69 - 16,153 KOPENSKI, Martin Leo, 1934SELECTED STUDIES OF MICHIGAN LEECHES Michigan State University, Ph.D., 1969 Zoology University Microfilms, Inc., Ann Arbor, Michigan SELECTED STUDIES OP MICHIGAN LEECHES By ■ Martin Leo Kopenski A THESIS Submitted to Michigan State University in partial fulfillment of the requirements for the degree of DOCTOR OP PHILOSOPHY Department of Zoology 1969 ABSTRACT SELECTED STUDIES OP MICHIGAN LEECHES By Martin Leo Kopenski This study is concerned with leeches present in Michigan, particularly in Marquette County waters. An attempt is made to correlate various chemical, physical and biological features of water with the abundance and distri­ bution of leeches. Leeches were taken from 50 locations in Marquette County in the Upper Peninsula of Michigan and from six each in Barry and Kalamazoo Counties in the Lower Peninsula. These leeches were collected over a period of six years (1961-1966) and were obtained from such habitats as lakes, ponds, bogs, rivers, and creeks. Tests to measure total alkalinity and pH were performed on waters from a represen­ tative number of the waters investigated. Several physical characteristics of the waters studied, such as depth, color, temperature, nature of the bottom, soil types, and whether lentic or lotic, were recorded. Certain biological factors, such as feeding habits, enemies, and reproductive habits, were noted. Nineteen species of leeches were found in Michigan during this study. Nine species are recorded in addition to the listing of Miller (1937) bringing the total number known for Michigan to 22. Most glossiphonids, piscolids, hirudids, and erpobdellids are abundant and widely dis­ tributed in the areas of the state which have been studied. Helobdella stagnalis was the most abundant leech found in Marquette County, 103 individuals having been taken, and Erpobdella punctata was encountered at 3 studied. of the sites The leech fauna of the Upper and Lower Peninsulas are similar and the species in Michigan compare closely to those in Ohio, Illinois, Minnesota, and Wisconsin. The glossiphonids have been found to be more common in waters with a high total alkalinity. Most hirudids were found most commonly in waters with low total alkalinities, while erpobdellids vary. The abundance and distribution of most leeches are restricted by waters with low pH; however, leeches appear not to be restricted by waters with high pH readings. Most leeches in Michigan prefer standing waters, with the glossiphonids being best adapted for living in running waters. Water temperature is a major limiting factor for the distributuion of only Nephelopsis obscura and Philobdella gracilis. Most leeches were found along a variety of bottom types with no preference shown. The presence or absence of submerged objects is a more important factor in their dis­ tribution than the nature of the bottom. The soils sur­ rounding a body of water probably exert an indirect effect on the abundance and distribution of leeches by altering the chemical nature of the water which may affect the abundance and distribution of the leeches' food organisms. The reproductive habits of leeches undoubtedly in­ fluence their abundance and distribution. The data indicate that the glossiphonids are the most prolific leeches in Michigan and consequently the most abundant and widely dis­ tributed. The erpobdellids are a less prolific group; how­ ever, their abundance and distribution are comparable to that of the glossiphonids. Predators probably exert some influence on the abundance of erpobdellids. The abundance and distribution of food organisms are undoubtedly the most important factors affecting the abundance and distribution of leeches. ACKNOWLEDGMENTS I wish to express my appreciation to Dr. T. Wayne Porter, chairman of my committee, and assistance in this work. for his unselfish guidance I wish also to thank Dr. Wilbert Wade, Dr. Marvin Hensley, and Dr. Robert Ball for their assistance and guidance. Special thanks go to Dr. William Robinson for the many hours he spent reading over and commenting on this work. I wish to express my thanks to other members of the Biology Department of Northern Michigan University for their helpful suggestions throughout this, study. Dr. Mark Keith and Dr. Marvin Meyer were helpful in confirming several identifications of specimens. To them I wish to express my thanks. I wish to acknowledge the help of Dr. K. H. Mann for providing me with the English translation of Sandner's (1951) work and also his reprints and expert advice. Thanks go to Mr. John Farrell and assistants of the Geography Department of Northern Michigan University, who assisted me in the construction of the maps and graphs in this report. I wish to express my thanks to other investigators of leeches for providing me with reprints of their work. I wish to thank my wife for her helpful suggestions and the proofreading of this work. ii TABLE OF CONTENTS Page LIST OF TABLES v LIST OF FIGURES vii CHAPTER I. II. INTRODUCTION AND LITERATURE REVIEW 1 METHODS AND MATERIALS 5 LImnological Techniques 5 Chemical Methods Physical Methods Biological Data Leeches 6 Collecting Methods Processing and Preserving Identification Cataloging III. THE STUDY AREAS 11 Introduction 11 Marquette County 13 Geologic-Pedologic Description Habitat Descriptions Barry and Kalamazoo Counties 46 Geologic-Pedologic Description Habitat Descriptions IV. RESULTS AND DISCUSSION 60 Abundance 60 Geographical Distribution 65 Ecological Factors Related to Leech Distribution Chemical Factors Total Alkalinity pH iii 78 CHAPTER Page IV (cont'd . ) Physical Factors Lentic versus Lotic Waters Water Temperature Water Color Water Depth Bottom Composition Soils Biological Factors Feeding Habits Parasites and Predators Reproduction V. SUMMARY AND CONCLUSIONS 92 105 127 SELECTED BIBLIOGRAPHY 131 APPENDIX 135 iv LIST OP TABLES TABLE 1. 2. 3. Page Physical Features of the Waters Investigated in Marquette County 37 Floral Inhabitants of the Waters Studied in Marquette County 40 Faunal Inhabitants of the Waters Studied in Marquette County H3 Physical Features of the Waters Investigated in Barry and Kalamazoo Counties 5. 6. 7. 8. 9. 10. 57 Floral Inhabitants of the Waters Studied in Barry and Kalamazoo Counties 58 Faunal Inhabitants of the Waters Studied in Barry and Kalamazoo Counties 59 A Comparison of Leeches Collected in the Upper and Lower Peninsulas of Michigan 72 A Comparison of Leeches Collected in Five Midwestern States J6 The Water Temperature Range Found for Eleven Species of Leeches 97 The Percentage of Occurrence of Eleven Species of Leeches along Four Types of Water Bottoms 102 11. Foods of Michigan Leeches 106 Al. A Taxonomic Classification of Michigan Leeches 135 A2. A Key to the Leeches of Michigan 137 A3. Marquette County Stations, Their Locations, Dates Sampled, Chemistry, Species and Number Collected 143 Barry County Stations, Their Locations, Dates Sampled, Chemistry, Species and Number Collected 151 A4. TABLE A5. Kalamazoo County Stations, Their Locations, Dates Sampled, Chemistry, Species and Number Collected A6. Habitats Compared In Regard to Total Alkalinity A7. Total Alkalinity Mean, Standard Deviation, and Range for Eleven Species of Leeches A8. Waters Tested and Their pH A9. pH Mean, Standard Deviation, and Range for Eleven Species of Leeches A10. The Occurrence of Eleven Species of Leeches Along Various Bottoms LIST OP FIGURES FIGURE 1. Page Upper and Lower Peninsulas of Michigan showing the location of the three counties involved in this study 12 Locations and soils of the waters studied (Marquette County) 15 Locations and soils of the waters studied (Barry and Kalamazoo Counties) 49 Total number of individual leeches collected from fifty sites in Marquette County 61 Percentage of occurrence of leeches collected from fifty sites in Marquette County 63 Locations and soils of the waters studied (Marquette County) 66 7. Geographical Distribution of the Glossiphonidae 67 8. Geographical Distribution of the Piscicolidae 68 9. Geographical Distribution of the Hirudidae 69 10. Geographical Distribution of the Erpobdellidae 70 11. Percentage of occurrence of eleven species of leeches at different alkalinities 79 Occurrence of eighteen species of leeches in relation to the pH of water 87 Percentage of occurrence of eleven species of leeches in thirty-nine lentic and ten lotic waters in Marquette County 93 2. 3. 5. 6. 12. 13. Al. The eye patterns of several leeches A2. Structure of leeches l4l ■142 vii CHAPTER I INTRODUCTION AND LITERATURE REVIEW Introduction Since no comprehensive study of the ecology of leeches has been performed in the United States, the present study was undertaken to determine what leeches are present in selected counties of Michigan and to describe their abundance and distribution. A comparison was made between the species inhabiting the waters of both peninsulas of Michigan. The species of leeches in Michigan were also compared with those of Ohio, Illinois, Minnesota, and Wisconsin since species listings have been compiled for these states. Several chemical, physical, and biological features of the habitats studied were considered in an attempt to determine the most important factors regulating leech abundance and distribution. Since the relationship of these factors to leech abundance and distribution have been extensively studied only in Europe, a comparison was made between the results of this study and those of the European studies. Literature Review The major leech studies completed in the United States have been mainly concerned with species compilations 2 for various states along with species descriptions and keys to their identification. Most of these works have been performed in the midwestern states. Nachtrieb, Hemingway, and J. P. Moore (1912) conducted an extensive survey of the leech fauna of Minnesota which has recently been extended by Keith (1954, i960). J. P. Moore (1901) compiled a species listing for the state of Illinois. Bere (1931) studied Wisconsin leeches and compiled a species listing for that state. Sapkarev (1967) did a quantitative study of the leeches of Lake Mendota to determine their vertical and horizontal distribution and seasonal varia­ tions in population density. Miller (1929) studied the leeches of Ohio and in (1937) worked on Michigan leeches. The works of Mullen (1926) and Mather (1963) concern the leeches of Iowa. The only- s-ignificant study concerned with leeches of Michigan is that of Miller (1937)- The leeches referred to in his study were secured from the Museum of Zoology of the University of Michigan, having been collected in Michigan in 1925, 1926, and 1927 by Dr. Y. Metzelaar and assistants of the University of Michigan. The majority of the collec­ ting sites listed in this paper are in the Lower Peninsula of Michigan with a few from Luce County in the Upper Penin­ sula. Miller lists thirteen species of leeches found in Michigan and includes a key to their identification. Only four major ecological studies have been com­ pleted on leeches. These have been performed by Pawlowski (1936), Bennike (19^3), Sandner (1951), and Mann (1955). According to Mann (1955), Pawlowski (1936) collected from 57 stations In the Polish Lake Wigryseen and In neighboring smaller bodies of water. First, he recorded information about the food habits of leeches; secondly, he related the leech fauna to the type of substratum; and thirdly, he classified his habitats as eutrophic, dystrophic, or oligotrophic, and considered which species of leeches occurred in each type. Bennike (19^3) collected from 215 habitats scattered over the whole of Denmark and investigated leech distribu­ tion mainly from the physical and chemical standpoints. He measured certain selected characteristics of the water and related these to the species present. He divided his habitats into lakes, ponds, bogs, and streams and described the leeches inhabiting each. Sandner (1951) made an extensive survey in the county of Lodz in Poland, classifying his habitats into tarns, small ponds, fish-ponds, peat-pools, etc. He also made a number of physical and chemical observations on the water and attempted to relate these to the presence or absence of certain species of leeches. Mann (1955) collected from 29 stations in Berkshire, and 29 in the lake district of Britain. He performed a number of physical and chemical tests on the water and classified his standing waters as oligotrophic, eutrophic, and dystrophic, and his running waters on the basis of current speed. He also used quantitative collecting methods by collecting at a given site for a certain period of time and he measured the area of his habitats. He pre­ sented data to show that the area of a given body of water has an effect on the species of leeches inhabiting such a body of water. Mann (1962) states that only in Europe has the ecology of leeches been studied intensively. Information for the rest of the leech fauna of the world consists of brief notes about food organisms with which the leech is associated and perhaps some information about the habitats in which specimens have been found. CHAPTER II METHODS AND MATERIALS Limnological Techniques Chemical M e t h o d s : Chemical water analyses were per­ formed at a representative number of the sites visited. The water tested was taken at the location of the leeches present. Two chemical characteristics of the water were measured, (1) pH and (2) total alkalinity. 1. pH: The pH of the water was measured in the laboratory by means of a Beckman pH meter calibrated m to correct for temperature change. The pH was determined immediately after returning from the field not later than two hours after it had been ob­ tained. 2. Total Alkalinity: The total alkalinity of the water was determined at the collecting site as soon as the water sample was obtained. The determination was made by adding four drops of phenolphthalein in­ dicator plus two drops of methyl orange to a 100 milliliter sample and titrating with 0.2N until the solution turned salmon pink. The number of milliliters of 0.2N I^SO^ used times 10 repre­ sented the total alkalinity in mg./L. Welch (1948). 5 CaCO^, Physical Me t h o d s : habitats were recorded. Several physical features of the The physical conditions described are those which existed at the exact location where speci­ mens were obtained. 1. Bottom Composition; The bottom descriptions included are based on the bottom classifications system of Roelofs 2. Depth; (1944). The depth was measured by means of a yard stick. 3. C ol or : The water color descriptions included are based on gross observations as no color analyses were performed. 4. Temperature: The water temperature was measured by means of a Centigrade thermometer. 5. Lentic vs. L o t i c : Leeches were collected from standing and running waters and the frequency of species inhabiting both types was compared. Biological D a t a : The dominant floral and faunal in­ habitants associated with the leeches collected at each site were recorded. Leeches Collecting M e t ho ds : The leeches referred to in this study were taken in the following ways: 1. Examination of the undersurfaces of submerged rocks, sticks, boards, logs, cans, and other sub­ merged objects in the body of water. 7 2. Agitation of the water and netting free- swimming leeches which were disturbed by such agi­ tation . 