and Hawaiian Islands, Central
and South America, Egypt, St. Helena (Gates, MS); and Taiwan (Tsai,
1964) . Its original home may be in China where it is widely
distributed (Gates, 1963).
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 61
P. hawayana has been reported from greenhouses in Illin ois (Smith,
1915); Massachusetts (Gates, 1942a); Illin o is,'Tennessee, Oregon (Gates,
1954); Maine, New Jersey (Gates, 1963). It was also reported from a
bait dealer in Michigan (Gates, 1954) and may have been found in an
outdoor habitat in Michigan (Stinauer, 1951), but the identification
has never been confirmed. In outdoor habitats it has been reported
from Louisiana, M ississippi (Gates, 1937C) ; Louisiana, Alabama, Florida,
California (Gates, 1954); and again from California (Gates, 1967).
Although the species has been introduced into the northern part
of the United States via greenhouses on numerous occasions, it
apparently has been able to establish itself outdoors only in the
warmer portion of the United States (Alabama, California, Florida,
Louisiana, M ississippi).
In Louisiana, _P. hawayana was found in 36 parishes at 128 sites
(Map 4). At 60 sites this species was found on the floodplains of
bayous, creeks, and rivers or the margins of lakes and swamps. Thirty-
one of the sites are in yards or similar sites, 10 sites are in woods,
6 of the sites are in dumps, and 4 sites are in botanical gardens.
Barnyards, cultivated farm land, and grassland were the sites in 3
instances each, _P. hawayana was found at 2 sites each in nurseries,
sawmills, and in rotting bagasse at sugar m ills. It was also found at
one site in the dried out sludge of a sewage plant and at one site in
a cemetery.
All of the collections of P. hawayana north of the northern
boundary of the Florida Parishes are from yards, dumps, barnyards or
nurseries where the worms were introduced by the activities of man. In
northern Louisiana, this species has not dispersed away from these areas
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 62
as it. has done in the southern portion of the state. With the passage
of time, this species can be expected to extend its range in the
northern part of Louisiana.
Clitellate, active specimens of this species were collected in
every month of the year in Louisiana, while immature specimens were
found only in February, March, April, November, and December in outdoor
habitats. Immatures were also secured at a nursery in August. In a
natural habitat cocoons apparently are produced from about October
until April, the species possessing the ability to produce cocoons at
any time when conditions are favorable. In an incomplete study of the
life history of P. hawayana, Stinauer (1951) found that each cocoon
contains one embryo but failed to secure any other data on the life
history of this species.
Localities :
Ascension Parish
2 mi W Dutchtown on La. Hwy 74, 12 Dec, 1964, 0-0-1.
3.9 mi S Sorrento on US Hwy 61, 30 Jan, 1965, 0-0-2.
Donaldsonville, Charles St. at Lafourche St., 28 A p ril 1965, 0-0-1.
Donaldsonville Fairgrounds, 28 A p r il 1965, 0-0-2.
E bank Miss. River at La. Hwy 22, 28 April 1965, 1-2-1.
Assumption Parish
Bellerose, on Bayou Lafourche, 14 April 1957, 0-0-1.
5.5 mi N Pierre Pass on La. Hwy 70, 14 April 1967, 0-1-5.
Near Lafourche Par. line on La. Hwy 1, 14 April 1967, 0-1-2,
4 mi S Napoleonville on La. Hwy 1, 14 April 1967, 1-9-12.
Avoyelles Parish
1 mi N Bunkie on US Hwy 71, 29 March 1965, 0-0-1.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 63
Marksville Prehistoric Indian State Park, 20 March 1966, 0-1-2.
Bienville Parish
Arcadia, Fourth St. at Thornhill St., 12 Aug. 1966, 0-1-5.
1.4 mi S Arcadia on La. Hwy 9, 12 Aug. 1966, 0-1-4.
Calcasieu Parish
0.3 mi W La. Hwy 109 on Nibletts Bluff Rd., 23 Aug. 1965, 0-0-3.
Moss B lu ff on La. Hwy 378, 25 March 1967, 0-0-4.
Cameron Parish
Cameron, old Cameron H otel, 21 S e p t. 1966, 0-0-9.
Grand Chenier, near post office, 21 Sept. 1966, 0-0-5.
Claiborne Parish
1.7 mi NE Homer on L a. Hwy 9, 11 Aug. 1966, 0-1-5.
East Baton Rouge Parish
Highland Rd. at Dentation Dr., 11 Nov. 1963, 0-0-3.
Highland Rd. near Highland Park Dr., 11 A p r il 1964, 0-2-2.
McDonald Ave. near Highland Rd., 15 A p ril 1964, 13-6-9.
L. S. U. Campus, beside Dodson Pond, 3 May 1964, 0-5-1.
L. S. U. Campus, beside Dodson Pond, 17 Sept. 1964, 0-0-1.
W Lakeshore Dr. near Corporation Canal, 3 Nov. 1964, 0-0-3.
Stanford Ave. at Yale Ave.,28 Nov. 1964, 0-0-5.
Ben Hur Rd. at Nicholson Dr., 6 D ec. 1964, 0-0-1.
0.3 mi N Kleinpeter on old Perkins Rd., 6 Feb. 1965, 1-0-1.
0.2 mi S Highland Rd. on Ben Hur Rd., 7 Feb. 1965, 0-3-6.
0.2 mi N Highland Rd. on Nelson Dr., 7 F eb . 1965, 0-1-7.
McDonald Ave. near Highland Rd., 9 Feb. 1965, 0-0-2.
Highlandia Dr. at Highland Rd., 11 Feb. 1965, 0-1-0.
Houston St. at Gail Dr., 26 Feb. 1965, 0-0-1.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 64
0.3 mi N Kleinpeter on old Perkins Rd., 3 March 1965, 0-0-1.
0.3 mi N Kleinpeter on old Perkins Rd., 21 March 1965, 0-1-0.
Gardens near State Capital, 1 April 1965, 0-0-1.
McDonald Ave. near Highland Rd., 7 April 1965, 0-2-8.
Highland Rd. at Dentation Dr., 8 April 1965, 0-1-7.
W Parker Blvd. at Dodson Ave., 8 A p r il 1965, 0-1-8.
Perkins Rd. near Lee Dr., 11 April 1965, 0-0-1.
L. S. U. Campus, Audubon Hall, 29 May 1965, 0-3-3.
Comite River at Greenwell Springs Rd., 23 Sept. 1965, 0-2-4.
0.3 mi N Kleinpeter on old Perkins Rd., 2 Oct. 1965, 0-2-3.
L. S. U. Campus, near golf course, 27 Nov. 1965, 0-0-1.
East Feliciana Parish
0.5 mi NW Clinton at Pretty Creek, 16 Oct. 1965, 0-2-6.
Iberia Parish
Avery Island, Jungle Garden, Timber bamboo, 27 A p ril 1965, 0-6-5.
Avery Island, Jungle Garden, Sunken garden, 27 A p ril 1965, 0-1-0.
Avery Island, Jungle Garden, Camélia garden, 27 A p r il 1965, 0-5-2.
Avery Island, Jungle Garden, Cactus garden, 27 A p ril 1965, 0-1-0.
New Iberia sewage plant on La. Hwy 83, 28 A p r il 1965, 0-1-2.
Jeanerette, E Main St., 5 March 1966, 1-0-6.
Iberville Parish
Plaquemine, Desorby St. at Calvin S t.,17 May 1964, 0-0-5.
2.7 mi NW Sunshine on La. Hwy 337, 6 Dec. 1964, 0-0-2,
1.6 mi NW Sunshine on La. Hwy 337, 12 Dec. 1964, 0-0-8.
Spanish Lake, 4 mi S L a. Hwy 427 on La. Hwy 928, 1 A p r il 1965, 0-0-1.
Jefferson Parish
Grand Isle, near church on Ludwig Lane, 9 A p r il 1967, 0-3-7.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 65
8 mi N Lafitte on La. Hwy 45, 29 April 1967, 0-3-4.
1.9 mi SW La. Hwy 18 on US Hwy 90, 29 April 1967, 0-1-6.
Jefferson Davis Parish
2.6*mi W Jennings on US Hwy 90, 26 March 1967, 0-0-2.
Lafayette Parish
Broussard, R. U. Bernard Park, 1 A p ril 1967, 0-3-10.
0.3 mi S Vermillion River on La. Hwy 182, 1 A p r il 1967, 0-1-1.
Lafourche Parish
8.5 mi W Larose on La. Hwy 24, 8 April 1967, 0-0-9.
0.7 mi S Lockport on La. Hwy 1, 9 A p ril 1967, 0-0-3.
Thibodaux, La. Hwy1 at Cherokee Ave., 9 A p r il 1967, 0-0-1.
2 mi NW Thibodaux on La. Hwy 1, 9 A p ril 1967, 0-1-3.
LaSalle Parish
1.3 mi W Jena on US Hwy 84, 20 June 1966, 0-0-5.
Lincoln Parish
Ruston, Rigdell's Nursery, 21 Aug. 1954, 1-0-1.
Ruston, Drigger's Nursery, 16 D ec. 1954, 0-0-3.
Livingston Parish
0.5 mi N Denson on La. Hwy 1039, 19 D ec. 1964, 2-6-9.
Port Vincent, old sawmill, 15 March 1965, 0-2-12.
1.2 mi NE Springfield on La. Hwy 42, 13 Nov. 1965, 0-0-1.
Orleans Parish
Canal Blvd. at Chappelle St., 26 June 1965, 0-2-14.
Audubon Park, 26 June 1965, 0-1-4.
Chartres St. at Ursulines Ave., 28 April 1967, 0-1-1.
City Park, Palm Dr. af Roosevelt Mall,28 A p r il 1967, 0-0-6.
City Park, Stadium Dr. at Roosevelt Mall,28 A p ril 1967, 0-1-2.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 65
L ittle Woods on Hayne Dr., 28 April 1967, 0-0-3.
Plaquemines Parish
Knobloch Plantation near Dalcour, 22 April 1967, 0-2-5.
3 mi S Pointe a la Hache on La. Hwy 39, 22 April 1967, 0-4-9.
1 mi S Venice on La. Hwy 23, 22 April 1967, 0-3-2.
5.8 mi N Boothville on La. Hwy 325, 22 April 1967, 0-0-1.
Belle Chase on La. Hwy 23, 23 A p ril 1967, 0-3-4.
Pointe Coupee Parish
1 mi N Jarreau on La. Hwy 413, 25 April 1964, 0-0-10.
Batchelor, Major's Landing, 14 July 1965, 0-0-2.
Rapides Parish
Pineville, barnyard, 27 Nov. 1960, 0-2-1.
Bayou Boeuf in Cheneyville, 25 Nov. 1965, 1-1-11.
Red River Parish
0.1 mi E Red River on US Hwy 84, 29 Aug. 1966, 0-0-3.
Saint Bernard Parish
2.4 mi SE Yscloskey on La. Hwy 624, 21 April 1967, 0-0-3.
2.9 mi N Delacroix on La. Hwy 300, 22 A p r il 1967, 0-0-2.
2.1 mi S Orleans Par. line on La. Hwy 47, 23 A p r il 1967, 0-1-6.
Chalmette, De la Ronde Park, 23 A p r il 1967, 0-4-3.
Chalmette, De la Ronde House, 23 April 1967, 0-0-1.