3. Examination of vertebrate hosts, such as, fish, frogs, and turtles which may harbor leeches. H. Uprooting aquatic plants and examining their stems, roots, and leaves for adhering leeches. 5. Scooping up debris from the bottom, such as, submerged leaves, algae, and organic sediment. This material was sometimes examined by picking through it with forceps or placing it on dry land for a period of time. As the material dried, leeches were picked up as they left it. Processing and Preserving: The leeches collected were taken to the laboratory in the living state. A label denoting the date and location of the collection site was inserted into each jar containing leeches. In the labora­ tory, the leeches were anesthetized by immersing them in a saturated solution of menthol. When the leeches would no longer respond to prodding, they were removed from the menthol solution. The mucus secreted by them during the preceding process was washed off. The erpbobdellid leeches were preserved internally by injecting 70# ethanol into the body cavity. This was done to insure preservation of the reproductive organs which might be necessary for positive identification. Next, the leeches were straightened and placed between two glass plates. A 8 rubber band was wrapped around the glass plates with suf­ ficient pressure to keep the leeches In an extended position without displacing or distorting the Internal This preparation was Immersed In a 10 % formalin organs. solution to fix the leeches In this position. After the leeches were sufficiently hardened so as to remain in this extended position, they were removed,from the formalin solution and placed in eight-dram vials containing 70% ethanol. Identifica~tlon: The identity of the leeches was determined by using one of several keys listed in the bibliography. One of the main characteristics used to identify these specimens was the number, shape, and posi­ tion of the eyes. If the eyes were not clearly visible due to heavy pigmentation in the head region, the head portion of the leech was immersed in a 5$ solution of potassium hydroxide to bleach out the pigments, thus ren­ dering the eyes more apparent, Mann (1962). In the case of the erpobdellids, it was necessary to dissect out the reproductive organs in some cases in order to make posi­ tive identifications. In regard to Haemopis spp. the presence or absence of jaws plus the number of teeth on the jaws were used in determination of species. After the leeches were determined, the members of one species from a single collection were placed in a vial. A tag designating the date of collection, locality, locus, county, state, catalog number, and collector was inserted into each vial. 9 A taxonomic classification of Michigan leeches is given in Table Al, pages 135 and 1 3 6 . A key to the identification of these leeches is included in Table A 2 , pages 137-140. A listing of the station location, dates on which col­ lecting was done, the species and number of each found, as well as the pH and total alkalinities of the waters studied is shown in Tables A3, A4, and A5, pages 143-152. Cataloging; The cataloging system used for the pre­ served leeches is as follows: Roman numerals were used to designate the collecting sites beginning with the first collecting site visited and proceeding to the most recent. If a site was visited more than once, the same Roman numeral was used to designate the site. Capital letters were used to designate the family to which a certain species belonged, Arabic numbers the genus, and small letters the species. Thus Redberry Pond, the first site studied, would be indi­ cated by IAia. A would represent the family Glossiphonidae, 1 the genus Glossiphonia, and a the species complanata. Leeches collected in Barry and Kalamazoo Counties were cataloged separately by- county in the same manner as described. 10 Catalog Code to Leech Collection Capital Letters = Family A Glossiphonidae B Erpobdellidae C D Hirudidae Piscicolidae Arabic Numbers = Genera 1 Glossiphon!a 7 Macrobdeiia 2 Helobdella 8 Haemopis 3 Placobdella 9 Therornyzon 11 Erpobdella 10 Illinobdella 5 Dina 11 Philobdella 6 Nephelopsis 12 Piscicola Small Letters = Species Species Genera Species Genera Glossiphonla a complanata Nephelopsis a obscura Helobdella a stagnalis Macrobdeiia a decora b fusea Haemopis a marmorata a hollensis b grandis b parasitica Therornyzon a rude c rugosa Illinobdella a alba d montifera b punctata Erpobdella a punctata Philobdella a gracilis Dina a parva Piscicola a milneri b fervida Placobdella CHAPTER III THE STUDY AREAS Introduction Fifty collecting stations were selected representing such habitats as: lakes, ponds, bogs, rivers, and creeks located throughout Marquette County. Leeches were also collected from six sites in Barry County and six sites in Kalamazoo County in the Lower Peninsula of Michigan. These leeches were collected from habitats comparable to those in Marquette County. The names used to designate collecting stations are those used on various maps of the areas. Where no known name could be found for a body of water, a name was derived in relation to a road or some permanent structure In the area. Figure 1 depicts the three counties involved in this study. Figure 2, page 15, shows the location and soil relationships of the waters studied in Marquette County. Only the soil types immediately surrounding the waters studied are shown in.Figure 2. Table 1, pages 37-39, lists several physical features of the waters investigated in Marquette County. Tables 2, pages 40-42 and 3, pages 43-45 , denote the dominant floral and faunal inhabitants of the waters studied in Marquette County. 11 12 MARQUETTE BARRY KALAMAZOO Fig. 1. Upper and Lower Peninsulas of Michigan showing the location of the three counties involved in this study. 13 Likewise, Tables 4, 5, and 6, pages 57-59, de­ pict corresponding Information for the waters of Barry and Kalamazoo Counties. Marquette County Geologic-Pedologic Description; The following geologic-pedologic description of Marquette County and the specific study areas is taken from Veatch (1953, pages l4l142). The Marquette area is a complex of rock knobs and deep-drift highland masses. Swamps and bogs in large and small basins are inter­ spersed; and narrow swampy valleys form a network pattern. Other features of the landscape are: both large and small lakes, diverse in origin, shape-y-and kind of shore lines, beaver meadows; enclosed dry and wet valley plains and basins, floored with sand, gravel, and boulders. Streams are relatively numerous, have eccentric courses, and flow through swampy valleys and m ea dows. The dominant soils of the highland are reddish, loams or silts in texture, and over­ lie coarse glacial drift characterized by small fragments and boulders of basalts, gabbros, diorites, and a variety of other rocks, especially granites, schists, slates, and red sandstones. The soils on the dry plains are mainly Stambaugh and Rubicon types. Much of the present bare rock surface on crests of knobs and hills was originally covered by a blackish, spongy humous soil, and brown, silty loam which was lost through fire and rain-wash after lumbering. The swamp soils are mainly raw, highly acid, peats of the Greenwood and Spalding types, with less of the darker more decomposed Carbondale peat. 14 Figure 2 Legend I. Red Drift. Bedrock basic igneous and granites. Podzol profiles, both weak and strong development of gray and brown horizons. II. Largely gray drift, high proportion of slate rocks. Podzol profiles have buff and yellow orterde horizons. Brown podzols common. III. Red Drift. Mostly granite rock, some sandstone and basic igneous rocks. Mostly normal podzol profiles including lithosols and brown podzols. IV. Red Drift. Mostly igneous and metamorphic rocks in drift, fairly strong influence from sand­ stones; very slight or none from limestone. Podzol soil profiles; strongly developed brown horizons, except some dry sands which have only weak develop­ ment and absence of a lower red clay. V. Fluvio-glacial and outwash deposits. Little or no limestone influence. Podzol soil profiles, variably weak and very strong development of gray and brown horizons. VI. Clayey Drift, red color; influence from sandstones, igneous and metamorphic rocks, little or no influ­ ence from limestone. Podzol profiles; variably weak, to strong development of gray and brown horizons. VII. Outwash, glacio-fluvial and lacustrine deposits, reddish color. No limestone influence, podzol pro­ files weakly or incompletely developed. VIII. Mostly sand and gravel. Podzols either gray horizon or brown horizon separately strongly developed. Little or no limestone influence. IX. Coarse red drift, hard compact sandy clay. Drift strongly influenced by local sandstone bedrock. (from Veatch, Soils and Land of Michigan) 15 COLLECTING SITES MARQUETTE COUNTY, MICHIGAN 34! MILES 44, l«T 33 32 124 '43 -o R 30 W RIIW 1.REDBERRY PONO 2.HARLOW LAKE 3.WALDO POND 4.SAUXHEAD LAKE 5.INDEPENDENCE LAKE 6.KAWBAWGAM LAKE 7.ROAD 510 POND 8.MANGUM ROAD DITCH 9.BAGDAD POND IQCEMETERY POND 11.RUSH LAKE I2H0WE LAKE Fig. 2. 13.MINE SHAFT POND 14. ROAD 553 CREEK 15. HORSESHOE POND 16. TEAL LAKE 17. CRANBERRY BOG 18.DEAD RIVER 19. SMALL PONDS 20.VERNAL POND 21.SMALL CREEK 22.C00PER LAKE 23.BANCROFT LAKE 24.B0ST0N LAKE 25 TILDEN LAKE 4 26 W 26.SCHOOLHOUSE LAKE 27. OGDEN LAKE 28. FISH LAKE 29.BACON LAKE 30.MORGAN CREEK 31.RAMSETH POND 32.HUMBOLDT POND 33.LAKE SUPERIOR 34.M0UNTAIN LAKE 35.PINE LAKE 36.GOOSE LAKE 37.R0D B GUN CLUB POND 38. WETMORE POND R 24W 39.WEST BRANCH CREEK 40.TOURIST PARK POND 41. DEAD RIVER 42.MINE SHAFT POND 43.CARP RIVER 44.DISHN0 CREEK 45.ARFELIN LAKE 46.DEER LAKE 47.BARNHARDT CREEK 48.GOLDMINE CREEK 49.MID0LE ISLE POINT BOG 50.CH0C0LAY RIVER Locations and soils of the waters studied. 16 Habitat Descriptions (Marquette County): Tables 1, 2, and 3 on pages 37-^5 denote several physical and biological characteristics of the waters investigated. The following habitat descriptions pertain mainly to the specific locations at which leeches were found with some information about the body of water as a whole. The criteria used to distinguish between lakes, ponds, rivers, bogs, creeks, and ditches are denoted. LAKES "Bodies of water having an area of open, rela­ tively deep water sufficiently large to produce somewhere on its periphery a barren, wave swept shore." (Welch, 1952, page 16) Station 2. Harlow Lake derives its water supply from several small streams and occupies an area of 75 acres. . It drains into Lake Superior by way of Harlow Creek. The bottom in the littoral zone consists of sand with small interspersed graveled patches. The bottom in the limnetic region is made up of fibrous and pulpy peat. An abundance and variety of aquatic plants are present con­ sisting mainly of sedges (Scirpus spp.) in the lit­ toral zone with pondweeds (Fotamogeton Richardsonii) and (P. Robbinsii) mainly dominating the limnetic area. A wide variety of invertebrates are present with clams (Anodonta sp.) and snails (Helisbma sp.) predominating. present. Few crayfish (Cambarus sp.) are also Yellow perch (P'erca flavescens) is the dominant species of fish with bluegills (Lepomis macrochirus) and northern pike (Esox lucius) also present. Progs (Rana spp.) occur in the littoral zone while snapping turtles (Chelydra serpentina) and painted turtles~*(Chrysemys p i c t a ) are present throughout the lake. Station k. Sauxhead Lake is fed by the Big Garlic River and drains into Lake Superior by way of this river. The bottom is mainly sand with pieces of submerged wood being abundant. Sedges (Scirpus spp.) dominate the littoral zone with yellow water lilies advena) and pondweeds (Nuphar (Potamogeton spp.) being prevalent in the limnetic waters. Clams (Elliptio sp.), snails (Campeloma sp.), and crayfish (Orconectes sp.) are the dominant invertebrates with the vertebrates represented by yellow perch (Perea flavescens), northern pike (Esox lucius), and walleyes (Stizostedion vitreum). Station 5. Independence Lake occupies an area of 1,860 acres, is fed mainly by the Yellow Dog River and drains into Lake Superior by way of the Iron River. The bottom consists of sand with patches of rubble and submerged wood. Pondweeds (Potamogeton spp.) are the dominant plants while clams and amphipods (Pisidium sp.) (Hyalella azteca) are abundant. Yellow perch (Perea flavescens) , northern pike (Esox lucius), and walleyes (Stizostedion vitreum) are the dominant vertebrates. Station 6. Kawbawgam Lake occupies an area of 153 acres, possesses mainly a sand bottom, and drains into Lake Superior by way of the Chocolay River. Pondweeds (Potamogeton spp.) are the dominant plants while snails (Physa sp.) and isopods (Asellus sp.) are abundant. Yellow perch (Perea flavescens) and northern pike (Esox lucius) are present in sizeable n um be rs . Station 11. Rush Lake is a clear deep oligotrophic lake with mainly a sand bottom with areas of overlying rubble and sticks. by rushes The littoral zone is occupied (Juncus sp.); no higher aquatic plants were visible in the limnetic area. Clams (Pisidium sp.) and crayfish (Orconectes sp.) are abundant with the dominant vertebrates being salmonids. Station 12. Howe Lake^ located near Rush Lake, resembles it in appearance and composition. The bottom is mainly sand covered with a 2-inch layer of silt, with an abundance of submerged sticks and intermittent accumulations of rubble. Rushes (Juncus sp.) dominate the littoral zone. The limnetic waters are deep with no higher aquatic plants evident. — - Clams (Pisidium sp.), crayfish (Orconectes sp.), and amphipods (Hyalella sp.) are present in small numbers with salmonids being the dominant vertebrates. Station 16. Teal Lake occupies an area of 505 acres. The bottom is composed of bedrock with overlying sand in the deeper areas. Stonewort (Chara sp.) is the dominant plant in the littoral zone with pondweeds - (Potamogeton spp.) dominating the limnetic waters. Planarians (Dugesia sp.) and crayfish (Orconectes sp.) are very abundant with chironomids sp.) present in smaller numbers. (Chironomus Yellow perch (Perea flavescens), pumpkinseeds (Lepomis gibbosus), and walleyes (Stizostedion vitreum) are the dominant fish. Station 22. Cooper Lake occupies an area of 3^ acres and is surrounded mostly by woodland with a small area of pasture land on the south shore. The bottom in the littoral zone is composed of sand with an abundance of submerged sticks and a few boulders. The bottom in the limnetic area is composed of fibrous and pulpy peat. Rushes (Juncus spp.) are abundant in the littoral zone with pondweeds (Potamogeton spp.), yellow water lilies (Nuphar adv en a), and bushy pondweed (Najas flexilis) dominating the limnetic waters. Amphipods (Hyalella sp.) and aquatic insects mainly gyrinids sp.) are abundant. and bullheads fish. (Gyrinus Yellow perch (Perea flavescens) (Ictalurus sp.) are the representative 20 Station 23. Bancroft Lake occupies an area of 28.5 acres. The bottom consists of pulpy peat covered with pieces of submerged concrete at the site of collection. Aquatic plants are sparse being repre­ sented by patches of pondweeds (Potamogeton spp.). Snails (Physa sp.) are very abundant with amphipods (Hyalella sp.) and crayfish (Orconectes sp.) being present in fewer numbers. Bullheads (Ictalurus sp.) are the dominant fish along with a few cyprinids. Station 2k. Boston Lake is a spring fed lake occupying an area of 50.5 acres. The bottom in the littoral zone consists of sand with intermittent accumulations of rubble. The bottom in the limnetic zone consists of pulpy peat. pondweeds Yellow water lilies (Nuphar advena) and (Potamogeton spp.) are abundant. Clams (Sphaerium sp.), snails (Helisoma sp.), and crayfish (Cambarus sp.) are the dominant invertebrates. White suckers (Catostomus commersonnii) are present in large numbers, while black crappies (Pomoxis nigromaculatus) are present in fewer numbers. Station 25. Tilden Lake occupies an area of 53 acres and reaches a maximum depth of 37 feet. The bottom con­ sists of sand, rubble, and numerous submerged logs. The entire shoreline of the lake is heavily wooded. Arrowhead (Saglttaria latifolia) and sedges spp.) are present but not abundant. (Scirpus Crayfish (Cambarus sp.) is the dominant invertebrate. 