Saint Charles Parish
1 mi NE US Hwy 61 on W Bonnet Carre Spillway le v e e , 13 March 1965,
0 -3-13.
2 mi NE US Hwy 61 on W Bonnet Carre Spillway levee, 30 O ct. 1965,
0- 0- 6 .
2 mi SW US Hwy 61 on La. Hwy 628, 20 D ec, 1965, 0-0-9.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 67
3 mi NE Norco on E Bonnet Carre Spillway levee, 20 D ec, 1965, 0-0-4.
2 mi NE Norco on E Bonnet Carre Spillway levee, 20 D ec. 1965, 0-0-3.
Luling on La. Hwy 52, 29 A p r il 1967, 0-0-1.
Saint James Parish
5.5 mi NW Gramercy on US Hwy 61, 13 Nov. 1965, 0-0-3.
2 mi NW Gramercy on US Hwy 61, 4 Dec, 1965, 0-0-6,
4 mi S Ascension Par. line on La. Hwy 44, 5 Dec. 1965, 0-0-5.
1 mi S Convent on La. Hwy 44, 5 Dec. 1965, 0-0-7.
5 mi S Vacherie on La. Hwy 20, 5 Dec. 1965, 0-0-4.
Saint Jolm The Baptist Parish
3.4 mi N Laplace on US Hwy 51, 13 Nov. 1965, 0-0-1.
4.2 mi N Laplace on US Hwy 51, 13 Nov. 1965, 0-0-7.
7.2 mi N Laplace on US Hwy 51, 13 Nov. 1965, 0-0-1.
2.3 mi S Laplace on La. Hwy 628, 30 April 1967, 0-1-1.
2.5 mi N Killona on La. Hwy 18, 30 April 1967, 0-0-3.
Saint Landry Parish
Krotz Springs on La. Hwy 105, 24 March 1965, 0-1-4.
1.3 mi N Krotz Springs on La. Hwy 105, 24 March 1965, 0-0-10.
2 mi W Melville on La. Hwy 10, 24 March 1965, 0-1-15.
14.2 mi BE Lebeau on US Hwy 71, 6 Nov. 1965, 0-0-1.
Bayou Cortableau at US Hwy190, 15 O ct. 1966, 0-1-3.
Saint Martin Parish
Lake Martin at La. Hwy 353, 20 F eb. 1965, 0-0-4.
6.7 mi SE Henderson on Dupuis Rd., 31 Oct. 1965, 0-0-1.
5.9 mi N Morgan City on La. Hwy 70, 7 April 1967, 0-0-1.
Saint Mary Parish
1 mi E Franklin on US Hwy 90, 28 April 1965, 0-1-2.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 68
4 mi N Charenton on levee rd., 31 Oct. 1965, 0-0-5.
S ain t Tammany P a rish
1.2 mi S Folsom on La. Hwy 25, 23 Dec. 1964, 0-0-1.
3.4 mi E Pearl River, La. on old US Hwy 11, 12 Dec. 1965, 0-0-1.
Talisheek Creek at La. Hwy 41, 22 Dec. 1966, 0-0-1.
Tangipahoa Parish
3.1 mi S Hammond on US Hwy 51, 5 Dec. 1964, 0-1-0.
Ponchatoula Creek at La. Hwy 22, 19 Dec. 1964, 0-1-7.
1 mi E H olten on La. Hwy 16, 29 D ec. 1964, 0-0-1.
Terrebonne Parish
Gray on La. Hwy 24, 22 Aug. 1966, 0-0-4.
Crozier, cemetery on La. Hwy 315, 7 April 1967, 0-3-3.
2 .3 mi SE G ibson on US Hwy 90, 8 A p r il 1967, 0-0-2.
Montegut, Bayou Terrebonne at La. Hwy 58, 8 April 1967, 0-7-16.
Vermillion Parish
0.2 mi S Kaplan on La. Hwy 35, 31 March 1967, 0-0-5.
0.8 mi W Esther on La. Hwy 82, 31 March 1967, 0-1-1.
Erath, sugar mill on W Landry St., 1 April 1967, 0-0-2.
Washington Parish
Bogalusa, St. Rosa Pearl Lane at Pearl River, 10 Oct. 1965, 0-1-0.
West Baton Rouge Parish
6 mi E Rosedale on La. Hwy76, 31 D ec. 1964, 0-3-5.
5 mi E Rosedale on La. Hwy 76, 31 Dec. 1964, 0-2-1.
1 mi N Addis on River Rd., 23 Oct. 1965, 0-0-2.
4.2 mi W Brusly on La. Hwy 989-1, 23 Oct. 1965, 0-0-1.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 69
Pheretima hilgendorfi (Michaelsen, 1892)
1892 Perichaeta hilgendorfi Michaelsen, (part) Arch.
Naturgeschichte 58:235-237 Fig. 15 (Excluding specimens
without midventral presetal patches of genital markings.
See Gates, 1958)
1892 Perichaeta rokugo Beddard, Zool. Jahrb. Syst. 6:756-759 Fig.
1-7.
1954 Pheretima hilgendorfi: Gates, Bull. Mus. Comp. Zool. 111:230-
234.
Diagnosis : Quadrithecal, spermathecal pores small transverse
slits with slightly whitened margins in 6/7-7/8 nearly 1/2 C apart.
Female pore single, midventral on xiv. Male pores usually absent;
when present minute and located on the lateral margin of an eversible
papilla, contained within a slit-like parietal invagination on an
elevated porophore, usually in the setal circle of xviii about 1/3 C
apart. Genital markings small raised circular tubercules (0.16 mm in
diameter), each with a minute central aperture, located inside unpaired,
midventral, presetal depressions (0.6-1.2 mm in diameter) in some of
vi-xi, occasionally in xvii-xviii, very infrequently in xix-xxi.
Clitellum annular 13/14-16/17, dorsal pores, setae and intersegmental
furrows obliterated. Setae slightly larger ventrally in the pre-
clitellar region, but with intersetal distances nearly equal, 17-30/ii,
18-41/iii, 35-43/iv, 40-52/vi, 46-62/viii, 48-66/xii, 50-62/xx,
46-60/xxx, vi/19-30, vii/20-31, viii/23-28, xviii/17-20. First dorsal
pore usually at 12/13. Prostomium epilobic with open tongue. Length,
85-215 mm. Diameter, 5-9 mm. Segments, 92-116.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 70
Septa 5/6-Ô/7 and 11/12-13/14 slightly thickened, 8/9-9/10 absent.
Intestine beginning in xv. Intestinal caeca manicate, arising in xxvii
and extending forward to about xxiii, with 6-10 secondary caeca, the
dorsal-most the longest. Typhlosole low and lamelliform, arising in
xxvii and ending in region of Ixxi-lxxix. Hearts of x esophageal when
present but usually lacking, those of xi-xiii latero-esophageal. Testis
sacs unpaired and ventral in x and xi. Seminal vesicles in xi and xii.
Prostates present in xvi-xx only when male terminalia are present,
with muscular U-shaped duct 5-6 mm long, thinning at both prostate and
papilla of male porophore. In absence of male terminalia, the vasa
deferentia end blindly in xvii-xxx without passing into parietes.
Spermathecae in vii and v iii. Spermathecal duct with thick muscular
wall, shorter than ampulla. Ampulla with thin wall and numerous
wrinkles, dorsoventrally flattened. Diverticulum arising from median
face of duct, longer than combined lengths of duct and ampulla, stalk
with muscular wall. Seminal chamber with thinner wall, slightly larger
in diameter than stalk. Genital marking glands coelomic, with long
stalk, one for each tubercule in the midventral, presetal patches.
Since the original description of Michaelsen (1892) in c lu d e s
specimens of at least one other species, the above diagnosis is a
composite taken from later additions to the morphology of the species
(H a ta i, 1930; G a tes, 1954, 1958; Kobayashi, 1937, 1938; Yamaguchi,
1930a, 1962). P. hilgendorfi is characterized by the midventral,
presetal patches of genital markings and identifications of worms
without these patches is unacceptable (Gates, 1958).
Discussion; The 54 specimens of _P. hilgendorfi collected in
Louisiana conform to the-above diagnosis with the exception of a
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 71
greater total length, 218 mm (Literature; 215 mm); in the number of
setae on various segments, 38-45/iv (35-43/iv), 45-55/vi (40-52/vi),
52-62/viii (46-62/viii), 53-68/xx (50-62/xx), vi/21-28 (vi/19-30),
vii/22-27 (vii/20-31), viii/23-30 (viii/23-28); and in a more posterior
extent of the typhlosole back to Ixxxii (Ixxix).
The clitellate, apparently nonautotomized specimens in this study
ranged from 81-218 mm long and from 5-9 mm in diameter, the number of
segments ranged from 97-114. The average length of 22 of these speci
mens was 114 mm, the average number of segments was 109.
This species is extremely activée and crawls rapidly when disturbed
but does not jump and thrash about as does _P. diffringens. It may
autotomize a posterior portion of its body when disturbed as did 23 of
the 54 specimens collected in this study. The level of autotomy
ranged from 50/51-99/100 and was always in the intersegmental furrow.
The recognition of regenerates in _P. hilgendorfi is difficult as
segments are apparently produced in a slow, one-at-a-time manner
similar to that in P. agrestis. In addition, the regenerated segments
are of the same length as the older segments, making recognition by the
usual means impossible. A growth zone of 2 segments, as indicated by
the dorsal pores and intersegmental furrows but without setae, was
recognized in regenerates of 84 and 93 segments. A similar growth zone
was found in adults of 97, 104, 107, 109, 110, 111, 112, 113, and 114
segments. Unless every specimen in this study was a posterior amputee
in the past, segments are added in mature adults in the same manner as
in regeneration.
The following list shows the number of specimens (in parentheses)
that possess the characteristic midventral, presetal patches of genital
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 72
markings on the segments indicated by Roman numerals: viii(20); i%(3);
viii and ix(28); viii and xviii(l); viii, ix and xvii(l); viii, ix and
x v i i i ( I ) .
The following list shows the number of tubercules (in parentheses)
within the midventral, presetal patches on the segments indicated by
Roman numerals: viii(l-13), ix(ll-28), xvii(17), xviii(30-33).
The specimen with patches in v iii and xviii has a well developed
male porophore on the left side of xviii and is from the Shreveport
site in Caddo Parish. Two aclitellate specimens from the Livingston
Parish site have very rudimentary male pores developing on the right
side. In one specimen the genital marking patches are in v iii, ix and
xvii; with the male pore in xvii. The other specimen possesses the
genital markings in v iii, ix and xviii; with the male pore in xviii.
From the above it would appear that there is a correlation between the
genital markings in xvii-xviii and the possession of male pores in
these Louisiana specimens. This observation has not been reported by
other workers, and it is impossible to determine from the literature if
it holds true in other areas.
Yamaguchi (1931) has recorded the variation in the location of the
male pores in P. hilgendorfi from Japan and found 10 per cent of the
specimens with male pores, which occurred in xvi, xvii, or xviii.
Specimens with male pores were also found in Korea (Kobayashi, 1937,
1938), but they have never been described adequately.