21 Salmonids are the dominant fish along with a wide variety of cyprinids. Station 26. Schoolhouse Lake Is a shallow lake surrounded by wooded hills. The bottom consists of muck with a 6-inch layer of hemlock (Tsuga canadensis) bark overlying it in the littoral zone. Bur reed (Sparganium androcladum) and arrowhead (Sagittaria latifolia) are abundant in the littoral zone with yellow water lilies limnetic waters. (Nuphar adyena) dominating the Planarians (Hyalella sp.), and snails abundant. (Dugesia sp.), amphipods (Physa sp.) are very No representative vertebrates were observed. Station 27. Ogden Lake is an elongated, shallow lake. The bottom consists of pulpy peat overlain with numerous sticks and chips of wood. americana), pondweeds Eel grass (Vallisneria (Potamogeton spp.), and yellow water lilies (Nuphar advena) are the dominant plants although they are not abundant. sp.), snails (Dugesia (Physa sp.), and crayfish (Orconectes sp.) are numerous. Station 28. Planarians No vertebrates were observed. Pish Lake occupies an area of 156 acres and possesses a sand bottom with numerous patches of overlying gravel. and horsetails Arrowhead (Sagittaria latifolia) (Equiselum sp.) crowd the littoral zone of the lake. Yellow water lily (Nuphar advena) is the dominant plant present in the limnetic water, 22 although it Is not abundant. Snails (Campeloma sp.), clams (Sphaerlum sp.), and amphlpods are abundant. (Hyalella sp.) The fish present In order of abundance are yellow perch (Perea flavescens), black crappies (Pomoxls nigromaculatus), bluegills (Lepomls macro- chlrus), and northern pike (Esox lucius) . Station 29. Bacon Lake fits In the category between lake and pond. It Is a shallow lake reaching a maximum depth of 15 feet. The bottom consists of sand over- lain with a 3-4 inch layer of fibrous peat. Bur reed (Sparganium androcladum) and bushy pondweed (Najas flexilis) are abundant. Snails (Physa sp.) and culicine larvae (Chaoborus sp.) occur in small numbers. Station 33. Cyprinids are the dominant vertebrates. The portion of Lake Superior examined is a bayou altered considerably by man for the purpose of boat storage and launching. The bottom is composed of sand with pieces of overlain concrete, also bricks, rubble, and boulders. Waterweed (Anacharis canadensis) is the only aquatic plant in this area and is not abundant. Snails (Campeloma sp.) and (Physa sp.) are abundant along with isopods (Asellus sp.). Fish are scarce in this area although fish from t h e •lake proper sometimes enter into this bayou. Station 34. _ . Mountain Lake is a clear, deep, oligotrophic lake with a sand bottom overlain with gravel and 23 intermittent pieces of wood. Sedges (Scirpus spp.) dominate the littoral zone of the lake. The lim­ netic waters are devoid of visible aquatic plants due to the great depth. amphipods Snails (Physa sp.) and (Hyalella sp.), as well as salmonids, are abundant. Station 35. Pine Lake is a deep oligotrophic lake with a sand bottom overlain with intermittent sticks. The littoral zone is crowded with sedges (Scirpus spp.) while pondweeds (Potamogeton spp.) dominate the limnetic zone. Snails (Physa sp.), clams (Pisidium sp.), and amphipods (Hyalella sp.) are the predomi­ nant invertebrates. This lake, along with Howe, Rush, and Mountain Lakes previously described, is located on the property of the Huron Mountain Club and is not accessible to the general public. Conse­ quently, these four lakes have not been subjected to the ravages of man and are possibly the least modi­ fied and in the author’s estimation the most beauti­ ful lakes in the State of Michigan. Station 3 6 . Goose Lake occupies an area of 446 acres and possesses a sand bottom covered with a thin layer of silt. This lake in past years was a receptacle for the sewage wastes of the City of Negaunee. This practice was discontinued ten years ago; however, extensive algal blooms still occur during the summer. Yellow water lilies (Nuphar advena) and pondweeds 24 (Potamogeton spp.) are present in large numbers. Copepods (Cyclops sp.) and cladocerans are exceedingly numerous. (Daphnia sp.) Yellow perch (Perea flavescens) and northern pike (Esox lucius) are the dominant vertebrates. Station 45. Arfelin Lake occupies an area of 66.5 acres, reaches a maximum depth of 35 feet, and drains into the Peshekee River. The bottom in the littoral zone consists of sand covered with gravel and submerged sticks. Pondweeds small numbers. (Potamogeton spp.) are present in Sponges (Spongilla spp.) are abundant adhering to submerged sticks. Planarians (Dugesia sp.), snails (Ferrisia sp.), and the chironomid (Chironomus sp.) are also numerous. fish are catostomids, Station 46. The dominant salmonids, and cyprinids. Deer Lake is a body of water containing many islands, bayous, inlets, and outlets. The bottom consists of sand with few submerged rocks. Blue green algal blooms of Anabaena sp. and Microcystis sp. occur frequently and there is a lush growth of pondweeds m i n o r ). (Potamogeton spp.) and duckweed (Lemna Oligochaetes and chironomids (Chironomus sp.) are extremely numerous while snails (Physa sp.) and amphipods numbers. (Hyalella sp.) are present in lesser Yellow perch (Perea flavescens) are the dominant fish. 25 PONDS "Very small, very shallow bodies of standing water In which quiet water and extensive occupancy by higher aquatic plants are common characteristics." (Welch, 1952, page 16) Station 1. Redberry Pond Is spring fed and reaches a maximum depth of ten feet. The bottom consists of fibrous peat overlain with numerous submerged l o g s . Yellow water lilies (Nuphar advena) are abundant and occupy the greater portion of the pond during the summer months. Snails (Physa sp.), amphipods (Hyalella sp.), and aquatic insects mainly dragon fly and damsel fly naiads are abundant. Yellow perch (Perea flavescens) are present in small numbers. Painted turtles (Chrysemys picta belli) are the pre­ dominant vertebrates with small populations of redspotted newts frogs Station 3. (Notopthalmus virldescens) and green (Rana clamitans) also present. Waldo Pond reaches a maximum depth of eight feet and possesses a bottom composed of pulpy peat with submerged wood. Yellow water lilies (Nuphar advena) are abundant in the limnetic waters, while sedges (Scirpus s p . ) dominate the littoral zone. vertebrates are represented by amphipods The in­ (Hyalella sp.), and aquatic insects mainly the dytiscid (Dytiscus s p .) . bluegills Yellow perch (Perea flavescens) and (Lepomls macrochirus) are the dominant 26 fish. Painted turtles (Chrysemys p i c t a ) are abundant throughout the pond. Station 7. A small, un-named pond along County Road 510 which is supplied by several small un-named creeks. The bottom consists of muck covered with leaves, sticks, and logs. Sedges (Scirpus sp.) and cattails (Typha latifolia) are present in small numbers. Snails •(Lymnaea sp.) are abundant along with a few isopods (Asellus sp.). Cyprinids are the dominant fish; other vertebrates are rare or absent. Station 9- Bagdad Pond possesses a bottom of muck with intermittent submerged logs. A portion of the pond has a bog margin of Sphagnum spp., leatherleaf (Chamaedaphne calyculata), pitcher plants (Sarracenia purpurea), and cranberries (Oxycoccus sp.). the pond, however, is surrounded by red maple Most of (Acer rubrum) , white birch (Betula papyrifera), and white cedar (Thuja occidentalis). Yellow water lilies (Nuphar advena) dominate the open waters along with sedges (Scirpus sp.). copepods Snails (Diaptomus sp.), and crayfish (Orconectes sp.) are abundant. . Frogs newts (Airin'!cola sp.), (Rana spp.), red-spotted (Notopthalmus viridescens), and painted turtles (Chrysemys p ic ta ) are also abundant. Station 10. The Park Cemetery Pond is a small pond reaching a diameter of about 30 feet. rounded by a mowed lawn. It is sur­ It Is completely crowded 27 with white water lilies (Nymphaea odorata) . Amphi­ pods (Hyalella sp.), isopods (Asellus sp.), and painted turtles (Chrysemys p i ct a) are abundant. Station 13. This is an un-named pond formed by the flooding of an old iron mine shaft. plied by underground springs. of clay and submerged logs. Water is sup­ The bottom consists Yellow water lilies (Nuphar advena) are present in small numbers. In­ vertebrates are sparse with snails (Physa sp.) being present in small numbers. No vertebrates were observed. Station 15. Horseshoe Pond is an out-pocketing of Horse­ shoe Lake. The bottom consists of sand with sub­ merged boulders. It reaches a depth of 15 feet. Most of its shoreline is surrounded by Sphagnum sp., leatherleaf (Chamaedaphne calyculata), sundews (Drosera spp.), and pitcher plants p urea) . (Sarracenla pur­ The northern shore borders U. S. Highway 4l and has been filled in with boulders. contains small growths of pondweeds The open water (Potamogeton spp.) and bladderwort (Utricularia vulgaris). Snails (Physa sp.), copepods (Cyclops sp.), and cladocerans (Bosmina sp.) are abundant. Bullheads (Ictalurus sp.) and cyprinids are the dominant fish with a few painted turtles Station 19. (Chrysemys p i c t a ) present. This study area consists of several inter­ connected ponds formed by the overflow of the Dead 28 River. The bottom.consists of sand with submerged wood and boulders. dominant plants. isopods Sedges (Scirpus spp.) are the Crayfish (Orconectes sp.) and (Asellus sp.) are present, but not abundant. Some cyprinids are present; however, frogs (Rana spp.) and toads (Bufo americanus) are the dominant vertebrates. Station 20. snow. This is a small vernal pond formed from melted The bottom consists of muck with submergent and emergent logs scattered throughout. Portions of the pond are surrounded by a sphagnum mat with white cedar (Thuja occidentalis) growing in it. Another portion is bordered by fill from a railroad grade. Eel grass (Vallisneria americana) and spike rush (Eleocharis sp.) are abundant. sp.), cladocerans Copepods (Cyclops (Daphnia sp.), and fairy shrimp (Eubranchipus sp.) are abundant. Progs (Rana spp.) and salamander larvae (Ambystoma spp.) are the dominant vertebrates. Station 31. Ramseth Pond is fed by a small un-named stream. The bottom consists of muck overlain with submerged sticks and leaves. Stonewort (Chara sp.) is very abundant throughout the pond, as well as a smaller population of pondweeds (Potamogeton spp.). Amphi­ pods (Gammarus sp.) and water boatmen (Sigara sp.) occur in sizeable numbers, as well as frogs (Rana 29 spp.), red-spotted newts (Notopthalmus viridescens), and painted turtles (Chrysemys p i c ta ). Station 32. Humboldt Pond was formed by the excavation of land and filled by surface water. The bottom con­ sists of sand with submerged boulders and sticks. Cattails (Typha latifolia) and horsetails (Equisetum sp.) are dominant in the shallows; no plants are evident in the deeper waters. Amphipods (Hyalella sp.) are present but not numerous. Vertebrates are seemingly absent. Station 37. This un-named pond is located on the Marquette Rod and Gun Club property and is formed by the back flow of the Dead River. and submerged wood. The bottom consists of muck Yellow water lilies (Nuphar advena) are abundant and dominate the open waters; sedges (Scirpus sp.) are abundant in the shallows'. Very few invertebrates are present with the exception of some aquatic insects mainly whirligig beetles (Gyrinus sp.). Frogs (Rana spp.) and toads (Bufo americanus) are abundant. Station 38. Wetmore Pond is fed by several small un-named streams. The bottom is composed of pulpy peat with overlain logs. Yellow water lilies are the dominant plants. (Nuphar advena) Backswimmers (Notonecta sp.), as well as painted turtles (Chrysemys p icta), are abundant. 30 Station 40. The Tourist Park Pond is formed by the back­ waters of the Dead River due to a dam. is mainly sand with a few boulders. The bottom This area is used as a swimming area; consequently, other than some eel grass (Vallisneria americana), aquatic plants are sparse. Clams (Pisidium sp.) and snails (Physa sp.) are present in small numbers, while vertebrates are absent in this area. Station 42. The Mine Shaft Pond resulted from the flooding of an abandoned mine shaft by spring w a t e r s . The bottom consists of clay. No higher aquatic plants or vertebrates are evident. Water scavenger beetles (Hydrophilus sp.) occur in small numbers. BOGS Bodies of standing water completely surrounded by a sphagnum mat containing such plants as leatherleaf (Chamaedaphne calyculata), sundews (Drosera sp.), pitcher plants (Sarracenia purpurea), and cranberries (Oxycoccus s p .). Station 17. This is a shallow body of water not exceeding four feet in depth. The bottom consists of fibrous and pulpy peat with little submerged wood. run-off is the main source of water. Surface It is sur­ rounded by a sphagnum mat which contains sundews (Drosera spp.), pitcher plants (Sarracenia purpurea) , leatherleaf (Chamaedaphne calyculata), and cranber­ ries (Oxycoccus s p O . rushes (Juncus sp.). The open water is dominated by Water boatmen (Sigara sp.) are 31 the dominant Invertebrates with some snails (Stagnlcola s p .) also present. No vertebrates were evident. Station 49. This bog receives its water from a small un­ named stream and also from springs. The bottom consists of fibrous peat with numerous submergent and emergent logs. The emergent logs are a favorite stand for painted turtles (Chrysemys p i c t a ) which are abundant and also mallards rhynchos) and black ducks (Anas platy- (Anas rubripes) which fre­ quent this body of water. The sphagnum mat surrounding this body of water contains the same plants listed for the previous habitat. The open water is heavily populated with yellow water lilies (Nuphar advena) . Amphipods (Hyalella sp.) and isopods (Asellus sp.) are the dominant inverte­ brates but are not abundant. RIVERS Bodies of running water whose width exceeds twenty feet along most of its course. Stations 18 and 4l. The Dead River is a long river which traverses most of Marquette County from West to East and empties into Lake Superior. Several hydroelec­ tric dams located along this river cause water back­ up into large basins. This river is approximately , 30.feet wide along most of its course and the depth varies from four feet along most of its course to 32 twenty feet behind the dams. The bottom is sand with overlying gravel, boulders, and