In the specimen with well developed male terminalia on the left
side, the male porophore is an elevated mound about 1.8 mm in diameter
located in the setal circle of xviii, its posterior border obliterating
p a rt o f 18/19. The central transverse aperture of the porophore leads
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 73
to a shallow parietal Invagination and is surrounded by numerous wrinkles
of the epidermis. The minute male pore is located on the lateral
margin of the papilla, which forms the roof of the shallow parietal
invagination. This papilla is apparently eversible through the opening
of the male porophore, as evidenced by its spongy internal nature, but
in the specimen at hand it is contained within the shallow parietal
invagination. There are 9 setae between the male porophore and the
nerve cord on xviii (Figures 16 and 17).
The prostate is present only on the left side extending from
16/17-20/21, divided into 3 main lobes, with several smaller lobes
composing each main lobe. The vas deferens passes into the prostatic
duct close to the connection of the latter with the prostate
(Figure 18). The muscular prostatic duct is about 5 mm long and at its
junction with the papilla of the male pore, it loses its muscular nature
and fuses with the tissue of the papilla (Figure 16).
The stalked genital marking glands, one for each tubercule in the
patch, extend well into the coelom as can be seen in Figure 18. The
internal anatomy of these glands was described by Beddard (1892a) and
Ohfuchi (1939).
Distribution; Pheretima hilgendorfi is widely distributed in
Japan, its presumed original home, having been reported from all four
main islands (Michaelsen, 1892; Beddard, 1892a; H a ta i, 1930; Yamaguchi,
1930a; Kobayashi, 1937). It has also been reported from Korea
(Kobayashi, 1938) .
In the United States, _P. hilgendorfi has been reported from
New York, Michigan, Virginia (Gates, 1954); Louisiana (Gates, 1958);
and Connecticut (Gates, 1966) in outdoor sites.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 74
111 the present study, P. hilgendorfi was collected in 7 parishes
at 8 sites, scattered in the northern and southern portions of the
state (Map 5). Two of the sites are the margin of a lake and the
floodplain of a bayou, where the worms may have been carried by
fishermen. Three of the sites are in flowerbeds or similar sites in
yards, where the worms may have been introduced with exotic plants.
This species was also found in a dump, a cemetery, and a sawmill, all
places where the activities of man may well be responsible for the
presence of the worms.
From the above it can be seen that the distribution of _P.
hilgendorfi in Louisiana is limited to areas influenced by the activities
of man and that it has not as yet dispersed over the state as _P.
diffringens has done.
This lack of dispersal may be due in part to the annual life
cycle described for it in Japan and Korea, in which the adult worms die
in the fall and the species overwinters in the cocoon (Kobayashi,
1937). Stinauer (1951) found that cocoons of P. hilgendorfi required
an incubation period of from 244-264 days in the laboratory in Michigan,
while Kobayashi (1937) found that the period from oviposition to
hatching required 196 days in the field in Korea.
In Louisiana, as well as the remainder of the United States, this
species appears to have the same annual life cycle as that described
in Japan and Korea, The largest specimen of P. hilgendorfi in this
study (218 mm by 8 mm) was collected on 28 October from the Livingston
Parish site and was kept in the laboratory until 6 December, when it
died. C litellate specimens were collected in Louisiana from April to
October, while aclitellate specimens were secured in the period of
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 75
March to June. Other United States records are from the period of
May to September.
L o c a lit ie s ;
Ascension Parish
5.5 mi W Port Vincent on La. Hwy 42, 3 June 1965, 0-0-9.
Caddo Parish
Shreveport, 0.1 mi S Cross Bayou on N Common St., 25 Aug. 1966, 0-0-2,
1 mi W US Hwy 171 on La. Hwy 511, 10 June 1967, 0-1-8.
East Carroll Parish
3.1 mi NW La. Hwy 596 on US Hwy 65, 11 July 1966, 0-0-3.
LaSalle Parish
1.3 mi W Jena on US Hwy 84, 20 June 1966, 0-0-1.
Livingston Parish
2 mi S Walker on Gaylord Rd., 29 Oct. 1964, 0-0-1.
2 mi S Walker on Gaylord Rd., 21 April 1965, 0-10-8.
2 mi S Walker on Gaylord Rd., 29 May 1965, 0-0-1.
Ouachita Parish
. Monroe, 0.2 mi S Forsythe Ave. on 19th St., 9 Aug. 1966, 0-5-2.
Saint Landry Parish
5.8 mi N Beggs on La. Hwy 182, 24 March 1965, 0-3-0.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 76
Pheretima hupeicnsis (Michaelsen, 1895)
1895 Perichaeta hupeiensis Michaelsen, Abhandl. Naturwiss. Verin
Hamburg 13:35-36 F ig . 11 and 12.
1899 Amyntas hupeiensis: Michaelsen, M itt. Naturhist, Mus. Hamburg
1 6 :6 -8 .
1955 Pheretima hupeiensis: Grant, Proc. U.S. N at. Mus. 105:49-63.
Diagnosis: Sexthecal, spermathecal pores minute and superficial,
on the anterior margins of vii-ix close to 6/7, 7/8 and 8/9, about 1/9
C apart. Female pore single, midventral on xiv. Male pores minute and
superficial in the center of small circular discs in the setal circle
of xviii, about 1/6 C apart. Genital markings always present, two pair
on 17/18 and 18/19 interrupting these intersegmental furrows, the
center of the markings slightly median to male pores. Clitellum
annular 13/14-16/17, dorsal pores and intersegmental furrows obliterated
but with setae present ventrally: 8-10/xiv, 5-10/xv, 10-16/xvi. Setae
small and closely crowded; 60-78/ii, 60-100/iii, 88-132/vi, 74-121/viii,
62-lüO/xii, 79-88/xx, 74-84/xxx, xviii/8-18, vi/10-16, vii/10-16,
viii/10-22. First dorsal pore in 11/12 or 12/13. Prostomium epilobic
with open tongue. Length, 45-222 mm. Diameter, 3-6 mm. Segments,
97-138.
Septa 4/5 thin, 5/6-13/14 present and more or less muscular.
Septa 8/9-9/10 are also present and muscular.' Intestine beginning in
X V . Intestinal caeca simple, arising in xxvii and extending forward
to about xxiv. Typhlosole simply lamelliform, scarcely recognizable
in xvi-xxvi, more evident following xxvii and ending in the region
Ixxxiii-xcii, Hearts of x-xiii latero-esophageal. Testis sacs
unpaired, U-shaped and vertical in x and xi. Seminal vesicles in xi
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 77
and xii, those of xi enclosed within the testis sac of xi. Prostates
in xvi, xvii-xviii, xix; with muscular C or S-shaped duct 2-4 mm long.
Vas deferens passing into prostatic duct close to the connection of the
latter with the prostate. Anterior pair of spermathecae located in vii,
posterior two pair usually contained within v iii. Muscular spermathecal
duct shorter than slightly enlarged ampulla. Ampulla not sharply
demarcared from duct. Diverticulum arising from anterior face of duct
at parietes, nearly twice as long as combined lengths of duct and
ampulla. Stalk of diverticulum with many internal convolutions.
Seminal chamber small and spherical at extreme ental end of diverticulum.
Genital markihg glands low and confined to, or extending only slightly
above parietes.
The above diagnosis is a composite taken from references covering
specimens from China, Japan, Korea, and the United States (Michaelsen,
1895, 1899b; Chen, 1933; Gates, 1937c, 1939, 1954, 1958; Kobayashi,
1938; Grant, 1955). Michaelsen (1895) stated that the species
possessed two pair of spermathecae but corrected this to three pair in
a later (1899b) publication.
Discussion; The 78 specimens of P. hupeiensis collected in
Louisiana conform well with the above diagnosis. The setal counts in
Louisiana specimens closely approximate those recorded (in parentheses)
in the literature: 91-103/iv (no record), 100-127/vi (88-132/vi),
92-119/viii (74-121/viii), 72-84/xx (79-88/xx), vi/8-15 (vi/10-16),
vii/8-16 (vii/10-16), viii/9-16 (viii/10-22), ix/9-15 (no record),
xviii/10-17 (xviii/8-18).
The 25 clitellate, nonautotomized specimens in this study ranged
from 111-150 mm long and from 3-5 mm in diameter, the number of
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 78
segments ranged from 121-146. The average length of these specimens
was 123 mm and the number of segments averaged 134. The 146 segm ents
found in a Louisiana specimen exceed the former maximum of138 segm ents
reported by Chen (1933), but the longest Louisiana specimen (15Ü mm) i s
somewhat shorter than the longest specimen Chen found in China (222 mm).
The posterior end of the typhlosole in Louisiana specimens (xc-c)
is somewhat more posterior than has been recorded in the literature
(Ixxxiii-xcii).
Eighteen of the specimens in this study autotomized when they
were collected, but no specimen was found in which regeneration had
o ccu rred .
The genital markings located in 17/18 and 18/19 measured from
0.48-0.72 mm in diameter in clitellate Louisiana specimens, the greater
diameter recorded in specimens with maximal clitella r development
(F igu re 19). The size of these genital markings has never been reported
in the literature.
Living specimens of P. hupeiensis in Louisiana were found with
the same green coloration and characteristic odor resembling that of
raw carrots, recorded by Chen (1933) and Gates (1937c). This species is
less active than the other species of Pheretima occurring in Louisiana,
and when removed from the soil it coils into a rather tight series of
convolutions and remains inactive. During rain, this species had been
collected while it was crawling on sidewalks of the LSU campus.
Distribution; Pheretima hupeiensis is widely distributed in China,
its presumed original home (Michaelsen, 1895; Chen, 1933). It has also
been reported from Japan (Michaelsen, 1899b); Korea (Kobayashi, 1938);
and Taiwan (Tsai, 1964) . In the United States it has been reported
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 79
from Pennsylvania (Chen, 1933); Maryland, D istrict of Columbia (Gates,
1937c); Louisiana (Harman, 1952); Arkansas (Causey, 1953); L o u isia n a ,
New York (G a tes, 1954); Connecticut, New York (Grant, 1955); M aryland,
New York (G a te s, 1958); Kentucky, Tennessee (Gates, 1963); Pennsylvania
( B h a t ti, 1965); and Utah (Gates, 1967) in outdoor sites. It has also
been reported from greenhouses in Maine, Connecticut, and New York
(G a te s, 1963).
In Louisiana, P. hupeiensis was collected from 11 parishes at 15
sites scattered in southern and central Louisiana (Map 6). Five of
the sites are the floodplains of bayous or rivers or the margins of
lakes, where it may have been carried by fishermen. Six of the sites
are in yards, where it may have been introduced with exotic plants. It
was also found at 2 sites in botanical gardens, which it may have
reached by the same means. It was found at one site each in a dump and
a roadside ditch. The Louisiana distribution of this species appears
to be the result of the activities of man, as it was seldom found in
areas away from the habitations or activities of man.
Specimens with slight clitellar development were found in
Louisiana in every season of the year, but maximal development of the
clitellum was found only in October. Specimens from December
collections appear to have a postsexual clitellum . From the above it
would appear that cocoons are produced "n October and hatch in the
spring. Immature specimens were found only in May.
Localities;
Avoyelles Parish
2.7 mi E Bordelonville on La. Hwy 451, 20 March 1966, 0-9-1.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 80
Beauregard Parish
L o n g v ille Lalce on La. Hwy 110, 1 May 1966, 0-7-1.
Concordia Parish
4 mi W Shaw on La. Hwy 910, 19 March 1966, 0-0-5.