submerged wood. Waterweed (Anacharis canadensis) is common in the eddies while stonewort (Chara sp.) occurs in patches in the faster moving waters. trichia sp.), chironomids Caddis cases (Leuco- (Chironomus sp.), and mayfly larvae (Hexagenia sp.) are abundant, as well as sponges (Spongilla sp.), snails (Physa sp.), planarians (Dugesia s p .), clams (Pisidium sp.), and crayfish (Cambarus sp.). This river is primarily a trout stream and also contains a wide variety and abundance of cyprinids. Station 43. The Carp River is also an extensive body of water which flows throughout most of the county, possesses many tributaries, and empties into Lake Superior. The bottom consists mainly of sand with intermittent graveled areas. The vegetation along its banks varies from one section of the stream to another but generally consists of tag alder (Ulnus incana), red-osier dogwood (Cornus stolonifera), and white cedar (Thuja occidentalis). The soil along this river contains an unusually large number of earthworms (Lumbrlcus sp.) which are washed into the stream during heavy rains. Aquatic plants are sparse, being represented by patches of waterweed (Anacharis canadensis) and eel grass (Yallisneria americana). Isopods (Asellus sp.), amphipods 33 (Hyalella sp.), crayfish (Orconectes sp.), and caddis larvae (Limnephilus sp.) are abundant. Salmonids, cyprinids, and catostomids are the most abundant fish present. Station 50. The Chocolay River spreads over 20-25 miles and possesses many tributaries. It is approximately 25 feet in width and varies in depth from 1-6 feet along its course. The bottom is mostly sand with intermittent stretches of gravel, rubble, and boulders. Pondweeds (Potamogeton spp.) and water­ weed (Anacharis canadensis) are present but not abundant. Planaria (Dugesia sp.), snails sp.), amphipods (Ferrissia (Hyalella sp.), and isopods (Asellus sp.) are abundant. Salmonids are the dominant fish with cyprinids and catostomids also present. CREEKS Bodies of running water whose width does not exceed 15 feet along most of its course. Station 14. This is a spring fed, un-named creek located along County Road 553 which varies within 2-3 feet in width. The bottom consists of sand with numerous submerged sticks and leaf litter. Stonewort (Chara sp.) is abundant and is the only higher aquatic plant in the area. Clams (Sphaerium sp.) and amphipods (Hyalella sp.) are present but not abundant. vertebrates were evident. No 34 Station 21. This creek is a small branch of the Yellow Dog River which meanders through a low marshy area. The bottom consists of muck with numerous submerged logs and tree stumps which project above the water. Sedges (Scirpus spp.) dominate the shoreline of the creek, whereas no higher aquatic plants are present in the open water. Snails (Physa spp.) and isopods (Asellus sp.)- are abundant along with some redspotted salamanders frogs Station 30. (Notopthalmus viridescens) and - (Rana spp.). Morgan Creek originates at Morgan Pond and empties into the Carp River. Alternate slow and fast moving water occur along its course. The bottom consists of sand covered with rubble and a few submerged sticks. The width varies within 3-10 feet while the depth varies from 1-3 feet. The slower waters are crowded with yellow water lilies (Nuphar advena) and pondweeds (Potamogeton spp.). Bur reed (Sparganlum androcladum) and spike rush (Eleocharis sp.) are abundant along the shoreline. Planaria (Dugesia sp.), snails (Hell'soma sp.), and blackfly larvae (Simulium sp.) are abundant. Cyprinids are the dominant fish with some salmonids also present. Station 39. West Branch Creek is spring fed and empties into the Escanaba River. It varies within 5-8 feet in width with a depth of 1-3 feet. The bottom is 35 sand with overlying rubble. pondweeds Sparse growths of (Potamogeton spp.) and bur reed (Spar- ganium androcladum) occur along its shoreline. Planaria (Dugesia sp.) are present in small numbers. Cyprinids are abundant and some salmonids also occur. Station 44. Dishno Creek originates from two small un­ named lakes and after traversing about six miles empties into the Peshekee River. The bottom is com­ posed of sand almost completely covered with rubble. This creek is approximately 12 feet wide and varies in depth within 1-3 feet. No higher aquatic plants were present at the collecting site. Sponges (Spongilla sp.), snails (Physa sp.), and blackfly larvae (Simulium sp.) are abundant. Cyprinids are the dominant fish present. Station 47. Barnhardt Creek is a spring fed and ground water seepage creek which is 10-12 miles long and empties into the Dead River basin. It possesses mainly a sand bottom with patches of rubble. No higher plants are present in the faster waters; yellow water lilies (Nuphar advena) are very abundant in the slower reaches of the creek. Aquatic insects are exceedingly numerous, particularly caddis larvae (Limnephilus sp.) and dobsonfly larvae (Corydalus cornutus). Planaria (Dugesia sp.) are also present in large numbers. This is primarily a trout stream with a wide variety of cyprinids also present. 36 Station 48. Goldmine Creek is a shallow, fast-moving creek. It is about six feet wide and two feet deep. bottom consists of sand and rubble. The No higher aquatic plants were observed in the collecting area. Snails (Physa sp.) and planaria (Dugesia sp.) are the predominant faunal inhabitants. DITCH A depression along a road which is fed by surface run-off and the overflow from a nearby stream. Station 8. The Mangum Road Ditch extends along the Mangum Road and is only 2-6 inches deep. composed of muck. The bottom is It is crowded with cattails (Typha latifolia) and sedges (Scirpus sp.). (Stagnicola sp.) and dragonfly naiads sp.) are present but not abundant. are the only vertebrates present. Snails (Helocordulia Progs (Rana spp.) 37 Table 1. Station Physical Features of the Waters Investigated In Marquette County. Bottom Depth (Ft.) Color T em p. °C Date 1 Fibrous Peat, Detritus 2 Brown 10 9/16/65 2 Sand, Gravel, Detritus 3 Clear 14 9/22/63 3 Pulpy Peat, Detritus 2 Brown 10 9/16/65 4 Sand, Detritus 4 Clear 10 10/10/65 5 Sand, Rubble, Detritus 2 Clear 15 10/10/65 6 Sand, Detritus 1 Clear 12 10/ 13/61 7 Muck, Detritus 3 Yellow 7 10/14/61 8 Muck Brown 9 10/ 16/61 9 Muck, Detritus 1 Yellow 18 8/28/65 10 Muck, Detritus 2 Brown 11 11/ 5/61 11 Sand, Rubble, Detritus 3 Clear 8 5/ 6/62 12 Sand, Rubble, Detritus 2 Clear 10 5/ 6/62 13 Clay, Detritus 3 Clear 8 5/13/62 14 Clay, Detritus 1 Clear 12 5/19/62 15 Sand, Detritus 3 Clear 8 8/25/62 16 Sand, Bedrock 3 Clear 8 8/26/62 17 Fibrous and Pulpy Peat 3 Brown 12 10/29/62 18 Sand, Gravel, Boulders 1 Clear 10 4/26/63 Sand, Detritus 2 Clear 12 4/27/63 Yellow 10 5/ 4/63 •5 20 Muck 21 Muck, Detritus 2 Brown 12 4/26/63 22 S and,,Rubble, Detritus 1 Clear 14 8/28/63 23 Pulpy Peat, Detritus 2 Clear 9 8/28/63 .5 Table 1 (cont ’d.) Station 24 Sand, 25 Bottom Rubble Depth (Ft. ) Color Temp. °C Date 1 Clear 18 9/ 1/63 Sand, Rubble, Detritus 2 Clear 17 9/ 5/63 26 Muck, Detritus 3 Clear 18 9/ 5/63 27 Pulpy Peat, Detritus 2 Clear 20 9/ 5/63 28 Sand, Gravel, Detritus 2 Clear 23 9/18/63 29 Sand, Detritus 2 Clear 24 9/18/63 30 Sand, 3 Clear 14 9/14/63 31 Muck, Detritus 1 Brown 16 5/ 8/64 32 Sand, Detritus 1 Clear 12 9/15/63 33 Sand, Rubble, Boulders .5 Clear 17 5/ 5/66 34 Sand, Rubble, Detritus 1 Clear 12 9/28/63 35 Sand, Detritus 2 Clear 11 9/28/63 36 Sand 12 Clear 10 8/25/63 37 Muck, Detritus 2 Brown 16 5/ 6/64 38 Pulpy Peat, Detritus 2 Brown 14 5/16/64 39 Sand, Rubble 1 Clear 13 6/28/64 40 Sand, Rubble 3 Clear 18 6/29/64 41 Sand, Rubble .3 Clear 8 9/16/65 42 Clay 2 Clear 12 7/ 5/64 43 Sand, Rubble 3 Clear 19 7/ 8/64 44 Sand, Rubble 1 Clear 10 8/ 2/64 45 Sand, Gravel, Detritus 2 Clear 22 8/ 2/64 46 Sand, Rubble 1 Green 10 8/ 4/64 47 Sand, Rubble 1 Clear 10 8/ 4/64 Rubble 39 Table 1 (c o n t ’d . ) Station Bottom 48 Sand, Rubble 49 Fibrous Peat, Detritus 50 Sand, Detritus Depth (Ft.) Color 2 Clear .4 Yellow 1 Clear Temp. °C Date 10 8/ 4/64 15 9/ 5/65 9 11/ 4/65 C ^ v j l - E r , oo U 1—1 j r O I—1 M I—1 O V O O O —q C T v V J l j=- uo ro I—1 w ct Flora Ceratophyllum Myriophyllum Floral I —’ I—’ 2. I—1 Table I—1 H* X o 3 X X X X X Nymphaea X X X X X X X X X X X X Nuphar Typha X X X Scirpus X Eleocharis X X X X X X Potamogeton X Najas Anacharls X X X X X Sagittaria Sparganlum X X Utrlcularla Juncus X Equlsetum X X X X Chara In X X Studied Valllsneria Inhabitants of the Waters Marquette County. X X u ru w m w h - £r UO IV) I— 1 O VO Station o j u j o j u j i j o u j u ) U o r o r o r o r o r o - < CT ' \ \ j - i - f c - u o r o h J o v o c o —a c ^ u i Flora C er a t op hy 11 um Myriophyllum Nymphaea > O ct 3 3* 3 CD ct «< s: • P ct CD 3 CO Centrarchidae CO ct X X X Cyprinidae X X Salmonidae X X X X X X X X X X X Salamanders X X X X X Frogs X X X X X Turtles 3 & HCD a UJ ff\ UJ UJ U1 UJ w C/3 i j o o o o o r o r o r o r o r o r o r o ro m o vo o o -j cr\ vji -t uj ro rvi w i—* w o h vo <+ fu ct- h* X X X X X >•* 'Isopoda X X x x x x x x x x x x x X x X x x xx X X X X X x X X X X Crayfish Insect Larvae Percidae Centrarchidae X Cyprinidae X Salmonidae x X X x x X X x X X X Amphipoda x x x x Salamanders x Frogs Turtles 3 (cont X X Clams X Tabl e X X Oligochaetes X X Fauna Planaria >> cd F-. o Ph 1—1 Floral Inhabitants of the Waters Studied In Barry and Kalamazoo Counties. 1—1 1—1 •ft ft o cd XI ft O p cd •H u o o u j >J s Fh cd cd x: d! s >> Eh ft ft ft to 3 ft Sh •H O to (0 •H Fh CtJ Xi o o CD rH W c o to •H p e ft rH 2 o •H Fh P D ft to to ft o C ft p <1) CO •rH ft O' w cd Fh cd XI o Barry County Station 1 X 2 X X 3 4 X X X X X X X X X X 5 X X 6 X X X X X X X X X X X X X Kalamazoo County 1 X X X X 2 X X X 3 X X 4 5 6 X X X X X X X X X X X X X X X X — 59 Table 6. Paunal Inhabitants of the Waters Studied in Barry and Kalamazoo Counties. G to <1> -p cd G 3 cd a cd a o o cd Td cd g o a CD o o •H Td cd Xi a rH O cd Td O a o to H ,G to •H cd PI •p o j o CO G 64 abundant leeches listed only Helobdella stagnalis is a glossiphonid. The size and activity of leeches would influence both the total number taken and the frequency with which they were encountered during the present study. The erpobdellids and hirudids are large, active leeches which are very responsive to water agitation and swim away in an undulating manner. On the other hand, the glossiphonids are smaller, more sedentary leeches which move mainly in a looping fashion by alternate attachment of the suckers. Thus, the latter group would be less conspicuous and would require more extensive searching to discover. These facts might ex­ plain why two erpobdellid leeches, Erpobdella punctata and Dina fervida, were the two most frequently encountered species. Too few piscicolid leeches were found to draw con­ clusions regarding their abundance and distribution. It was found during the present study that yellow perch (Perea flavescens) are a favorite host of Illinobdella alba and I. punctata. Keith (i960) also lists yellow perch as a host of I. alba but not of I_. p unctata. Perch collected from several locations during the present study were found to be parasitized by these two leeches. The examination of many cyprinids, salmonids, catostomids, and osmerids during the present study failed to produce any piscicolids, although fish in these families are attacked by these leeches 65 according to other authors, Thompson (1927), Meyer (1940), Keith (195^), and Rupp (1954) et al. Geographical Distribution Figure 6 denotes the location of the fifty study areas in Marquette County. This is a duplicate of Figure 2, page 15, for reference to the following figures. Figures 7, 8, 9, and 10 on pages 67-70 show the geographical distribution of the members of four families of leeches in Marquette County. The data show that the members of three families, namely, Glossiphonidae, Hirudidae, and Erpobdellidae, are uniformly distributed throughout the county. Too few specimens of piscicolid leeches were collected to draw conclusions regarding their distribution. Illinobdella punctata and I. alba are probably widely distributed throughout Marquette County, as well as throughout Michigan corresponding with the wide distribution of yellow perch (Perea flavescens). Piscicola m i l n e r i , on the other hand, seems to parasitize less widely distributed fish, such as smelt (Osmerus m o r d a x ), burbot (Lota lota maculosa), and whitefish (Coregonus spp.), and is probably more restricted in its distribution. 66 COLLECTING SITES MARQUETTE COUNTY, MICHIGAN 341 V MILES T I 4 • o N 147• >33 32 >3' T •!* 42 4 7 N 26 M t 4 8 N R 30 W R 28 W 1.REDBERRY POND 2.HARLOW LAKE 3.WALDO POND 4.SAUXHEAD LAKE 5.INDEPEN0ENCE LAKE 6.KAWBAWGAM LAKE 7.ROAD 510 POND 8.MANGUM ROAD DITCH 9.BA6DAD POND IQCEMETERY POND 11.RUSH LAKE 12.H0WE LAKE Fig. 6. 13.MINE SHAFT POND 14.ROAD 553 CREEK 15. HORSESHOE POND 16. TEAL LAKE 17. CRANBERRY BOG IB.DEAD RIVER 19. SMALL PONDS 20.VERNAL POND 21.SMALL CREEK 2ZC00PER LAKE 23.BANCR0FT LAKE 24.B0ST0N LAKE 25lTILDEN LAKE R 28 W 26.SCHOOLHOUSE LAKE 27. OGDEN LAKE 28. FISH LAKE 29. BACON LAKE 30.M0RGAN CREEK 31.RAMSETH POND 32.HUMBOLDT POND 33.LAKE SUPERIOR 34.M0UNTAIN LAKE 35.PINE LAKE 36.GOOSE LAKE 37.R0D B GUN CLUB POND 38.WETMORE POND 39.WEST BRANCH CREEK 40. TOURIST PARK POND 41. DEAD RIVER 42.MINE SHAFT POND 43.CARP RIVER 44.DISHN0 CREEK 45.ARFELIN LAKE 46.DEER LAKE 47.BARNHARDT CREEK 48.GOLDMINE CREEK 49.MIDDLE ISLE POINT BOG 50.CH0C0LAY RIVER Locations and soils of the waters studied. 67 DO GEOGRAPHICAL DISTRIBUTION GLOSSIPHONIDAE □O OO OO OO OO □ PLACOBDELLA RUGOSA O GLOSSIPHONIA COMPLANATA O PLACOBDELLA PARASITICA O HELOBDELLA STAGNALIS ■ PLACOBDELLA HOLLENSIS • HELOBDELLA FUCSA A THEROMYZON RUDE Fig. 7 68 GEOGRAPHICAL DISTRIBUTION PISCICOLIDAE ■ ILLINOBDELLA A PISCICOLA ALBA • ILLINOBDELLA MILNER! Fig. 8 PUNCTATA 69 GEOGRAPHICAL DISTRIBUTION A ▲ HAEMOPIS GRANOIS • MACRQBDELLA HIRUDIDAE ■ HAEMOPIS DECORA Fig. 9 MARMORATA 70 GEOGRAPHICAL DISTRIBUTION a ▲ DINA A NEPHELOPSIS ERPOBDELLIDAE PARVA • ERPOBDELLA OBSCURA ■ DINA FERVIDA Pig • 10 PUNCTATA 71 Table 7 shows a comparison between leeches collected in the Upper and Lower Peninsulas. The data Indicate, with few exceptions, the presence of the same species in both peninsulas. As only twelve bodies of water were examined in the Lower Peninsula, the collection records of Metzelaar et al. (1925, 1926, and 1927) in Miller (1937) are included in the comparison. Most of Metzelaar et al. work was done on Lower Peninsula waters, all their Upper Peninsula records having been obtained from Luce County. Seemingly significant species differences between the Upper and Lower Peninsulas and the factors thought to be involved will be described later. The species differences between the two peninsulas and their geographical distribution patterns in the midwest are as follows. Placobdella hollensis; This leech was found at only two locations in Marquette County and was not encountered in the Lower Peninsula by either Metzelaar or in the present study. This leech is known to be present in Minnesota, Nachtrieb, Hemingway, and J. P. Moore (1912), but not in the states of Illinois, Wisconsin, or Ohio. Perhaps it is a northerly species in regard to its distribution in the Midwestern states. Placobdella p i c t a : This leech was collected at one location in the Lower Peninsula by Metzelaar in 1926. It was not found in either peninsula during the present study. Bere (1931) reported it for Wisconsin. Sapkarev (1 9 6 7 ) found only one specimen of this leech in Lake Mendota. 72 Table 7. A Comparison of Leeches Collected in the Upper and Lower Peninsulas of Michigan. M = K = M-K = Collected by Metzelaar etal. (1925-1927) Collected by Kopenski (1961-1966) Collected by Metzelaar et al. and Kopenski Species Upper Peninsula Lower Peninsula M-K M-K Helobdella stagnalis K M-K Helobdella fusca K K Theromyzon rude K K Placobdella montifera K M Placobdella hollensis K --- Placobdella rugosa M-K M-K Placobdella parasitica M-K M-K Placobdella picta --- M Illinobdella alba K --- Illinobdella punctata K --- Piscicola milneri K --- Macrobdella decora K M-K Haemopis marmorata M-K M-K Haemopis lateralis --- M M M Haemopis grandis M-K M-K Philobdella gracilis --- K Erpobdella punctata M-K M-K Glossiphonia complanata Haemopis plumbeus 73 Table 7 (cont’d.) Species Upper Peninsula Lower Peninsula Dina parva K K Dina fervida K M-K Nephelopsis obscura K --- 74 Not enough information is available to postulate a geographical pattern in its distribution. Haemopis lateralis: Metzelaar collected this leech at two locations in the Lower Peninsula in 1926 and 1927* It was not found in either peninsula during the present study. This leech has been reported for Ohio, Miller (1929), Illinois, J. P. Moore Moore et al. (1901), and Minnesota, J. P. (1912). Haemopis pl umbeus: Metzelaar found this leech in Luce County in the Upper Peninsula in 1925 and in Mont­ morency County in the Lower Peninsula. It has been re- ported for Ohio, Miller (1929), and Minnesota, J. P. Moore et al. (1912). Philobdella gr acilis: Several specimens of this leech were collected from Otis Lake in Barry County in the Lower Peninsula during this study, but none were found in Marquette County waters. It has been reported for Illinois, J. P. Moore (1901), but not for Wisconsin nor Minnesota. Perhaps it is confined to the southern regions of the midwest area. Nephelopsis obscura: Marquette County waters. This leech is abundant in It was not found in the Lower Peninsula during the present study nor by Metzelaar. It has been reported for Wisconsin, Bere (1931), Sapkarev (1967), and Minnesota, J. P. Moore et al. (1912). also abundant in Canadian waters, Oliver (1958), It is 75 J. E. Moore (1966). It is not listed for Illinois nor Ohio. It appears to be a northerly species. As previously noted, too few specimens of piscicolids were obtained to draw conclusions as to their distribution. The same species are probably present in both peninsulas of Michigan since the fish which they parasitize are present in both peninsulas. When one compares the leeches present In a larger geographical area, such as the states of Michigan, Minnesota, Wisconsin, Illinois, and Ohio as illustrated in Table 8 , one finds a uniformity of species distribution with a few exceptions as discussed in the preceding paragraphs. The smaller number of species listed for some states is un­ doubtedly due to a lack of investigation rather than to a lack of species present. Consequently, an attempt to ex­ plain the presence or absence of a given species in each state would be non-justifiable. The large number of leeches known for Minnesota is due to the early thorough investiga­ tions of J. P. Moore et al. Keith (195^, I960). (1912) and more recently by To my knowledge, no up-to-date species listing has been compiled for Illinois. 76 Table 8 . A Comparison of Leeches in Five Midwestern St at Species Ohio 111. (Glossiphonidae) Glossip'honia complanata Minn. W i s e . Mict • X X X X X X Glossiphonia heteroclita X X X X X X X X X X Helobdella stagnaTis X Helobdella fusca Helobdella nepheloidea X X Helobdella punctata X X Helobdella lineata X Helobdella elongata Theromyzon rude X Theromyzon occidentalis X X X Hemiclepsis carinata X Hemiclepsis occidentalis Placobdella parasitica X X X X X Placobdella rugosa X X X X X Placobdella montifera X X X X X X Placobdella pediculata Placobdella picta X X X Placobdella hollensis Placobdella phalera X Actinobdella inequiannulata X X X X X X (Piscicolidae) Piscicola punctata Piscicola milneri X X X X X 77 Table 8 (cont ’d.) Species Ohio 111. Minn. W i s e . Mi cl X Piscicola geometra X Illinobdella alba X Illinobdella richardsoni X Illinobdella punctata X X X X (Hirudidae) Macrobdella decora X X X Haemopis marmorata X X X X X Haemopis grandis X X X X Haemopis lateralis X Haemopis plumbeus X X X X Haemopis latero-maculatum X X X X Philobdella gracilis X X (Erpobdellidae) Erpobdella punctata X X Nephelopsis obscura Dina fervida X X Dina parva Dina microstoma (total) X X 19 li» X X X X X X X X X X X 29 22 22 78 Ecological Factors Related to Leech Distribution Chemical Factors Total Alkalinity: A listing of the waters tested and their total alkalinity is shown in Table A 6 , page 1 5 3 . Table A7 on page 15^ shows the number of stations sampled, the mean total alkalinity, standard deviation, and range found for eleven species of leeches. Figure 11 depicts the percentage of occurrence of eleven species of leeches in relation to total alkalinity of the water. The alka­ linity ranges used are patterned after those of Mann (1955). Analysis of the data by families is as follows: Glossiphonidae: The glossiphonids were most frequently encountered in waters having a total alkalinity of lOOppm. or more. Forty-six percent of these waters contained at least one species of glossiphonid, whereas only 33% contained erpobdellids and 8% hirudids. An average, irrespective of species, of three glossiphonids were found in each body of water with a total alkalinity of less than 74ppm. On the other hand, an average of nine glossiphonids were found in each body of water with a total alkalinity of 7^ppm. or more. Although all glossiphonids studied were found more frequently in the higher alkaline waters, they were also present in waters with a lower total alkalinity as shown in Figure 1 1. Helobdella stagnails was found in 33% of the lentic waters in the l8- 59ppm. range and in 39%> of the waters in 79 % % so 40 30 20 10 A B C A BC Glosslphonia complanata H elobdella s t a g n a lls % 90 ■ •0 ■ 70 ■ AO - % 50 • 40 • 30 - 20 - 10 • AB C A B C Placobdella parasitica P laco b d ella rugosa A = Intermediate Standing Waters l8-59ppm. CaCOg B = Hard Standing Waters 60-99ppm. C = Hard Standing Waters CaC03 100- +ppm. CaCC>3 Fig. 11. Percentage of occurrence of eleven species of leeches at different alkalinities. 80 A B C A B C Maorobdella decora Haemopis grandls A B C A B C Haemopis marmorata Erpobdella punctata A * Intermediate Standing Waters l8-59ppm. CaCOg B = Hard Standing Waters 60-99ppm. CaCOg C = Hard Standing Waters 100- +ppm. CaCO^ Pig. 11 (Cont'd.) 81 40 30 20 10 A B C A B C Dina parva Dina fervlda U A B C Nephelopsls obscura A = Intermediate Standing Waters l8-59ppm. CaCOg B = Hard Standing Waters 60-99ppm. CaCC>3 C = Hard Standing Waters 100- +ppm. CaC03 Pig. 11 (Cont'd.) 82 the 60-250ppm. range. Mann (1955) found this leech in 60$ and 75$ respectively of the waters In the same ranges In England. Considering both lentic and lotic waters, Helob­ della stagnails wa-s found within an alkalinity range of 47-200ppm. in comparison to M a n n ’s range of 8-249ppm. Glosslphonia complanata was encountered in 25% of the lentic waters in the l8-59ppm. range and in 44$ -in the 60-250ppm. range. Mann (1955) found this leech in 42$ and 62$ respectively, of the waters in the same ranges in England. This leech was encountered most frequently in waters in the 60-99ppm. range which agrees with the findings of Mann. Considering both lentic and lotic waters, G. com­ planata was found within an alkalinity range of 24-200ppm. in comparison to M a n n ’s range of 10-280ppm. Sizeable percentage differences between the results of this study and that of Mann (1955) regarding the two leechesdiscussed are evident. This might be accounted for in several ways. (1) Mann's percentages for the l8-59ppm. waters are based on nineteen waters in comparison to twelve waters used in this study. (2) Eleven of the sixteen 60-250ppm. waters considered by Mann had total alkalinities of over lOOppm., four of which were over 200ppm. and con­ tained one or both of the two leeches concerned. In com­ parison, only present six of eighteen waters considered in the study had total alkalinities above lOOppm., one of which was over 200ppm. and contained neither of the two leeches in question. Since the conclusion has been reached 83 that both Helobdella stagnalis and Glosslphonla complanata prefer waters with a high alkalinity, their frequency of occurrence would undoubtedly be higher when a larger number of highly alkaline waters Is considered. (3) The time spent collecting at each site was not uniform In these studies, and (4) a variation in other chemical, physical, and biological characteristics of the waters considered in the two studies. Although percentage differences are evident, a definite pattern is indicated in both studies, i,.e., the frequency of occurrence of both species increases from the l8-59ppm. waters to the 60-250ppm. waters. Undoubtedly, the relationship between the distribu­ tion of the two leeches discussed and others is confined to the abundance of food organisms. One of the major food organisms of G. complanata and H. stagnalis is snails. Snails are most abundant in alkaline waters as they need CaCO^ to construct their shells. This may explain why these two species of leeches are more common in the higher alka­ line waters. Glosslphonla complanata and Helobdella stagnalis are the only species common to North America and Europe in which alkalinity relationships have been studied and upon which direct comparisons can be made. All subsequent alkalinity comparisons between European and North American leeches will be on a generic basis. 84 Placobdella parasitica and Placobdella rugosa exhibit distribution patterns similar to each other in re­ gard to total alkalinity. in waters above lOOppm. They were most frequently found Considering lentic and lotic waters P. parasitica was found within an alkalinity range of 24-250ppm. with a mean of 93ppm. P. rugosa was found within a range of 23-195ppm. with a mean of 89ppm. The main food of these two leeches consists of turtle blood. As turtles are widely distributed and do not appear to be restricted by alkalinity, the distribution of these two species is undoubtedly correlated with the distribution of turtles. Turtles would also act as disseminators of these leeches as they move from one body of water to another. No alkalinity studies on the distribution of the genus Placob­ della in relation to water alkalinity have been done in Europe; thus comparisons cannot be made. Hirudidae: Haemopis marmorata and H. grandls exhibit distribu­ tion patterns similar to each other in regard to total alkalinity. Both were found more commonly in waters with a total alkalinity of less than lOOppm. H. m a r m o r a t a , which was taken at six stations, was not collected in waters with a total alkalinity above lOOppm. H. gr a n d i s , recorded at four stations, was found in only one of the six waters above lOOppm. Similarities in distribution between these two species might be explained by the fact that their feeding habits, which will be discussed later, are 85 alike. The alkalinity range and mean of\ Haemopis spp. are narrower than those of the glossiphonids. Interestingly, Mann (1955) reported a related leech, Haemopis sanguisuga, more commonly in waters of less than lOOppm. in England. Macrobdella decora, on the other hand, was frequently encountered in waters of high alkalinity, as well as in lower alkalinities. Undoubtedly, the distribution differ­ ence between this species and the previous two hirudids lies in their feeding habits. Macrobdella decora is a voracious parasite which will attack fish, frogs, salamanders, turtles, and other vertebrates. Many of its hosts appear not to be restricted by alkalinity. This may explain why this leech is distributed over a wide range of alkalinities. Erpobdella punctata was found with almost equal fre­ quency (50-60$) over the three alkalinity ranges shown in Figure 11. The findings during this study for E. punctata compare closely with those of Mann (1955) and Sandner (1951) for a related species, E. octoculata, in Europe. Both E. punctata and E. octoculata are found over a wide range of alkalinities. Dina p a r v a , found at five stations, occurred with equal frequency throughout the three alkalinity ranges as shown in Figure 11. It was found within a range of 24-l40ppm. On the other hand, Dina fervida was found more often in waters below lOOppm. Twelve of the thirteen stations at which this leech was found had an alkalinity of less than lOOppm. Dina lineata, a related species in Europe, has not 86 been thoroughly Investigated In regard to alkalinity; thus comparisons cannot be made. Nephelopsis ob s cura, although found only at four stations, appeared most commonly in waters in the 60-100ppm. range and was not found in waters above lOOppm. No corresponding genus of leeches is present in Europe for comparison purposes. Piscicolidae: Not enough data were obtained to draw conclusions regarding the relationship of members of this family to total alkalinity. The data indicate that most of the leeches present in Michigan are tolerant of a wide range in alkalinity. The data also indicate that the percentage of occurrence of most species of the leeches studied is highest above a total alkalinity of 60ppm. This is particularly true of the glossiphonids and compares closely with the data obtained by Pawlowski (1936) and Sandner (1951) in Poland, Bennike (19^3) in Denmark, and Mann (1955) in England. pH: A listing of the waters tested and their pH is shown in Table A8 on page 155. The pH mean, standard deviation, and range found for eleven species are illustrated in Table A9 on page 156. Figure 12 shows the pH values for waters in which each of eighteen species of leeches occurred based on 38 waters. Most of the pH readings obtained for both Upper and Lower Peninsula waters were in the alkaline —1 bi ->i b» a> o as» o nt 9.5 i Fig. 12. Occurrence of in relation to the pH of water. 