East Baton Rouge Parish
L. S. U. Campus, beside Dodson Pond, 3 May 1964, 0-5-2,
L. S. U. Campus, beside Library, 4 Oct. 1964, 0-0-10.
L. S. U. Campus, beside Dodson Pond, 30 D ec. 1964, 0-0-2,
McDonald Ave. near Highland Rd., 7 A p ril 1965, 0-5-7.
L. S. U. Campus, Audubon Hall, 29 May 1965, 1-1-2.
L. S. U. Campus, beside Library, 10 July 1965, 0-0-1.
L. S. U. Campus, beside football stadium, 15 Dec. 1965, 0-2-2.
Iberia Parish
Avery Island, Jungle Garden, Timber bamboo, 27 April 1965, 0-0-1.
Iberville Parish
2.7 mi NW Sunshine on La. Hwy 337, 6 Dec. 1964, 0-0-4.
Orleans Parish
Audubon Park, 26 June 1965, 0-0-3.
Pointe Coupee Parish
3.2 mi N Oscar on La. Hwy 1, 14 March 1965, 0-3-0.
Saint Charles Parish
2.4 mi NW Norco on US Hwy 61, 21 Feb. 1965, 0-0-1.
Terrebonne Parish
2.3 mi SE Gibson on US Hwy 90, 8 April 1967, 0-2-0.
West Baton Rouge Parish
1 mi N Addis on River Rd., 23 Oct. 1965, 0-0-1.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 81
Pheretima minima (Horst, 1893)
1893 Perichaeta minima Horst, ^ Weber, M., Zool. Ergeb, Reise
Niederlandisch Ost-Indien 3:66-68 F ig . 27.
1942 Pheretima humilis Gates, Bull. Mus. Comp. Zool. 89:120-122.
1961 Pheretima minima: Gates, Z ool. Meded. Rijksmus. Leiden 37:298-
304.
Diagnosis: Bithecal, spermathecal pores minute and superficial,
widely separated in 5/6. Female pore single, midventral on xiv. Male
pores minute and superficial, each on a circular porophore in the setal
circle of xviii, about 1/3 C apart. Genital markings, when present,
small circular tubercules with a central translucence, paired and
presetal near spermathecal pore level or more medially in some of
v-viii, rarely postsetal in vi near spermathecal pore level; unpaired
and median, presetal in some of xvii, xix-xxi. Clitellum annular
13/14-16/17, dorsal pores and intersegmental furrows obliterated, with
3-12 setae ventrally in xvi. Setae closely crowded in preclitellar
region, further apart posteriorly, 40-55/ii, 40-47/iii, ca. 75/vi,
60/vii, 45-58/viii, 46-52/xii, 38-44/xx, v/26-29, vi/27-33, xviii/6-10.
First dorsal pore at 11/12-12/13. Prostomium epilobic with open
tongue. Length, 20-56 mm. Diameter, 1.5-2 mm. Segments, 75-97.
Septa 5 /6 -7 /8 and 10/11-11/12 somewhat thickened, 8/9 c o m p lete,
or present only ventrally, membranous,9/10 absent. Intestine
beginning in xv. Intestinal caeca simple, arising in xxvii and
extending forward to about xxiv. Typhlosole simply lamelliform,
arising in xxvii and extending back to region of xlviii-lxiv. Hearts
of x-xi (latero?) esophageal, those of x ii-x iii latero-esophageal.
Testis sacs unpaired, U-shaped to annular in x and xi. Seminal
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 82
vesicles in xi and xii, those of xi included within testis sac of xi.
Prostates in some or all of xvi-xix, muscular ducts 1-2 mm long,
twisted or in hair pin loop. Spermathecae in vi. Non muscular
spermathecal duct as long or longer than ampulla. Diverticulum as long
or longer than combined lengths of duct and ampulla, arising from
median face of duct. Ental portion of diverticulum constricted into 2
or 3 seminal chambers, separated from each other by a short neck-like
region. Genital marking glands stalked and coelomic or bound down to
p a r i e t e s .
Since the original description of Horst (1893) lacks many details,
the above diagnosis is taken from the references of Gates (1942b, 1961,
MS), who has examined the type specimen. In the reexamination of the
type material. Gates (1961) states that the spermathecae are in vi
opening at 5/6 and not in vii as recorded by Horst (1893).
Discussion: The 23 specimens of _P. minima collected in Louisiana
conform well with the above diagnosis. Without exception, these
Louisiana specimens lack genital markings. Genital markings were also
lacking in a series of 6 specimens described by Gates (1942b) from
Rangoon, Burma.
The ranges of setal numbers in Louisiana specimens correspond
closely with those recorded (in parentheses) in the literature;
34-40/ii (40-55/ii), 39-44/iii (40-47/iii), 40-51/iv (no record),
50-60/vi (ca. 75/vi), 48-60/viii (45-58/viii), 38-44/xx (38-44/xx)
(38-44/xx), v/21-24 (v/26-29), vi/22-25 (vi/27-33), xviii/2-8
(xviii/6-10).
The typhlosole in these Louisiana specimens ends in the region of
Ivi-lix, well within the reported range for the species (xlviii-lxiv).
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 83
The 12 nonautotomized specimens in this study ranged from 21-35
mm long and from 1.5-2 mm in diameter, the number of segments ranged
from 76-97. The average length of these specimens was 28 mm, and
the number of segments averaged 84.
P. minima is a very active species that jumps and thrashes about
on the ground when disturbed. In such instances it may autotomize a
posterior portion of the body as did 4 of the 23 specimens collected
in this study. The level of autotomy ranged from 50/51-71/72 and was.
always in the intersegmental furrow.
Three posterior regenerates were observed in this study. In the
first worm of 66 segments, the last 7 segments had been regenerated.
The second of 72 segments had regenerated 15 segments. The third of
76 segments had regenerated 10 segments. The regenerated segments can
be recognized by their shorter length and smaller diameter.
Only one of the 22 clitellate specimens possessed setae on the
clitellum and in that instance there were only 2 setae ventrally on
x v i.
Five specimens of _P. minima from Louisiana possess spermathecae,
four are from the Avery Island collections in Iberia Parish, and one
specimen with a single spermatheca on the left is from the Lincoln
Parish collection. The remaining 18 specimens that lack spermathecae
are parthenogenetic A morphs according to the terminology of Gates
(1956). One specimen has a single spermatheca on the right at 5/6 and
is an intermediate (I) morph. Two specimens possess a single sperma
theca on the left at 5/6, also I morphs. Two specimens possess a pair
of spermathecae at 5/6, in one specimen sperm were observed in the 3
seminal chambers after the spermatheca had been cleared in xylene and
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 84
mounted in Harleco synthetic resin. The species appears to reproduce
in the normal hermaphroditic, biparental manner at Avery Island, at
least those specimens with spermathecae may, since sperm had been
exchanged.
All of the spermathecae in the specimens from Avery Island are
like that illustrated in Figure 20 with 3 seminal chambers, while that
of the Lincoln Parish specimen is somewhat abnormal, the diverticulum
is shorter than the combined length of the duct and ampulla, the seminal
chamber is not completely divided into 3 seminal chambers.
The testis sac of x and xi were found to be annular in one
specimen. The testis sac of xi was found to be annular in two other
specimens, and in all the specimens the seminal vesicles of xi were
included within the testis sac of xi, though the determination is
difficult due to the small size of the specimens. The seminal vesicles
are small vertical bodies in all of the Louisiana specimens and they do
not fill the coelomic cavities of their respective segments.
Septum 8/9 was present only as a membranous ventral rudiment
under the gizzard in 2 specimens. In the remainder of the specimens
8/9 was apparently absent. Septum 9/10 was absent in all of the
Louisiana specimens.
Distribution; P. minima was erected for specimens from Java and
has since been reported from Java, Australia, Burma, and Hawaii (Gates,
1961) . The present specimens represent the first records of the
species in the continental United States.
In the present study, _P. minima was found in 5 parishes at 10
sites all of which are areas where the worms could have been introduced
with exotic plants (Map 7). Four of the sites are in a botanical
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 85
garden, four of the sites are in flower beds in yards or parks, and two
sites are in nurseries. P. minima was never collected in a site away
from the activities of man in Louisiana.
C litellate specimens were collected in March, April, June, and
August. Nothing can be said concerning the life history of the species
from the present series because no immature specimens were collected.
Localities;
East Baton Rouge Parish
McDonald Ave. near Highland Rd., 7 A p ril 1965, 0-0-1.
W Parker Blvd. at Dodson Ave., 8 April 1965, 0-0-2.
Iberia Parish
Avery Island, Jungle Garden, Timber Bamboo, 27 A p ril 1965, 0-0-3.
Avery Island, Jungle Garden, Sunken garden, 27 A p ril 1965, 0-0-5.
Avery Island, Jungle Garden, Camélia garden, 27 A p r il 1965, 0-0-6.
Avery Island, Jungle Garden, Cactus garden, 27 A p r il 1965, 0-0-1.
Lincoln Parish
Ruston, Rigdell's Nursery, 21 Aug. 1954, 0-1-1.
Orleans Parish
Audubon Park, 26 June 1965, 0-0-1.
Chartres St. at Ursulines Ave., 28 April 1967, 0-0-1.
Sabine Parish
Hodges Gardens, Nursery, 26 March 1965, 0-0-1.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 86
Pheretima morrisi (Beddard, 1892)
1892 Perichaeta morrisi Beddard, Proc. Zool. Soc. London 1892:166.
1937 Pheretima m orrisi: Gates, Bull. Mus. Comp. Zool. 80:361-363.
Diagnosis: Quadrithecal, spermathecal pores minute and superfi
cial in 5/6-6/7, nearly 1/2 C apart. Male pores minute and superficial
on small discoidal porophores in the setal circle of xviii, about 1/3
C apart. Female pore median on xiv, occasionally paired. Genital
markings, very rarely lacking, small discs; presetal, unpaired and
midventral nearer setal circle than intersegmental furrows in some or
all of v-ix and xviii-xix; presetal, paired (1-3 pairs), just median to
the spermathecal pore levels or more medially in some of vi-ix; two
just median to each male porophore disc, one presetal and the other
postsetal forming a triangle surrounded by the circular wrinkles of
the male porophores; 1-8 presetal in some of xvii-xix, 1-8 postsetal on
xvii-xviii, nearer to intersegmental furrows than to segmental equators
and median to male pores. Clitellum annular 13/14-16/17, occasionally
failing to reach 13/14 or 16/17, dorsal pores and intersegmental
furrows obliterated, some setae usually present ventrally in xiv-xvi.
Setae: 23-39/iii, 40/v, 42-56/viii, 42-56/xii, 46-59/xx, vi/16-28,
xviii/10-17. First dorsal pore at 10/11. Prostomium epilobic with
open tongue. Length, 40-150 mm. Diameter, 2.5-6 mm. Segments, 75-100,
but usually 92-98.
Septa 8/9-9/10 absent, none thickly muscular. Intestine beginning
in X V . Intestinal caeca simple with short ventral lobes, arising in
xxvii* and extending forward to about xxiv. Typhlosole quite
rudimentary or lacking. Hearts of x (latero?) esophageal, those of
xi-xiii latero-esophageal. Testis sacs paired and ventral in x and xi.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 87
Seminal vesicles medium sized in xi and xii. Prostates in some of
xvii-xxiii, with muscular U or C-shaped ducts 2-3 mm long. Spermathecae
in vi and vii. Spermathecal duct elongate and slender, shorter than
pear-shaped ampulla. Diverticulum arising from anterior face of duct
at parietes, shorter or longer than combined lengths of duct and
ampulla, slightly and gradually widened entally to form seminal chamber.