0> o Erpobdella punctata Dina fervida Dina parva Placobdella parasitica Placobdella rugosa Placobdella hollensis Glosslphonla complanata •: • t • • t •• • Macrobdella decora Haemopis grandis • •• • eighteen Haemopis marmorata Nephelopsis obscura Theromyzon rude species Helobdella stagnalis Helobdella fusca of Illinobdella alba leeches Illinobdella punctata Piscicola milneri Philobdella gracilis • • • •I OO 88 « range. Because only a few acid readings were obtained, only a general comparison can be made. The data show that leeches are abundant in alkaline waters and that the mean pH for most species was 7*5* Bennike (1943) states that his investigations regarding leech distribution and pH indicate that leeches are not abundant in acid waters. During this study Glosslphonla complanata and Placobdella parasitica were found at a pH of 5.9, while Placobdella hollensls, Haemopis grand!s, Erpobdella punctata, and Nephelopsis obscura were encountered at a pH of 4.5, but were not abundant in these waters. Helobdella stagnalis was found in waters ranging in pH f r o m -6.9-8.5. Mann (1955) found this leech within a pH range of 5.3-9*4 in England. Bennike (1943) found this leech at such low pH levels as 4.0 and 4.2. He states that sphagnum bogs are the only areas where this species is absent. In the present study, H. stagnalis was encountered in Middle Isle Point bog in Marquette County, a body of water surrounded by a sphagnum mat which could be considered a bog although the pH of the water was 7.3. Bennike's con­ clusion is based on acid sphagnum bogs and does not hold true for bogs containing alkaline waters. Bennike also states that H. stagnalis is especially prevalent in polluted waters. During this study, this leech was found in the Kalamazoo River, as well as in Deer Lake in Marquette County and Crooked Lake in Barry County. The Kalamazoo River re­ ceives the wastes from several paper mills along its shores and also sewage materials from various sources. Organisms characteristic of polluted waters, such as tubificids and chironomids, are abundant as are the blue green algae, Oscillatorla sp., Microcystis sp., and Anabaena sp. Deer Lake received the sewage wastes of the City of Ishpeming for several years. Oligochaetes and chironomids are abun­ dant as are blue green algae, Anabaena sp. and Microcystis sp. Certain sections of this lake are completely covered by a layer of duckweed (Lemna m i n o r ) probably due to the unnatural addition of nitrates and phosphates. Crooked Lake probably receives sewage materials by way of seepage from septic tanks along its shores. Oligochaetes are abundant and blue green algal blooms of Anabaena sp. and Microcystis sp. are common. The finding of numerous specimens of H. stagnalis in these modified waters during this study substantiates B en n i k e ’s conclusion that this species is especially prevalent in polluted waters. How­ ever, H. stagnalis is also present in clear, unmodified waters, such as Gull Lake in Kalamazoo County and Teal and Sauxhead Lakes in Marquette County. Thus, H. stagnalis is a very euryoecious leech which is able to adapt to many habitats and whose presence is not an indication of polluted waters. Sandner (1951) also found H. stagnalis to be a versatile leech in regard to its distribution in Poland as did Mann (1955) in England. _ Glosslphonla complanata was found over a pH range of 5.9-8.5 during this study. Mann (1955) found this species 90 over a range of 6.1-9.^. for this leech as 6.3-7.2. Bennike (19^3) lists the pH range He also states that this leech is not found in dystrophic waters. This conclusion would depend on the criteria used to designate a dystrophic lake. During the present study, this leech was taken from Purdy Lake in Barry County and Middle Isle Point bog in Marquette County, two bodies of water which could be classified as dystrophic on the basis of being shallow, poorly productive waters with heavily sedimented bottoms and crowded with Nuphar advena and other emergent aquatic plants. This leech is widely distributed in Europe as well as in North America. Helobdella stagnalis and Glossiphonia complanata are the only two leeches common to Europe and North America whose distribution in relation to the pH of water have been studied. The majority of leeches collected were found in waters which ranged in pH from 7*0-8.5. A large number and diversity of species were collected in waters which had a pH in the 8.1-9.3 range, e.g.., Teal, Gull, and Crooked Lakes. Mann (1955) also found leeches to be most abundant over this range in England. Although the frequency of occurrence of leeches in acid waters was much less than in alkaline waters, most sprecies were encountered over a wide range of pH. Undoubtedly, the relationship of pH, which is directly correlated with total alkalinity, and leech distri­ bution are directly linked to the distribution of food 91 organisms. This relationship has also been described by Pawlowski (1936), Bennlke (19^3), Sandner (1951), and Mann (1955). It is difficult to draw conclusions regarding leech distribution in relation to only two chemical characteris­ tics of water. Undoubtedly, total alkalinity and pH of water do have an effect upon the distribution of leeches indirectly by affecting the food organisms able to survive in such waters. Sandner (1951) states that in the majority of cases the influence of chemical factors on leech distribution may not appear at all. It may be true that a combination of many chemical factors operating collectively, not individually, indirectly regulates the distribution of leeches by directly affecting the food organisms present. 92 Physical Factors Lentic versus Lotic W a t e r s : Figure 13 shows the percentages of occurrence found for eleven species of leeches in thirty-nine lentic and ten lotic waters in Marquette County. A listing of the specific habitats com­ pared is found on page 66. A comparison between the occurrence of leeches in these two types of waters is con­ sidered by families as follows: Glossiphonidae: The glossiphonids were found to be the best adapted to the two types of waters. No glossiphonid varied more than Q% in its distribution in lentic and lotic waters. Glossiphonia complanata was found to be 2% more common- in lotic waters during this study. Mann (1955) found this leech to be 29 % more common in lotic waters in England; however Sandner (1951) found G. complanata more frequently in lentic waters in Poland. Bennike (19^3) encountered this leech with equal frequency in lentic and lotic waters in Denmark. From the results of the present study and those of the three European works, it would seem that G. com­ planata is equally adapted for life in both standing and running w at e r s . Helobdella stagnalis was found to be 6% more common in lentic waters than in lotic during the present study. Mann (1955).found this leech to be only 1% more common in lentic waters while Bennike (19^3) found it with equal frequency in lentic and lotic waters. From the results of Fig. 13. Percentage of occurrence of eleven species of leeches in thirty-nine lentic and ten lotic waters in Marquette County. PERCENTAGE Glossiphonia complanata Helobdella stagnalis Placobdella parasitica Placobdella rugosa Haemopis grandis Haemopis marmorata Macrobdella decora Erpobdella punctata Dina fervida \ t Dina parva Nephelopsis obscura mini VO (jU 94 the present study and those of the European works, it seems that H. stagnails has no preference for either standing or running waters. Placobdella rugosa was encountered 5% more frequently in lotic waters, whereas Placobdella parasitica was found to be 8% more common in lentic waters. These percentage differences for Placobdella spp. are s.mall and probably in­ consequential. Pawlowski (1936) considers current the most important factor limiting the abundance of leeches. In this connec­ tion, one might expect that the glossiphonids would be best adapted for living in lotic waters for several reasons as follows: (1) The glossiphonids are dorsoventrally flat­ tened; therefore they present running water. less surface area to the Also their venters are flat and smooth enabling them to better adhere to submerged objects. (2) Mann (1955) suggests the mode of reproduction of leeches as a factor determining their habitats. Glossi- phonia complanata is able to cement its cocoons to rocks, thereby decreasing the chances that they would be washed away by currents. (3) Another factor would be the care afforded the young by the parents. The young of the glos­ siphonids remain attached to the venters of the parents for a variable period of time and in this way tected somewhat from the actions of currents. would be pro­ 95 Hirudidae: Haemopis grandis and Haemopis marmorata were en­ countered slightly more often in lotic waters. These leeches are not dorsoventrally flattened nor do they possess the other adaptive characteristics of the glossi­ phonids. Consequently, one would not expect them to be abundant in lotic waters. This may be explained by the fact that these two leeches were usually collected in the eddies of the stream and not in the faster moving waters, whereas glossiphonids were commonly found in the faster moving waters. Cocoons of H. grand!s were found in holes in logs along the shoreline of Waldo Pond, Marquette County. Since Haemopis spp. are able to deposit their cocoons on moist soil, their progeny would not be subjected to the action of the current. Macrobdella decora was encountered in 15$ of the 39 lentic waters sampled and was not found in any of the 10 lotic habitats. It is by no means inferred that this leech is not present in running waters, since it has been ob­ served in running waters not considered in this study. Erpobdellidae: Erpobdella pu nctata, Nephelopsis obscura, and to a lesser extent, Dina fervida were more frequently encountered in lentic waters. N. obscura was not found in any of the ten lotic waters examined, while E. punctata was found in only one. It may be that current does influence the dis­ tribution of these three leeches. On the other hand, 96 Dina parva was found more commonly in lotic waters, pri­ marily in the eddies. Since D. parva is a smaller leech than the other three and presents less surface area to the currents, it is therefore theorized that this species is not as affected by the currents. In summary, water current could be an important regulating factor to the distribution of some hirudids and erpobdellids, but not to any great extent to the glossi­ phonids . Water Temperature: Table 9 shows the water tempera­ ture range found for eleven species of leeches. The tem­ perature ranges listed are based on temperatures taken at the time of collecting. Undoubtedly, these waters would reach higher and lower temperatures than indicated; conse­ quently it is not contended that these ranges are fixed. The data indicate that water temperature is not a major limiting factor to leech distribution. Two exceptions to this may be Nephelopsis obscura and Philobdella g racilis. N. obscura, found at nine stations, was not found in waters above 16°C. This leech was not found in the Lower Penin­ sula, nor has it been reported for Ohio or Illinois. How­ ever, it is common in the Upper Peninsula, Wisconsin, Bere (1931), Sapkarev (1967), and Minnesota, Nachtrieb, Hemingway, and Moore (1912). It is also common in Saskatchewan, Oliver (1958), and Alberta, J. E. Moore (1966). It seems therefore that N. obscura prefers colder waters. P. gracilis was encountered along the shore of 97 Table 9- The Water Temperature Range Pound for Eleven Species of Leeches Leeches Temperature Range °C Glossiphonia complanata 7-27 Helobdella stagnails 8-27 Placobdella parasitica 8.5-27 Placobdella rugosa 7-27 Haemopis grandls 8-24 Haemopis marmorata 8-27 Macrobdella decora '8.5-27 Nephelopsls obscura 8-16 Erpobdella punctata 8-27 Dina fervlda 8-24 Dina parva 8-25 98 Otis Lake, Barry County. The water temperature was 27°C and eight leeches were found underneath moist logs on shore two feet from the water's edge. The inference, therefore, is that these leeches took refuge out of the water because of the high temperature. This leech has been reported in Illinois by J. P. Moore (1901), but was not found in Minnesota, Wisconsin nor Canada. A postulation hereupon assumes that this leech is a warm water dweller and that water temperature is a limiting factor in its distribution. Sandner (1951) does not consider water temperature a major limiting factor to leech distribution. however, Bennike (19^3), considers water temperature a controlling factor of leech distribution since it affects their reproduction. He states that leech eggs will not mature in waters whose summer maximum temperature is less than 11°C. This low summer maximum temperature listed by Bennike is atypical of most waters and probably only pertains to springs. Bennike found that high water temperature was of no importance in Danish waters. In small ponds, however, temperatures may become so high as to be critical for certain leeches. Bennike's conclusion may fit the situation previously described for Philobdella gracilis; however, the leech acclimated to the high temperature problem by moving tem­ porarily to a moist and probably cooler area on land. 99 Aside from the two exceptions mentioned, water tem­ perature did not appear to constitute a limiting factor to distribution of the leeches studied. Water C o l o r : Since no quantitative measurements were made of color, only a few cursory remarks can be made, on the basis of gross observations. In most cases a greater number and variety of leeches were found in clear waters, e.g^., Teal, Independence> and Mountain Lakes, Marquette County, and Gull Lake, Kalamazoo County. Mann (1955) also found a greater number and variety of leeches in clear waters. However, exceptions did occur during this study, whereby a large number and variety of leeches were found in brown waters with high amounts of dissolved organic matter, namely Otis Lake, Barry County, and Ramseth Pond, Marquette County. Therefore, a suggestion is proposed that water color plays a minor role if any in determining the abundance and distribution of leeches. Neither is it a good criterion for predicting what leeches may be present in a body of water. Water D e p t h : Most collecting during this study was done at depths from 0 to 1.2 meters, the exceptions being some piscicolid leeches which were removed from fish taken at depths of 3.6 and 6.0 meters. Eckman dredge sampling was attempted both from a boat and through the ice, but proved ineffective in obtaining a good representation of the leech fauna within a body of water. 