Genital marking glands stalked and coelomic, composite.
Since the original description of Beddard (1892b) lacks many
details, the above diagnosis is taken from the works of Gates (1937c,
1939, MS), who has examined material referred to the species by
Beddard, and from other references covering the species (Righi and
Knepper, 1965; Sims, 1964; Tsai, 1964).
_ Discussion; The 138 specimens collected in Louisiana conform well
with the above diagnosis. The number of setae on various segments
correspond closely with those reported (in parentheses) in the
literature, 29-41/iv (no record), 32-51/vi (no record), 35-54/viii
(42-56/viii), 48-64/xx (46-59/xx), v/15-25 (no record), vi/20-28
(vi/16-28), vii/19-29 (no record), xviii/8-20 (xviii/10-17).
The typhlosole in Louisiana specimens is low and barely
lamelliform but recognizable, arising in xxvii and ending in the region
of Ixv-lxxiv.
The clitellate, nonautotomized specimens in this study ranged
from 59-144 mm long and from 3-5.5 mm in diameter, the number of
segments ranged from 79-97. The average length of 43 of these speci
mens was 90 mm and the number of segments averaged 92.
This species is very active and when disturbed will jump and
thrash about on the surface of the ground. It may autotomize a
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 88
posterior portion of the body in such circumstances as did 20 of the
138 specimens collected in this study. The level of autotomy ranged
from 68/69-89/90, and was always in the intersegmental furrow.
• Posterior regenerates, which are recognizable by the shorter
length and smaller diameter of the regenerated segments, were
recognized at various levels between 67/58 and 89/90. At the anterior
level 16 segments were regenerated, while at the more posterior level
3 segments were added. Sixteen segments was the maximum number of
regenerated segments observed in this study.
Records of the location of the genital markings (GMs) were kept
on 111 specimens. The most common pattern of the midventral GMs was
one in each of vi, vii and v iii (Table 10). Thirty-nine specimens
lacked midventral GMs, while 11 specimens lacked all preclitellar GMs
(Tables 10 and 11). Eight specimens possessed midventral GMs but
lacked the lateral preclitellar GMs (Table 11). The postclitellar GMs
are quite variable in their location and numbers (Table 12), but without
exception the Louisiana specimens all possess the two GMs just median
to each male pore, one presetal and the other postsetal inside the
wrinkles of the male porophore. The GMs listed in Table 12 are in
addition to the midventral GMs listed in Table 10. A common pattern of
GMs can be seen in Figures 22 and 23.
Distribution; _P. morrisi is widely distributed in China, which
may be its original home (Gates, 1963). It has also been reported
from Burma, India, Thailand, Malay Peninsula, Sumatra, Hawaii, Barbados,
Peru, Brazil, Argentina, Chile, Cape Verde Islands, St. Helena (Gates,
MS); Ascension Island (Sims, 1964); and Taiwan (Tsai, 1964). In the
United States it has been reported from greenhouses in New York
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 89
(Gates, 1954); Maine (Gates, 1963); and Michigan (Gates, 1966). In
outdoor sites it has been reported from Mississippi, Louisiana, Texas
(Gates, 1954).
Although P. morrisi has been introduced into the northern United
States by way of greenhouses on numerous occasions, it apparently has
been able to establish itself outdoors only in the southern United
States (Florida, Louisiana, M ississippi, Texas).
In the present study, _P. morrisi was found in 23 parishes at 40
sites in southern, central, and northern Louisiana (Map 8). Nineteen
of the sites are the floodplains of bayous and rivers or the margins of
lakes and swamps. Fifteen of the sites are in flower beds or similar
sites in yards, where it may have been introduced with exotic plants.
P. morrisi was also found at one site each in a cemetery, a botanical
garden, the edge of a salt marsh, in cultivated soil, under grassland,
and in the rotting bagasse at a sugar mill.
In the extreme western and northern part of its range in Louisiana,
_P. morrisi is limited to yards and has not dispersed away from these
sites of introduction as it has done in the southeastern portion of the
state. With the passage of time, _P. morrisi can be expected to increase
its range in the western and northern parts of the state.
C litellate specimens of P. morrisi were collected in every month
of the year except January, July, and November. Nothing can be said
concerning the life history of this species in Louisiana, because no
immature specimens were collected.
Localities;
Ascension Parish
Donaldsonville, Charles St. at Lafourche St., 28 April 1965, 0-0-1.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 90
Donaldsonville Fairgrounds, 28 A p ril 1965, 0-0-8.
Assumption Parish
Bellerose, on Bayou Lafourche, 14 A p r il 1967, 0-1-0.
Near Lafourche Par. line on La. Hwy 1, 14 A p r il 1967, 0-0-1.
Avoyelles Parish
1 mi N Bunkie on US Hwy 71, 29 March 1965, 0-0-2.
Calcasieu Parish
Moss B lu ff on La. Hwy 378, 25 March 1967, 0 -5 -4 ,
Cameron Parish
Cameron, o ld Henry Home, 20 S e p t. 1966, 0-0-22.
Grand Chenier, near post office, 21 S e p t. 1966, 0-0-3.
East Baton Rouge Parish
L. S. U. Campus, beside Dodson Pond, 3 May 1964, 0-0-1.
McDonald Ave. near Highland Rd., 7 A p r il 1965, 0-0-2.
Highland Rd. at Dentation Dr., 8 April 1965, 0-0-2.
W. Parker Blvd. at Dodson Ave., 8 April 1965, 0-0-3.
Iberia Parish
Avery Island, Jungle Garden, Timber bamboo, 27 A p r il 1965, 0-2-4.
Iberville Parish
1 mi NW Sunshine on La. Hwy 337, 18 Oct. 1964, 0-0-2.
1.6 mi NW Sunshine on La. Hwy 337, 12 Dec. 1964, 0-0-10.
Jefferson Parish
Grand Isle, Chighizola Lane, 9 A p r il 1967, 0-0-1.
Jefferson Davis Parish
2.6 mi W Jennings on US Hwy 90, 26 March 1967, 0-0-1.
Lafourche Parish
0.7 mi S Lockport on La. Hwy 1, 9 April 1967, 0-0-1.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 91
2 mi NW Thibodaux on La. Hwy 1, 9 April 1967, 0-0-1.
Orleans Parish
Audubon Park, 26 June 1965, 0-1-6.
Chartres St. at Ursulines Ave., 28 April 1967, 0-1-0.
City Park, Palm Dr. at Roosevelt Mall,28 A p r il 1967, 0-0-2.
City Park, Stadium Dr. at Roosevelt Mall, 28 April 1967, 0-0-1.
L ittle Woods on Hayne Dr., 28 April 1967, 0-1-3.
Plaquemines Parish
3 mi S Pointe a la Hache on La. Hwy 39, 22 April 1967, 0-0-2.
5.8 mi N Boothville on La. Hwy 325, 22 April 1967, 0-3-5.
Rapides Parish
Alexandria, 12 April 1955, 0-0-4.
Red River Parish
0.4 mi E Red River on US Hwy 84, 26 Aug. 1966, 0-0-1.
0.1 mi E Red River on US Hwy 84, 29 Aug. 1956, 0-3-2.
Saint Charles Parish
2.4 mi NW Norco on US Hwy 61, 21 Feb. 1965, 0-0-1.
1 mi NE US Hwy 61 on W Bonnet Carre Spillway levee, 13 March 1965,
0 -1 -4 .
2 mi NE Norco on E Bonnet Carre Spillway levee, 20 Dec. 1965, 0-1-5.
Saint James Parish
1 mi S Convent on La. Hwy 44, 5 Dec. 1965, 0-0-1.
Saint John The Baptist Parish
2.5 mi N Killona on La. Hwy 18, 30 April 1967, 0-0-1.
Saint Landry Parish
Bayou Cortableau at US Hwy 190, 15 Oct. 1966, 0-1-2.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 92
S ain t Mary P arish
Morgan C ity , cem etery on Young D r., 2 A p ril 1967, 0-0-4.
Terrebonne Parish
2 .3 mi SE Gibson on US Hwy 90, 8 April 1967, 0-0-2.
Vermillion Parish
0.2 mi S Kaplan on La. Hwy 35, 31 March 1967, 0-0-1.
Erath, sugar mill on W Landry St., 1 April 1967, 0-0-1.
West Baton Rouge Parish
1 mi N Addis on River Rd., 23 Oct. 1965, 0-0-1.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 93
C. Key to L ouisiana Species of Pheretim a
1. Intestinal caeca glove-shaped (manicate)...... 2
1'. Intestinal caeca simple ...... 3
2(1). Spermathecal pores at 5/6-7/S, with paired areas of thin,
wrinkled brown epidermis in some of iv-xi . . P. agrestis
2' Spermathecal pores at Ô/7-7/8, GMs in midventral presetal
patches in some of vi-xi and xvii-xxi . . P. hilgendorfi
3(1'). Male pores invaginate ...... P. californica
3'. Male pores superficial...... 4
4(3'). Four pairs of spermathecal pores at 5/6-8/9. _P. diffringens
4 ' . Less than four pairs of spermathecal pores...... 5
5(4'). Three pairs of spermathecal pores ...... 6
5'. Less than three pairs of spermathecal pores ...... 7
6(5). Spermathecal pores at 5/6-7/8 ...... P. hawayana
6'. Spermathecal pores at or just behind
Ô /7-8/9 ...... P. hupeiensis
7(5'). Two pairs of spermathecal pores at 5/6-6/7. . P. morrisi
7'. One pair of spermathecae in vi...... P. minima
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. IV. ECOLOGY
Ranked in order of the number of sites at which they were
collected, the most common species of Pheretima in Louisiana is _P.
diffringens, followed by P. hawayana, P. californica, P. morrisi. P.
agrestis, P. hupeiensis. P. minima, and P. hilgendorfi.
Table 1 shows the 38 combinations of the species of Pheretima
that occurred at 448 sites in Louisiana. A total of 307 collections
contained only one species, 89 collections contained 2 species, 34
collections contained 3 species, 14 collections contained 4 species,
3 collections contained 5 species, and one collection contained 6
s p e c ie s .
Every species of Pheretima in Louisiana, except P. hilgendorfi,
was found occurring singly in at least one site. _P. diffringens was
found singly at 274 sites and in association with at least one other
species at 102 sites. All of the other species were found in associa
tion with other species more often then they were found occurring
singly (Table 1).
Table 2 shows the total number of times that each species was
found in association with every other species. P. diffringens was
found in association with every other species of Pheretima in this
study. It occurred with P. minima at 4 of 10 sites (40%) and was found
more frequently with each of the other species. P. hawayana was found
in association with every other species except P. agrestis. P.
californica, _P. hupeiensis, P. minima, and _P. morrisi were found in
association with every other species in this study except P. agrestis
94
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 95
and hilRendorfl. This is due to the small number of times that the
latter two species were collected and that P. agrestis is limited in its
distribution to northern Louisiana (Map 1). agrestis was found in
association with P. diffringens and P. hilgendorfi. P. h ilg e n d o rfi was
found in association with _P. a g r e s t i s , P. diffringens, and P. hawayana
(Table 2 ) .
While P. diffringens was found to be the most common species based
on the number of sites at which it occurred, it was not consistently
found in greater numbers than the other species with which it was
associated. Neither was any of the other species found to be distinctly
predominant in numbers over any other species when all sites at which
they were associated are considered.
At 62 of the sites in the present study, Lumbricidae were found
in association with the various species of Pheretima. At 66 of the
sites Diplocardia and Pheretima were found in association. At 63 sites
Lumbricidae, Diplocardia, and Pheretima were found in association. At
3 sites Microscolex and Pheretima were found in association. At 3 sites
Sparganophilus and Pheretima were found in association. At 197 sites
other genera of earthworms were found in association with Pheretima,
while at 251 sites Pheretima was found occurring alone. If Pheretima is
indeed able to eliminate other species of earthworms from areas in which
it occurs (see Stephenson 1930, p. 681), this work can be used for
comparison with later surveys.
At 20 sites in which Pheretima were collected the pH of the soil
was measured. _P. californica was found in a pH range of 7.5 to 8.1.
_P. diffringens was found in a pH range of 6.0 to 8.5. P. hawayana was
found in a pH range of 6.5 to 8.5. P. morrisi was found in a pH range
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 96
of 6.8 to 8.5. P. hupeiensis was found in a pH of 7.0. Schread (195i)
has reported P. hupeiensis from soils in New York with a pH value
ranging from 4.72 to 6.35. Olsen (1928) reported earthworms from soils
in Ohio with pH values ranging from 5.45 to 8.7, but his report
contained no Pheretima.
The topography of Louisiana varies from a southern coastal plain,
which includes a coastal marsh some 20 miles wide and slightly above
sea level, to an upland pine area with elevations up to 500 feet. The
wide deltaic plain of the Mississippi River runs down the east boundary
of the state and spreads out to about 100 miles wide at the level of
New Orleans. A more nearly complete account of the topography of the
state can be found in Davis (1965).
The various species of Pheretima in Louisiana are not limited to
any one drainage system in the state and each species is also found in
several different soil and vegetation types (Maps 1-9). There is no
correlation between the natural ecological divisions, the vegetation
types, or the soil types in Louisiana and the distribution of Pheretima
species in the state.
The presumed mode of introduction of the various species of
Pheretima into the United States was in soil around the roots of
imported plant material. Eighty-seven of the species of exotic woody
plants reported from Louisiana by Brown (1961) are listed by Bailey
(1949) as native to the regions where Pheretima has its highest degree
of endemicity (China, Japan, Burma, Malaysia). Sixty-four of these
plant species would probably have been imported as rooted cuttings, the
remainder being more easily propagated from seed (Brown, Personal
Communication). The number of times that Pheretima could have reached
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 97
Louisiana with imported plant materials would thus appear to be
extremely high. Gates (1965a) has also listed 6 other methods by
which exotic earthworms could have entered Louisiana.
Gates (1963, 1966) has listed 10 species of Pheretima from
greenhouses or nurseries in the United States "... from whence the
worms could have been distributed more or less widely in accordance with
sales patterns of potted plants." This list includes all of the
species of Pheretima occurring in Louisiana except _P. minima. In the
present study P. minima is reported from a botanical garden and 2
nurseries, all sites that it may have reached with imported plants.
Thus the total number of Pheretima species reported from greenhouses in
the United States now stands at 11.
Three species included in the list of greenhouse pheretimas
(P. bicincta, JP. levis, and _F. rodericensis) were not found in
Louisiana. JP. bicincta has been reported in the United States only
from greenhouses in Maine (Gates, 1963), while JP. levis has been
reported from natural outdoor sites in New Jersey (Davies, 1954) and
Pennsylvania (Bhatti, 1965). JP. rodericensis has been found at an
outdoor site in Florida (Gates, 1954), P. posthuma has also been found
occurring in Florida (Gates, Personal Communication). In the future
these 4 species may reach Louisiana in plant materials shipped by
nurseries or by the methods listed by Gates (1965a).
Four other species (P. elongate, P. houlleti, P. robusta, P.
schmardae) known in various islands of the West Indies will probably
never reach Louisiana because of the prohibition against bringing soil
into the country and the fumigation of imported plant material enforced
by the Plant Quarantine Division.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. V. SUMMARY
The distribution of 8 species of the introduced earthworm genus
Pheretima (P. agrestis. P. californica. P. diffringens. P. hawayana.
_P. hilgendorfi. P. hupeiensis. P. minima. P. morrisi) is reported from
a total of 448 sites in all 64 parishes of Louisiana. All of these
species have previously been reported from the state except P. minima.
which is first reported from the continental United States and
Louisiana in this study.
P. diffringens was collected at 376 sites and is widely distributed
over the entire state. _P. hawayana was found at 128 sites, commonly
distributed in the southeastern portion of the state, but limited to
sites of introduction in the northern and western parts of the state.
P. californica was obtained at 70 sites and is widely distributed in
the southeastern portion of the state. JP. morrisi was secured at 40
sites, scattered in the southeastern portion of the state and at sites
of introduction in the northern and western extremes of the state.
_P. agrestis was collected at 17 sites in the northern portion of the
state, largely limited to areas of introduction. _P. hupeiensis was
found a t 15 sites in southern Louisiana, largely limited to areas of
introduction. JP. minima was obtained at 10 sites of introduction,
scattered over the state. _P. hilgendorfi was secured at 8 sites of
introduction, likewise scattered over the state.
The genus Pheretima has rather broad ecological requirements and
occupies several different soil and vegetation types in Louisiana. It
also occurs in all of the natural ecological divisions of the state
98
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 99
and has been able to cross drainage systems in the state. At 141 sites
two or more species of Pheretima were found in association, while at
one site 6 species of Pheretima were found in association.
At 197 sites other genera of earthworms were found in association
with Pheretima, while at 251 sites Pheretima was found occurring alone.
This information can be used in comparison with future surveys to
determine what effect, if any, Pheretima has in suppressing or
supplanting other earthworms which occur in Louisiana.
Five species (P. californica, P. diffringens, _P. hawayana, P.
hupeiensis, P. morrisi) remain active and clitellate all year in
Louisiana. _P. minima was secured on so few occasions that nothing can
be said concerning its life history. _P. agrestis and P. hilgendorfi
have annual life cycles in which the adults die in the fall and the
species overwinters in the cocoon.
All of the species of Pheretima in Louisiana, except P.
hupeiensis, are extremely active and all of them will autotomize when
disturbed. The percentage of specimens that autotomized when collected
ranged from 8.5% in P. californica to 42.6% in P. hilgendorfi.
Regeneration was noted in every species except jP. hupeiensis.
The manner of regeneration in P. agrestis and _P. hilgendorfi differs
from that of the other species since each regenerated segment attains
a size comparable to that of the old segments, before another new
segment is completed. These two species also appear to add segments at
the posterior end of nonautotomized adults in the same manner as in
regeneration.
In the Louisiana species of Pheretima there appears to be two
groups. One group (P. agrestis, P. hilgendorfi) has manicate
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 100
intestinal caeca, an annual life cycle, and slow regeneration. The
other group (^. californica, _P. diffringens. P. hawayana, _P. morrisi)
has simple intestinal caeca, a life history with year-round activity,
and rapid regeneration. These combinations of characteristics would
appear to provide a basis for the division of the genus into subgenera.
Subgenera are not erected in the present study since this apparent
dichotomy of the genus would not stand when all of the species of the
genus are considered.
The location of the genital markings of those species that possess
them yas recorded. The characteristic genital markings, while variable,
have a distinct diagnostic value. The genital marking glands of P.
diffringens are found to be composed of a secretory end piece supported
by a stalk that contains numerous ducts. In the sections at hand, each
cell in the secretory end piece appears to have an individual elongate
duct leading to the exterior through the stalk.
The male pores of P. agrestis and hilgendorfi, which are
located inside parietal invaginations, are described from the first
United States specimens to possess male terminalia.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. LITERATURE CITED
Bahl, K. N. 1922. On the development of the 'enteronephric' type of nephridial system found in Indian earthworms of the genus Pheretima. Quarterly Journal of Microscopical Science 66(1); 49-104.
Bailey, L. H. 1949. Manual of cultivated plants. 2nd ed. New York: MacMillan Co., 1116 pp.
Baird, W. 1869. Description of a new species of earthworm (Megascolex diffringens) found in North Wales. Proceedings of the Zoological Society of London 1869:40-43.
Beddard, F. E. 1890. Contributions to the anatomy of earthworms, w ith d e s c rip tio n s of some new s p e c ie s . Q u a rte rly Jo u rn al of Microscopical Science 30(4):421-479.
______.1892a. On some Perichaetidae from Japan. Zoblogische Jahrblicher, Abtheilung für Systematik, Géographie und Biologie Thiere, Jena 6(5): 755-766.
______.1892b. On some species of the genus Perichaeta (sensu stricto) . Proceedings of the Zoological Society of London 1892:153-172.
.1895. A Monograph of the Order of Oligochaeta. Oxford; Clarendon P re ss, 769 pp.
Bhatti, H. K. 1965. Earthworms of Swarthmore, Pennsylvania, and vicinity. Proceedings of the Pennsylvania Academy of Science 39:8-24.
Brown, C. A. 1961. Check list of woody plants of Louisiana, native, naturalized, and cultivated. 2nd ed. Louisiana Forestry Commission Bulletin, No. 8:1-16.
Causey, D. 1952. The earthworms of Arkansas. Proceedings of the Arkansas Academy of Science 5:31-41.
______.1953. Additional records of Arkansas earthworms. Proceedings of the Arkansas Academy o f Science 6:47-48.
Chen, Y. 1933. A preliminary survey of the earthworms of the lower Yangtze valley. Contributions from the Biological Laboratory of the Science Society of China, Nanking, Zoological Series 9(6): 177-296.
101
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 102
C o g n e tti, L, 1914. Oligochaeta recueillis pendant la "Nederl. Nieuw Guinea Expeditie" (1903). Nova Guinea, Zoology5(5): 543-564.
Davies, H. 1954. A preliminary list of the earthworms of northern New Jersey with notes. Breviora, No. 26:1-13.
Davis, E. A. 1965. Louisiana, a narrative history. 2nd ed. Baton Rouge: Claitor's Book Store, 394 pp.
Gates, G. E. 1929. A summary of the earthworm fauna of Burma with descriptions of fourteen new species. Proceedings of the United States National Museum 75:1-41.
______.1935a. New earthworms from China with notes on the synonymy of some Chinese species of Drawida and Pheretima. Smithsonian Miscellaneous Collections 93(3):1-19.
.1935b. On some Chinese earthworms. Lingnan Science Journal 14(3):445-457.
.1937a. Indian earthworms. I. The genus Pheretima. Records of the In d ian Museum, C a lc u tta 39:175-212.
.1937b. Notes on some species of Drawida and Pheretima with descriptions of three new species of Pheretima. Bulletin of the Museum of Comparative Zoology Harvard College 80(7): 305-335.
.1937c. The genus Pheretima in North America. Bulletin of th e Museum o f Com parative Z oology Harvard C o lle g e80(8): 339-373.
.1939. On some Chinese earthworms with special reference to specimens collected in Szechwan by Dr. D. C. Graham. P ro ceed in g s o f th e U n ited S ta te s N a tio n a l Museum 85:405-507.