100 Bennike (19^3) found leeches down to a depth of 12 meters and discontinued sampling beyond this depth for lack of sufficient findings. He cites lack of vegetation, absence of substratum for attachment, and low oxygen content as reasons for the scarcity of leeches at greater depths. Bere (1931) took Helobdella stagnails from a depth of six and seven meters in Trout Lake, Wisconsin. leeches were adhering to a gill net. These Bennike (19^3) found H. stagnalis at 5 and 12 meters in Denmark. Sapkarev (1967) found H. stagnalis at a depth of 12 meters in Lake Mendota. During the present study, leeches were found to be most abundant within a depth range of 0 to 1.2 meters. Sapkarev (19 6 7 ) found that the maximum density of glossi­ phonids occurred at a depth of 0 to 3 meters and that the maximum density of erpobdellids occurred at 0 to .5 meters. He found that all species inhabited the littoral zone and that in all cases the number of species decreased with an increase in depth. Therefore, leeches are probably restricted by great depth for the reasons cited by Bennike (1943). Bottom Composition: Table A10, page 157, shows the bottom composition of the fifty waters studied in Marquette County and the number of times each of eleven species of leeches occurred along these bottoms. Too many variables exist to draw statistical conclusions regarding the rela­ tionship of leeches to particular bottoms. The primary 101 problem encountered is the accuracy with which a given bottom can be described. Other variables are differences in the amount of time spent collecting at each site, the number of times each site was visited, and the few sites representing certain bottoms. Because of these problems, only general conclusions can be made. The data indicate that all eleven species of leeches were present along sandy bottoms with some combination of submerged matter and along all the bottoms without a sand base. Discounting disturbed leeches, none were found along a pure sand bottom without submerged objects. Likewise, few species occurred along clay, peat, or muck bottoms without the presence of submerged materials. Although few bottoms were found without submerged objects, probably plain peat and muck bottoms would be more suitable to leeches since they would be better able to burrow into them. The difference between the submerged materials lying above the base materials is too slight to show preference for one over the other. Consequently, a comparison was made between the percentage of occurrence of a given species along four types of bottoms irrespective of the submerged materials overlying each. comparison are shown in Table 10. The results of this The data indicate that Placobdella rugosa, Haemopis marmorata, and Dina parva occurred slightly more often along sand bottoms. Glossi- phonla complanata, Helobdella stagnalis, Haemopis grandis, Erpobdella punctata, and Dina fervida occurred more 1 0 2 Table 10. The Percentage of Occurrence of Eleven Species of Leeches along Pour Types of Water Bottoms. (Numbers in Parentheses Represent Number of Waters Sampled) Species Sand (31) Clay (3) Peat (7) Muck Gloss iphonia complanata 19 0 57 44 Helobdella stagnalis 26 0 29 22 Placobdella parasitica 26 0 43 22 Placobdella rugosa 19 0 14 11 Haemopis grandis 13 33 29 22 Haemopis marmorata 29 0 0 22 Macrobdella decora 10 0 14 22 Erpobdella punctata 39 0 71 11 Dina fervida 23 33 29 22 Dina parva 23 0 0 22 Nephelopsis obscura 13 33 14 33 103 commonly along peat bottoms, while Nephelopsis obscura and Macrobdella decora tended to be more common along muck bottoms. Clay bottoms were discounted as only three of the habitats studied possessed clay bottoms. In most cases leeches were found under the submerged objects typical of water bottoms. However, some more interesting locations of certain leeches can be mentioned. Helobdella stagnalis were found adhering to the undersides of Nuphar advena leaves. Dina parva were found fastened to the roots of Typha latifolia, and submerged pieces of concrete produced an abundance of Glossiphonia complanata. An interesting correlation may be exhibited by the latter situation. There is a possibility that the lime dissolving out of the concrete acts-as an attractant to snails, as lime is needed for shell construction. Since snails are the main source of food for G. complanata, this might account for its abundance in this situation. In summary, it is difficult to assign a given leech to a particular type of bottom. The main requirement for the occurrence of any leech, regardless of the nature of the bottom, appears to be the presence of submerged objects, such as wood, rocks, debris, rooted aquatic plants, etc. Seldom were leeches found along plain bottoms where no attachment materials were available. Submerged materials protect leeches from predation, enhance their resistance to currents, provide habitats for their food organisms, and 104 provide a substratum upon which they may deposit their cocoons. Soils: A description and designation of the soils surrounding the fifty bodies of water sampled in Marquette County is found on pages 14 and 15. A Chi-square test was performed attempting to correlate the soils surrounding each body of water with the leeches inhabiting it. Due to the same variables encountered when attempting to correlate leeches with bottom types, it was impossible to obtain a valid statistical answer. Only a general conclu­ sion regarding the relationship of leeches to soils can be given. Certainly the nature of the surrounding soils would be reflected in the chemical characteristics of the water due to the leaching out of materials from the soils by rain, streams, and springs. Waters in limestone areas would undoubtedly have a high total alkalinity and conse­ quently a high pH. On the other hand, waters in igneous rock areas would likely have a low total alkalinity and thus a low pH. Therefore, surrounding soils would only exert an indirect effect on leech distribution by altering the chemical nature of the water. Moreover, the chemical nature of the water, which has been discussed previously, probably exerts only an indirect effect by regulating the abundance and distribution of the food organisms of leeches. 105 Biological Factors Feeding H a b i t s : Table 11 shows the preferred foods of twenty-two leeches Inhabiting Michigan waters. This listing was compiled from personal observations as well as from the findings of J. P. Moore et al. (1937), Meyer (1912), Miller (1954), Rupp (1954), Keith (i960), J. E. Moore (1964 and 19 6 6), and Sapkarev (1967). Glossiphonidae: Members of this family possess a pore-like mouth through which a muscular proboscis is extruded. The proboscis is inserted into the tissues of the host and blood or the entire soft parts of the animal are withdrawn. Glossiphonia complanata was observed feeding on the snail, Physa sp. Sapkarev (1967) observed this leech preying upon the snails Physa gyrina and Planorbis pa rv u s . It is generally agreed by other workers that snails of several -species are the main food items of this leech. Neither Helobdella stagnalis nor H. fusca was observed feeding during this study. Other investigators list aquatic insects and snails as the favored foods of Helobdella spp., plus others listed in Table 11. The few specimens of Theromyzon rude encountered during this study were found in the non-parasitic state. Other workers list the blood of aquatic birds as the main source of nutrition for this leech. Meyer (1954) found this leech parasitizing waterfowl in a pond in Manitoba and suggests that they may cause the death of small 106 Poods of Michigan Leeches. CO TS •H rH CO p CD < < P o H& P O' U1 a P 4 P w M -o •a o o o' o' Q- & CD CD 1-* M h-1 H* P & 'O P CD P ■ —' rs o ct P ct P Tl S' H* M O O' 3 P O 4 o o' p& CD CD I-1 M 1 —1 M P P (TC tr 1 cd CD o 2 CD CO Oi CD O P o O H- 4 I-1 P H* CO X Other Leeches Table X Earthworms 11 Aquatic annelids (cont Foods X X X X X X X X X X X Snails Crustaceans Aquatic insects X 1 X X Fish X Salamanders X Frogs X Turtles Birds X X X Mammals 108 aquatic birds. J. E. Moore (1966) lists grebes (Podllymbus spp.) as well as the gadwall (Anas strepera), pintail (Anas acuta), shoveler (Spatula clypeata), baldpate (Marcea americana), coot (Fulica americana) , and other aquatic birds as hosts of T. rude in Alberta. Infestation occurs mainly in the nasal cavities of these birds and J. E. Moore also suggests that this leech may be responsible for a con­ siderable degree of mortality in young birds. Since T. rude seems to feed exclusively on the blood of aquatic birds, its abundance and distribution would tend to be correlated with the abundance and distribution of waterfowl. Turtle blood is the main food of Placobdella spp. During this study, Placobdella parasitica and P. rugosa were found adhering to the following turtles: western painted (Chrysemys plcta be lli), midland painted (Chrysemys picta marginata), Blandings (Emydoidea blandingi), ♦ snapping (Chelydra serpentina), spotted (Clemmys guttata), eastern box (Terrapene Carolina), and musk (Sternothaerus odoratus). Twenty-five specimens of P. parasitica were re­ moved from one musk turtle found along the shore of Pleasant Lake, Barry County. This turtle died in the laboratory the following day and it is conceivable that death may have been due to a loss of blood. Placobdella spp. attach mainly in the axillae of turtles, undoubtedly because the skin is more tender and moist in these areas and possibly more vascularized. Besides attacking turtles, one specimen of P. parasitica was removed from the head of 109 a red-spotted newt (Notopthalmus Virldescens louisianensis) taken from Redberry Pond, Marquette County. P. hollensis was not observed feeding, while P. plota and P. montlfera were not encountered during this study, although three specimens of P.-montlfera were removed from the skin of a leopard frog (Rana piplens) taken from Chicagoan Creek, Iron County. Other foods of Placobdella spp. as mentioned by other Investigators are aquatic annelids, snails, aquatic insects, fish, and waterfowl. P. parasitica and P. rUgosa are abundant and widely distributed in Michigan. This is undoubtedly correlated with the abundance and distribution of turtles in the state. P. hollensis, P. p i c t a , and P. m on tl f e r a , however, are not common in Michigan waters. Since these three species also parasitize turtles, their abundance and distri­ bution are probably regulated by some factor or factors other than food. Unfortunately these species were not found often enough to determine what the regulating factor or factors might be. Piscicolidae: The members of this family possess a pore-like mouth through which a muscular proboscis may be protruded. The proboscis is inserted into the bodies of fish where blood and body fluids are extracted. Illinobdella alba and I . punctata were found ad­ hering to the bodies of yellow perch (Perea flavescens). They were found attached mainly to the fins, in some cases 1 1 0 to the gills, and in one case to the lining of the mouth. The pectoral and pelvic fins were the favored sites of attachment possibly due to the nature of their vascular supply. As many as seven or eight leeches were removed from a single fin in some cases. Individuals of both species were found adhering to the same fin at the same time on some perch. They were found on perch ranging from four to ten inches in length and no preference for any par­ ticular size could be determined. Keith (i9 6 0 ) also found I. alba on yellow perch in Minnesota. The examination of a total of twenty-five northern pike (Esox lucius) from Goose, Bear and Sauxhead Lakes, Marquette County, and ten walleyes (Stizostedion vitreum) from Teal Lake, Marquette County, produced no piscicolids. Yellow perch in Goose and Teal Lakes, however, were heavily parasitized by I. alba and I. punctata. The preference shown for yellow perch might be due to the fact that their fins are thinner than the other two species and probably the blood vessels sup­ plying them are closer to the surface. Yellow perch may spend more time near the bottom or shore than either northern pike or walleyes and thus be more prone to leech attachment. Lewis E. Peters (personal communication) took I. punctata from the fins of walleyes in Portage Lake, Houghton County. It seems therefore that although yellow perch are the favored host of these two leeches, walleyes and possibly other fish are also attacked. Piscicola milneri was found adhering to the body of a burbot (Lota Ill lota maculosa) taken from Lake Superior. Attachment to the body of this fish Is undoubtedly not Impeded by Its small cycloid scales. Meyer (195*0 took this leech from land­ locked salmon (Salmo salar sebago) In Maine. Keith (i9 6 0 ) lists smelt (Osmerus m o r d a x ) and whitefish (Coregonus spp.) as hosts of this leech in Minnesota, and J. E. Moore (1964) found P. milnerl on white suckers (Catostomus commersonnii) in Alberta. The examination of fifty smelt and twenty-five white suckers during this study failed to produce this leech. I. alba and 1^. punctata were encountered in only two lakes during this study probably because yellow perch from these two lakes were extensively examined. It is suspected that these two leeches are abundant and widely distributed in Michigan correlating with the abundance and’ distribution of yellow perch. On the other hand, P. milneri might be more closely associated with Lake Superior and waters in close proximity to it. The three fish, namely smelt (Osmerus mordax), whitefish (Coregonus spp.), and burbot (Lota lota maculosa), known to be attacked by P. milneri are typically Lake Superior fish. To my knowledge this leech has been found in only two. Lake Superior states^ Minnesota and Michigan; thus the previous conclusion is based on these findings. Hirudidae: The members of this family possess a large mouth surrounded by lips and do not possess a muscular proboscis. 1 1 2 The mouths of Macrobdella dec or a, Haemopis marmorata, H. lateralis, and Philobdella gracilis contain jaws bearing numerous teeth. Consequently, these leeches are able to bore Into the tissues of various hosts or devour whole organisms. Other representatives, namely Haemopis grand!s and H. plumbeus, lack jaws and must feed on whole organisms or attach to wounds created by other leeches. Several observations were made regarding the feeding habits of H. m ar mo ra t a. Two specimens were found adhering to the detached cheliped of a crayfish. dividual was observed devouring an earthworm. One in­ Two specimens were encountered feeding on a portion of a dead fish. specimen, 82 mm. in length, was discovered with a specimen of Nephelopsis obscura, 72 mm. its mouth. One in length, protruding from Several individuals were observed breaking up frog egg masses and devouring the eggs. These findings during this study show that the feeding habits of this leech are varied. J. P. Moore (1912) and J. E. Moore (1966) reached the same conclusion in regard to the feeding habits of this leech. this study. H. grandis was not observed feeding during According to other investigators, its feeding habits are similar to those of H. m a rm or at a . Rupp (195*0 observed H. grandis attached to the wounds on brook trout (Salvelinus fontinalis) caused by Macrobdella decora in a pond in Maine. The feeding habits of M. decora were studied in an aquarium in the laboratory. Several guppies were 113 Introduced into an aquarium containing several specimens of this leech. One leech fastened to a guppy, pulled it to the bottom, and killed it. Since guppies are not associated with this leech in nature, this incident shows that this leech can adapt to a new source of food. attacked green frogs Several individuals (Ran a cl ami tans-)-, adults and tadpoles, and spotted salamanders (Ambystoma maculatum) when they were introduced into the aquarium. In the case of the salamanders, attachment was centered primarily on the digits, whereas no pattern of attachment was evident in the case of the frogs. In neither case were the hosts able to dislodge the leeches despite attempting to scratch them off with their limbs. Rupp (195*0 observed M. decora attacking and killing brook trout (Salvelinus fontinalls) weighing from one to two pounds in a Maine pond. Philobdella gracilis was not observed feeding, while Haemopis lateralis and H. plumbeus were not found during this study. The main foods of these leeches, as listed by other workers, are earth­ worms, aquatic annelids, and aquatic insects. The data indicate that H. marmorata, H. grandis, and M. decora feed on a variety of animals which are abundant and widely distributed in Michigan waters. This probably accounts for the great abundance and wide distribution of these species in the state. The foods of H. lateralis and H. plumbeus, as determined by other investigators, do not seem to be as varied as those of H. marmorata and H. grandis. Perhaps this accounts for the scarcity of these im leeches in Michigan waters. Philobdella gracilis was en­ countered only once during this study. It has previously been noted that water temperature, rather than food, seems to be the most important limiting factor to the distribu­ tion of this leech. Erpobdellidae: The members of this family possess a large jawless mouth with rounded lips and are not equipped with a pro­ boscis. Therefore, they must swallow their food whole. No representatives of this family were observed feeding. A listing of their preferred foods, according to other authors, includes aquatic annelids, snails, crustaceans, and aquatic insects. Erpobdella punctata feeds on a wide variety of organisms which enables it to inhabit many types of waters. This possibly accounts for its great abundance and its having been encountered more frequently than any other leech during this study. Dina fervida and Dina parva seem to have a more restricted diet than E. punctata. These two species, however, feed on aquatic insects which are probably never absent in any body of water and consequently these leeches are widely distributed in Michigan. The abundance and distribution of Nephelopsis obscura, although probably"Influenced somewhat by its food organisms, seems to be more regulated by high water temperature. 