.1942a. Checklist and bibliography of North American earthworms. American Midland Naturalist 27(1):86-108.
.1942b. Notes on various peregrine earthworms. Bulletin of th e Museum o f Com parative Z oology Harvard C o lle g e89(3): 63-144.
.1953a. Further notes on the earthworms of the Arnold Arboretum, Boston. Breviora, No. 15:1-9.
.1953b. On the earthworms of the Arnold Arboretum, Boston. Bulletin of the Museum of Comparative Zoology Harvard College 107(10):501-534.
.1954. Exotic earthworms of the United States. Bulletin of the Museum of Comparative Zoology Harvard College 111(6):219- 258.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 103
.1956. Reproductive organ polymorphism in earthworms of the oriental Megascolecine genus Pheretima Kinberg 1867. E v o lu tio n 10(2):213-227.
.1958. On some species of the oriental earthworm genus Pheretima Kinberg, 1867, with key to species reported from the Americas. American Museum Novitates 1888:1-33.
.1959. On a taxonomic puzzle and the classification of earthworms. Bulletin of the Museum of Comparative Zoology Harvard College 121(6):229-261.
.1960. On Burmese earthworms of the family Megascolecidae. Bulletin of the Museum of Comparative Zoology Harvard College 123(6):203-282.
.1961. On some species of the oriental earthworm genus Pheretima Kinberg, 1867. Zoologische Mededelingen Rijksmuseum van Natuurlijke Historié Leiden 37(18): 293-312.
.1963. Miscellanea megadrilogica. VII. Greenhouse earthworms. Proceedings of the Biological Society of Washington 76:9-18.
.1965a. Louisiana earthworms. I. Apreliminary survey. Proceedings of the L o u isian a Academy of Sciences 28:12-20.
.1965b. On variation in an anthropochorous species of the oriental earthworm genus Pheretima Kinberg 1866. Proceedings of the Biological Society of Washington 78:1-16.
.1966. Requiem-for megadrile utopias. A contribution toward the understanding of the earthworm fauna of North America. Proceedings of the Biological Society of Washington 79:239-254.
.1967. On the earthworm fauna of the great American desert and adjacent areas. Great Basin Naturalist 27(3): 142-176.
______.M .S. Unpublished manuscript entitled "Burmese Earthworms." 1195 pp.
Goto, S., and S. Hatai. 1899. New or imperfectly known species of earthworms. No. 2 . Annotationes Zoologicae Japonenses 3 (1 ): 13- 24.
G rant, W. C., J r . 1955. An anatomical study of the peregrine Megascolecid earthworm Pheretima hupeiensis in the eastern United States. Proceedings of the United States National Museum 105:49-63.
Harman, W. J. 1952. A taxonomic survey of the earthworms of Lincoln Parish, Louisiana. Proceedings of the Louisiana Academy of Sciences 15:19-23.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 104
Harper, E. H. 1905. Reactions to light and mechanical stimulation in the earthworm Perichaeta burmudensis (Beddard). Biological Bulletin 10:17-34.
Hatai, S. 1930. Note on Pheretima agrestis (Goto and Hatai), together with the description of four new species of the genus Pheretima. Science Report Tohoku Imperial University, Series 4. 5:651-667.
Horst, R. 1893. Earthworms from the Malay Archipelago. _In Weber, M., Zoologische Ergebnisse elner Reise in NiederlRndisch Ost- Indien 3:28-77.
Howell, G. D. 1939. The responses to light in the earthworm Pheretima agrestis Goto and Hatai, with special reference to the function of the nervous system. Journal of Experimental Zoology 81(2):231-259.
Kinberg, J. G. H. 1866. Annulata nova. Ofversigt af Kongliga Vetenskaps-Akademiens Forhandlingar, Stockholm 23:97-102.
Kindred, J. E. 1929. The leucocytes and leucocytopoietic organs of an oligochaete, Pheretima indiea (Horst). Journal of Morphology 47:435-478.
Kobayashi, S. 1937, On the breeding habit of the earthworms without male pores. Science Report Tohoku Imperial University, Series 4. 11:473-485.
______.1938. Earthworms of Korea. I. Science Report Tohoku Imperial University, Series 4. 13:89-170.
Liebmann, E. 1942. The correlation between sexual reproduction and regeneration in a series of Oligochaeta. Journal of Experimental Zoology 91:373-389.
Michaelsen, W. 1892. Terricolen der Berliner Zoologischen Sammlung. II. Archiv fUr Naturgeschichte 58:209-261.
_ .1894. Die Regenwurm-Fauna von Florida und Georgia. Zoblogische Jahrblicher, Abtheilung für Systematik, Geographic und Biologie der Thiere, Jena 8(2):177-194.
_ .1895. Zur Kenntnis der Oligochaeten. Abhandungen aus dem Gebiete der Naturwissenschaften herausgegeben vom n a tu r w is se n s c h a ftlic h e n V erin in Hamburg 13:1-37.
.1899a. Revision der Kinberg'schen Oligochaeten-Typen. Ofversigt af Kongliga Vetenskaps-Akademiens Forhandlingar, Stockholm 56(5):413-448.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 105
.1899b. Terricolen von verschiedenen Gebleten der Erde, M itteilu n g en aus dem N a tu rh isto risc h e n Museum in Hamburg 16; 1- 1 2 2 .
.1900a. Oligochaeta. Das Tierreich. Berlin; R. Friedlhnder & Sohn, 10;1-575.
.1900b. Zur Nomenclâtur der Oligochaeten, eine Rechtfertigung. Zoologischer Anzeiger 23(627);566-568.
.1903. Die geographische Verbreitung der Oligochaeten. Berlin; R. Friedlhnder & Sohn, 186 pp.
.1907. Oligochaeta. In Michaelsen, W., and H. Hartmeyer. Die Fauna SUdwest-Australiens, Jena 1(2); 117-232.
.1928. Die Oligoch&ten Borneos. Arkiv Fbr Zoologi 20A(3) ; 1-60.
.1934. Oligochaeta from Sarawak. Quarterly Journal of Microscopical Science 77(1);1-48.
Ohfuchi, S. 1937. Descriptions of three new species of the genus Pheretima from north-eastern Honshu, Japan. Research Bulletin S a ito Ho-on K ai Museum S en d a i, Japan 12;13-29.
______.1939. On the variability of the position and the number of the genital papillae and on their histological structure in Japanese Pheretima. (In Japanese, with English summary); Volumen Jubilare Pro Professore Sadao Yoshida, Osaka Natural History Society, Osaka Imperial University 1;392-407.
Olsen, H. W. 1928. The earthworms of Ohio. Bulletin of the Ohio Biological Survey 4(2);47-90.
Righi, G., and C. Knepper. 1965. 0 genero "Pheretima" Kinberg no estado do Rio Grande do sul (Oligochaeta, Megascolecidae). Revista Brasileira de Biologia, Rio de Janerio 25(4);419-427.
Rosa, D. 1891. Die exotischen Terricolen des k.k. naturhistorischen Hofmuseums. Annalen des K.K. naturhistorischen Hoffmuseums, Wien 6;396-397.
Schmarda, L. K. 1861. Neue wirbellose thiere, beobachtet und gesa- mmelt auf eih&r reise um die erde 1853 bis 1857 von Ludwig K. Schmarda. Leipzig; W. Engelmann, l(ii);13.
Schread, J. C. 1952. Habits and control of the oriental earthworm. Bulletin af the Connecticut Agricultural Experiment Station 556: 1-15.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 106
Sims, R. W. 1964. Oligochaeta from Ascension Island and Sierra Leone including records of Pheretima and a new species of Dichogaster. Annals and Magazine of Natural History, Series 13. 7:107-113.
Smith, F. 1915. Two new varieties of earthworms with a key to described species in Illinois. Bulletin of the Illinois State Laboratory of Natural History 10(v iii): 551-559.
______.1928. An account of changes in the earthworm fauna of Illinois and a description of one new species. Illinois Natural History Survey Bulletin 17(x): 347-362.
Stebbings, J. H. 1962. Additions to the Illinois and Missouri earthworm fauna. American Midland N aturalist. 67(2):504-505.
Stephenson, J. 1923. Oligochaeta. Fauna of British India. London; Taylor and Francis, 518 pp.
______.1930. The Oligochaeta. Oxford; Clarendon Press, 978 pp.
.1933. Oligochaeta from Australia, North Carolina, and other parts of the world. Proceedings of the Zoological Society of London 1932:899-943.
Stinauer, R. 1951. Life histories and culture of earthworms. Unpublished thesis for M.S. degree. Department of Zoology, Michigan State University, East Lansing.
Templeton, R. 1844. Description of Megascolex caeruleus. Proceedings of the Zoological Society of London 1844:89.
Tsai, Chu-Fa. 1964. On some earthworms belonging to the genus Pheretima Kinberg collected from Taipei area in North Taiwan. Quarterly Journal of the Taiwan Museum 17(l);l-35.
Ude, H. 1895. Beitrage zur Kenntnis der Enchytraeiden und Lumbriciden. Zeitschrift fur wissenschaftliche Zoblogie 61:110- 129.
.1932. Beitrage zur Kenntnis der Gattung Pheretima und ihrer geographischen Verbreitung. Archiv für Naturgeschichte 1:114- 190.
Yamaguchi, H. 1930a. On the variability of the capsulogenous glands in the earthworm (Pheretima hilgendorfi, Michaelsen). Trans actions of the Sapporo Natural History Society 11(2):89-95.
______.1930b. Preliminary report on several species of earthworms found in Sapporo, Hokkaido. (In Japanese); Dobutsugaku Zasshi, Tokyo 42(469): 49-59.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 107
.1931. On the normal position, and the variability in position and number of the male pores in Pheretima hilgendorfi Michaelsen. (In Japanese); Dobutsugaku Zasshi, Tokyo 43(511); 393-399.
.1962. On earthworms belonging to the genus Pheretima, collected from the southern part of Hokkaido. Journal of Hokkaido Gakugei University 13(1):1-21.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. CD ■ D O Q. C g Q.
"O CD
C/) 3o' O
8 "O T.OBLE 1 (O'3" i Number of Sites at which Species of Pheretima Occurred Singly and Together in Louisiana 3 CD Number o f Species 1 2 3 !,
3. X XX 3" P. agrestis X CD P. californica X X X XXXXX X XXX XX X XX ■DCD O P. diffringens X XXX XXXXX X X X X XX X XXX CQ. a P. hawayana X X XXXXX XXXXX X X X X XX X XX X o 3 T3 P. hilgendorfi XX XX O P. hupeiensis X X X XX XX XXXXX
CD Q. P . minima X X XX X XXXX
P. morrisi XX XXX XX XX X XXXXXX
274 1 8 2 6 h 2 4 1 1 4 1 14 1 1 1 1 1 1 2 6 10 1 1 1 1 1 1 1 1 T3 Number of Sites 6 5 17 5 1 19 38 5 CD I W(/) Total Number of Sites 448 o'
O 00 109
TABLE 2
Total Number of Associations of Pheretima Species at 448 Sites*
0 w Ü C U-i W O u •H C ÙÛ o CO u C C 13 q •H o •H nj C <3 td CO U-i Q) B •H •H 0£ o •H rH s r4 a C U (0 •H cd •H q •H o O B B P.| fUj P^\ P.! P.1 f^l f^i
P. agrestis (17)
P. californica - (70)
P. diffringens 9 35 (376)
P. hawayana - 56 75 (128)
P. hilgendorfi 3 - 5 1 (8)
P. hupeiensis - 7 7 8 - (15)
P. minima - 5 4 9 - 3 (10)
P. m o rris i - 23 25 33 - 7 5 (40)
Numbers in Parentheses are Total Numbers of Sites.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 110
TABLE 3
Preclitellar Genital Markings of P. diffringens
T o ta l Number o f GMs Number o f Specim ens
16 1
14 3
13 7
12 48
11 26
10 47
9 39
8 49
7 50
6 330
5 93
4 184
3 59
2 64
1 31
None 56
T o ta l 1087
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 111
TABLE 4
Preclitellar Genital Markings of _P. diffringens
Location of GMs Specimc
P re se ta l in v i - i x 71
Presetal in vii-ix 609
P re se ta l in v i i - v i i i 118
P re se ta l in v i i i - i x 146
P re se ta l in v i i i 72
Presetal in ix 10
P re se ta l in vi and viii 1
Pr^esetal in vi-ix, postsetal in v i i 1
P re se ta l in vii-ix, postsetal in v i 1
P re se ta l in vii-ix, postsetal invi-viii 1
P re se ta l in vii-viii, postsetal in vi-viii 1
Absent, (no GMs) 56
T o tal 1087
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 112
TABLE 5
Frequency of Preclitellar Genital Markings in 1031 Specimens
of P. diffringens
A BCDE F
v 0 --- - 0
v i 3 24 32 34 18 1
v i i 3 • 215 638 635 210 3
v i i i 2 225 956 940 222 2
ix 181 763 746 192
A and F Postsetal GMs of right and left sides respectively.
B and E Presetal lateral GMs of right and left sides respectively.
C and D Presetal median GMs of right and left sides respectively.
No GMs possible at this location,
v-ix Segments on which GMs are located.
Arabic numbers represent number of times the particular marking was
found at this location in 1031 specimens.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 113
TABLE 6
Preclitellar Genital Markings of P. hawayana
Number o f GMs Number o f Specim ens
None 174
1 49
2 170
3 8
4 8
T o tal 409
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 114
TABLE 7
Preclitellar Genital Markings of _P. hawayana
Location of GMs Number o f Specimens
v i 1
v i i 221
v i i - v i i i 10
v i i i 3
Absent (no GMs) 174
T o tal 409
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 115
TABLE 8
Postclitellar Genital Markings of P. hawayana
Number of GMs Number of Specimens
None 2
1 19
2 132
3 64
4 103
5 39
6 24
7 11
8 9
9 1
10 3
11 1
18 1
T o tal 409
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 116
TABLE 9
Postclitellar Genital Markings of P. hawayana
Number o f GMs on Number o f Specim ens R ig h t-L e ft
0-0 2
0-1 9
1-0 10
2-0 1
1-1 131
1-2 29
2-1 35
1-3 1
3-1 4
2-2 98
2-3 24
3-2 15
3-3 24
3-4 5
4-3 6
4-4 4
4-5 1
4-6 1
5-3 5
5-5 2
5-6 1
9-9 1
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 117
TABLE 10
Midventral Genital Markings of P. morrisi
Location of GMs Specimens
v i i 6
v i - v i i 5
v i i - v i i i 5
v - v i i 2
v i - v i i i 29
v - v i i i 2
v i - i x 1
vii and xviii 2
vii-viii and xviii 1
vi-viii and xviii 6
x v i i i 9
x ix 3
xviii-xix 1
Lack midventral GMs 39
T o tal 111
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 118
TABLE 11
Lateral Preclitellar Genital Markings of P. morrisi
Location of GMs Specimens
v i i 76
v i - v i i 8
v i i - v i i i 7
v i - v i i i 1
No preclitellar GMs 11
Midventral, but no lateral GMs 8
T o tal 111
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 119
TABLE 12
Postclitellar Genital Markings of _P. morrisi*
L o ca tio n Number o f GMs Specimens
Postsetal xvii 4 1
Postsetal xviii 1 1
2 1
5 1
P r e s e ta l x v i i i 0 40
1 11
2 17
3 10
4 18
5 10
6 4
7 1
P re s e ta l x ix 0 54
1 8
2 27
3 8
4 10
5 0
6 2
7 2
^These are in addition to the two GMs in each male porophore,
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 120
TABLE 13
Climatological Data for New Orleans
1931-1960
Mean High Low Mean T o ta l Temp. °F Temp. °F Temp, °F Precipitation in Inches
Jan . • 56.0 84 18 4.31
F eb . 58.4 85 19 4.48
Mar. 62.6 87 26 6.03
Apr. 69.1 89 35 5.18
May 75.7 98 47 4.96
June 81.2 98 58 5.76
J u ly 82.5 99 61 8.10
Aug. 82.7 100 61 6.53
S e p t. 79.4 97 53 6.24
O ct. 71.8 94 38 3.30
Nov. 61.7 86 26 3.08
D ec. 57.0 83 21 4.59
Annual 69.8 100 18
Mean Annual Mean Annual Snowfall in Precipitation In ch es in In ch es
0.2 62.56
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 121
TABLE 14
Climatological Data for Shreveport
1940-1960
Mean^ High Low Mean T o tal Temp. F Temp. °F Temp. °F Precipitation in Inches
Jan. 47.8 85 8 4.45
F eb . 51.0 84 2 3.81
Mar. 56.5 91 15 4.23
Apr. 66.0 92 35 5.25
May 73.7 99 42 5.42
June 80.9 101 55 3.37
J u ly 83.5 106 60 3.15
Aug. 83.4- 106 57 2.65
S e p t. 77.3 104 43 2.78
O ct. 67.4 97 31 3.28
Nov. 55.5 88 21 3.59
D ec. 49.0 84 14 4.27
Annual 66.0 106 2
Mean Annual Mean Annual Snowfall in Precipitation Inches in Inches
1.7 46.25
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 122
4-3 6 u rH t: m
' i % •H
a
A: FU •
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 123
&
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 124
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 125
a •H4-' Û) U O'
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 126
*î
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 127
to •H Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 128 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 129 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Figure 1. Spermatheca of Pheretima agrestis.* 1. ampulla 2. duct 3. stalk 4. seminal chamber 1 and 2. main axis 3 and 4. diverticulum Figure 2. Longitudinal section of the male porophore of Pheretima a g r e s t i s . 1. prostatic duct 2. papilla of male pore 3. parietal invagination of male porophore ^Figures 1-23 drawn with the aid of a camera lucida. Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. F ig u re 1 F ig u re 2 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Figure 3. Modified epidermis of Pheretima agrestis, 1. wrinkles in patch Figure 4. Male pore region of Pheretima agrestis. 1. male porophore 2. genital marking Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. v i i F ig u re 3 Fig u re 4 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Figure 5. Manicate Intestinal caecum of Pheretima agrestis. 1. intestinal tract 2. main caecum 3. secondary caecum Figure 6. Internai maie terminalia of the specimen of Pheretima agrestis illustrated in Figure 2 and Figure 4. 1, prostate gland 2, prostatic duct 3, vas deferens 4. genital marking gland Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. X X V 0.1 XXVI XXV xxiv x x iii F igure 5 XVI x v ii # XVlll XIX XX XXI F ig u re 6 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Figure 7. Male porophores of Pheretima californica, retracted on left and partially everted on right. Figure 8. Everted male porophores of Pheretima californica. Figure 9. Simple intestinal caecum of Pheretima californica. 1. intestinal tract 2. simple caecum Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. x v i i x v i i i F igure 7 X V I 1 I F igure 8 x x v ii XXIV F igure 9 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Figure 10. Hypothetical total battery of genital markings in Pheretima diffringens. Figure 11. Genital markings of Pheretima diffringens. Figure 12. Genital markings of Pheretima diffringens. 1. postsetal genital markings 2. lateral presetal genital markings 3, median presetal genital markings.— 4. spermathecal pores Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 40 40 43 ix v i i i v i i v i Figure 10 IX V l l l V ll VI F igure 11 "V o - o O -O . O XX V ixi v i i VI Figure 12 Reproduced with permission of the copyright owner. Further reproduction prohibited wi: without permission. Figure 13. Histology of genital marking gland of Pheretima diffringens. (Composite drawing from several slides). 1. openings of numerous ducts 2. minute ducts 3. cross section of minute ducts 4. secretory end piece 5. epidermis 6. circular muscle 7. longitudinal muscle Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. F ig u re 13 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Figure 14. Preclitellar genital markings of Pheretima hawayana. 1, genital marking Figure 15. Male pore region of Pheretima hawayana. 1. male pore 2. postclitellar genital marking Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. I VI v i l Figure 14 x v i i i \ F igure 15 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Figure 16. Longitudinal section of the male porophore of Pheretima hilgendorfi. 1. prostatic duct 2. papilla of male pore 3. parietal invagination of male porophore Figure 17. Male pore region of Pheretima hilgendorfi. 1. genital marking 2. papilla in genital marking Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. m m F igure 16 x v ii X V l l l I t F ig u re 17 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Figure 18. Internal male terminalia of the specimen of Pheretima hilgendorfi illustrated in Figures 16 and 17. 1. prostate gland 2. prostatic duct 3. vas deferens 4. genital marking gland Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 1 mm xvi x v ii x v i i i x ix XX F ig u re 18 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Figure 19. Male pore region of Pheretima hupeiensis. 1, genital marking 2. male pore Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. x v i i i F ig u re 19 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Figure 20. Spermatheca of Pheretima minima. 1. ampulla 2. duct 3. seminal chamber 4. stalk Figure 21. Internal male terminalia of Pheretima minima. 1. prostate gland 2. prostatic duct 3. vas deferens Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. -1 3 4 F ig u re 20 F ig u re 21 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. r Figure 22, Preclitellar genital markings of Pheretima m orrisi. 1. midventral genital marking 2. lateral genital marking Figure 23. Male pore region of Pheretima m orrisi. 1. genital marking in male porophore 2. male pore Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. VI v i i v i i i Figure 22 I xvix Figure 23 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 154 APPENDIX A series of each species of Pheretima in this study and slides of the genital marking glands of P. diffringens are deposited in the Louisiana State University Museum of Zoology, Invertebrate Collection. The museum numbers of th ese specimens are 595-602. Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 155 VITA Richard Elvin Tandy was born July 19, 1937, at Pontiac, Michigan. He graduated from Farmington High School, Farmington, Michigan, in 1955 and immediately entered Anderson College, Anderson, Indiana, from which he graduated with the Bachelor of Arts Degree in 1960. He was admitted as a graduate student in Zoology by Louisiana Polytechnic Institute at Ruston, Louisiana, in 1961 and was awarded the Master of Science Degree in 1963. In September, 1963, he entered the Louisiana State University Graduate School and was an Instructor in the Department of Zoology and Physiology for the 1967-1968 session. He was married in 1960 to Elva Williams of Baton Rouge, Louisiana and has a son. Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. EXAMINATION AND THESIS REPORT Candidate: Richard Elvin Tandy Major Field: Invertebrate Zoology Title of Thesis: The Earthworm Genus Pheretima Kinberg, 1866, In Louisiana (01 i gochaeta:Megasoleci dae) Approved: L a I ' Majpw Professor and Chairman Dean of the G raduate School EXAMINING COMMITTEE: Date of Elxamination: 9 September 1968 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.