115 Parasites and Predators of Leeches: Parasit e s . Information dealing with organisms parasitizing leeches is scanty. Mann (1962) discovered the protozoans Entamoeba aulastoml in the gut of Haemopls sp. and Orcheobius herpobdellae in the testes of Erpobdella sp. Mann also states that many species of Trypanosoma spp. from fresh water fishes are transported by Pisclcola spp. J. E. Moore (1964) reports collecting Haemopls marmorata in Alberta with numerous metacercariae in its body wall. He also found several specimens of Erpobdella punctata and Nephelopsis obscura containing the metacercariae of strigeid trematodes in their body walls. Predators. J. P. Moore (1923) lists crayfish, turtles, snakes, crows, kingfishers, and mink as predators of leeches based on observations made at leech farms in the environs of Philadelphia. Mann (1962) states that the erpobdellids in particular are preyed upon by trout, perch, tench, sticklebacks, eels, herons, swans, ducks, and bit­ terns in England. He also suggests that aquatic amphibians and mammals as well as carnivorous Hemiptera and Odonata may also feed on leeches. J. E. Moore (1966) found a single specimen of Erpobdella punctata and three specimens of Nephelopsis obscura in the esophagus of a Lesser Scaup (Aythya affinis) in Alberta. Certain species of leeches have been observed feeding on others. J. P. Moore (1923) observed Haemopls grandis devouring specimens of Macrobdella decora in the 116 Palisades Park area of New York. Mann (1962) cites Glossiphonia complanata as a predator of Erpobdella octoculata in England. G. complanata may possibly attack E. punctata In North America. During the present study, Haemopls marmorata was observed eating a specimen of Nephelopsis obscura. J. E. Moore (1966) reports finding H. marmorata feeding on Helobdella stagnails, Erpobdella punctata, and Dina dubia in Alberta. Haemopis spp., being large in size (6-12") with large mouths, probably take more leeches than any of the others. Erpobdellids, being medium sized (2-4") active leeches, are probably preyed upon more than others. • Due to our incomplete knowledge of leech parasites and predators, it is not possible to determine what effect they may have on the abundance and distribution of leeches. The parasites probably exert little or no effect. Fish, aquatic birds, and other leeches, being the principal predators, may exert some effect especially in the case of the erpobdellids. Reproduction: The leeches encountered in the breeding condition during the present study are discussed with comparisons being made with the findings of other in ­ vestigators. An attempt is made to relate the reproductive characteristics of three leech families to their abundance and distribution in Michigan waters. 117 Glossiphonidae: Most glossiphonids lack an eversible penis and sperm is transferred by means of spermatophores. The spermatophores are attached to the body of another leech. These spermatophores are attached at various positions along the dorsal surface of the leech. Placobdella rugosa was found during the present study with spermatophores attached near the anus which is some distance from the reproductive organs. The sperm pass through the epidermis into the dermal connective tissue and from there to the coelomic spaces in which the ovaries lie, Mann (1962). In some cases, e.g. Glossiphonia complanata, the thin-walled cocoons produced are cemented to some submerged object and the adults cup themselves over the cocoons, thereby shel­ tering them with their bodies. In other cases, e.g. Helobdella spp., the thin-walled cocoons remain attached to the venters of the adults. In both cases, when the young develop, they fasten to the venter of the parent by means of their embryonic attachment organs. Six specimens of Helobdella stagnalis were found in Harlow Lake, Marquette County, on May 15, 1968, with cocoons containing developing young attached to their venters. These cocoons contained the following number of embryos: 37, 37, 37, 31, 15, and 13. Probably the latter two cocoons ruptured during collecting or preservation as free embryos were observed floating in the medium. This species was found with young attached in the Dead River and 118 Deer Lake, Marquette County, on July 7, 1964, and August 4, 1964, respectively. H. stagnails were also taken with young from Gull Creek, Kalamazoo County, on July 29, 1965. The number of young attached ranged from 12-24 averaging 18 per adult. Undoubtedly, this leech produces two broods a year or is a continuous breeder through the summer months judging from the extended period over which it was found with young attached. This agrees with the findings of J. P. Moore (1912) in Minnesota, Mann (1962) in England, and J. E. Moore (1964) in Alberta. Sapkarev (1967) found H. stagnails with eggs as early as May 10 in Lake Mendota. His collections of this species during the period from May 20 to May 30 showed 80$ of the individuals with eggs, 10$ with young, and 10$ with neither eggs nor young. During the period June 1-9, Sapkarev found that 60$ of the individuals captured had young attached, 30$ were with eggs, and approximately 10$ had neither eggs nor young. He found that the breeding period of H. stag- nalis covered less than two months (May and June). Bennike (1943) found H. stagnails with young attached from June 1 — V‘ to September 11 in Denmark with the number of young averaging 20 per parent. J. E. Moore (1964, 1966) found this species with young attached from May to September in Alberta. He found that the number of young ranged from 17-23 averaging 20. According to J. E. Moore the eggs hatch in five days, embryos are present for three days, and the young are carried by the parent for 15-30 days. Of interest 119 Is the fact that the number of young found attached during the studies of Bennike, J. E. Moore, and Kopenski averaged 20. Since four cocoons examined during the present study contained over 30 developing embryos, some embryos must fail to develop into young. It appears that the breeding period of H. stagnails and the number of young produced is similar in North America and Europe. Helobdella fusca were found with young attached in Crooked Lake, Barry County, on July 22, 1965, and in the Kalamazoo River, Kalamazoo County, on July 27, 1965. The number of young found attached averaged 16 per adult. J. E. Moore (196 6) .found four specimens of this leech on August 10 in Alberta, each bearing egg capsules. H. fusca probably also produce two broods a year and a similar number of young as H. stagnalis. No specimens of Glossiphonia complanata were found with eggs or attached young during this study. Sapkarev (1967) found this leech with eggs from the end of May through July and encountered young during all of July and August. Bennike (19*13) found this leech with young attached in June and July in Denmark. The number of young he found ranged from 25-67 averaging 45. Mann (1962) found G . complanata in a breeding condition from March to May in England with an average of 26 young produced per adult. suggests that this leech produces two broods a year. He J. E. Moore (1964, 1966) found this leech In a breeding condition from May to July in Alberta. He found one specimen 1 2 0 covering 121 eggs (apparently in four capsules) which Is considerably more than produced by this leech In Europe. He found no evidence that G. complanata produced two broods a year in Alberta. Placobdella parasitica was found with young attached in Cooper Lake and Redberry Pond, Marquette County, on August 28, 1963, and November 11, 1961, respectively. This species was also found with young attached in Augusta Greek, Kalamazoo County, on June 20, 1963 . The specimen taken from Cooper Lake had 98 young attached to its venter. The specimen collected on November 11 was kept in an aquarium in the laboratory until it died on April 19 of the following year. During this time the young remained attached to it. Metzelaar et al., in Miller (1937)* found P. parasitica with young attached from March 15 to July 22 in Michigan. Miller suggests that this leech produces several broods per year. The finding of P. parasitica with young attached in June and November during the present study seems to substantiate Miller's suggestion. Three specimens of Placobdella rugosa were collected with spermatophores attached in Harlow Lake on May 15, 1968. This species was found with young attached in Otis Lake, Barry County, and Barnhardt Creek, Marquette County, on July 26, 1965, and August 4, 1964, respectively. The specimen taken from Otis Lake had 50 young attached while the three specimens collected from Barnhardt Creek had an average of 91 young per adult. The latter number compares closely with the 98 young removed 1 2 1 from a specimen of P. parasitica. J. E. Moore (1964) found a specimen of P. rugosa in Alberta carrying 95 young. It appears, therefore, that P. parasitica and P. rugosa produce 90-100 young and may produce more than one brood a year. Placobdella montifera, P . holiensis, and Theromyzon rude were not encountered in a breeding condition during this study. Since all members of the genus Placobdella whose reproductive habits are known reproduce in a similar manner, it is suspected that the breeding habits of P. mon­ tifera and P. hollensis may be similar to those described. J. E. Moore (1964) found T. rude with young attached from June through August in Alberta. One specimen found in the Kakisa River, Northwest Territories, had 234 young attached to it. Another specimen taken from a pond near Edmonton produced four capsules containing a total of 191 young. J. E. Moore states that the young of T. rude remain attached to the parent for a month or longer. The data for the glossiphonid leeches indicate that most species produce a sizable number of young. This is particularly true of Placobdella parasitica, P . rugosa, and Theromyzon rud e. These three leeches are mainly parasitic and tend to follow the general biological pattern of parasitic animals producing a large number of young. Helob­ della spp. and Glosslphonia complanata appear to produce a somewhat smaller, but yet substantial number of young. These leeches are not commonly parasitic, but feed on other 1 2 2 leeches, aquatic annelids, snails, and aquatic insects. Undoubtedly, the great abundance and wide distribution of P. parasitica, P. rugosa, H. stagnails, and G. complanata are somewhat due to their high reproductive capacities. The survival rate of the young of these leeches is also probably high since they are carried by the parents for a variable length of time. H. fusca and T. rude were not found frequently enough to determine a relationship between their distribution and reproductive characteristics. Hirudidae: The members of this family possess an eversible penis by which sperm transfer takes place. The cocoons produced are rounded with small papillae at each end where they were sealed. Since members of this family abandon their cocoons after depositing them, it is difficult to obtain information concerning their reproductive charac­ teristics. Consequently, no information was obtained about the breeding habits of Haemopls marmorata, Macrobdella dec­ ora , or Phllobdella gracilis. J. E. Moore (1966) reports collecting a large specimen of H. marmorata with a swollen clitellum on July 31 in Alberta. He also states finding juvenile individuals on October 16. These two records constitute the only information on the breeding habits of this leech in Alberta. Five cocoons of Haemopls grandis were found in holes in a log on the shore of Waldo Pond, Marquette County, on July 15, 196*1. The examination of the contents of two cocoons revealed no developing young. An 123 attempt was made to continue the development of the other three in the laboratory without success. J. P. Moore (1923) found cocoons of Macrobdella decora in June and July in New York. The eggs hatched in about three weeks and young were frequently found in July and August. Erpobdellidae: The erpobdellids, like the glossiphonids, produce spermatophores which are deposited by one leech onto another. The cocoons produced are flattened and are cemented to the undersurface of submerged rocks and water­ logged wood. The cocoons are deserted by the parents after deposition. Twenty cocoons of Nephelopsis obscura were collected from a bayou of Lake Superior, Marquette County, on August 7, 1967. Examination of these cocoons showed a range in the number of young from 3-17, averaging seven young per cocoon. It is suspected that the number of young per cocoon was probably closer to 17 as some young were partial­ ly out of some cocoons and others had probably left. It could not be determined whether a given adult had deposited one or several cocoons. The young leave the cocoon by way of the two papillae on the ends where the cocoon has been sealed. The cocoon does not rupture releasing all the young at one time. It was also noted that different cocoons contained young in varied stages of development, whereas all the young in a given cocoon were similar in appearance. J. E. Moore (196^) found N. obscura young from May 5 to 12H August 25 in Alberta. Sapkarev (1967) found young of this leech in July and August in Lake Mendota. Therefore this leech reproduces during the spring and summer months and might produce more than one brood a year. Although spent cocoons of Erpobdella punctata were commonly found attached to submerged rocks and wood, none were found with eggs or developing young in them. As leech cocoons do not deteriorate rapidly, it was impossible to determine when they had been deposited. J. E. Moore (1966) states that the breeding habits of E. punctata are similar to those of N. obscura, commencing in May and continuing through much of the summer. Sapkarev (1967) found cocoons with eggs from May through August in Lake Mendota. The number of eggs per cocoon varied with most containing 2-6. He found that most young leave the cocoon in July and that young appear in July and August. No information was ob­ tained concerning the reproductive habits of Dina fervida or Dina p arva. In summary the glossiphonids are prolific and exhibit a high degree of parental care, whereas the hirudids and erpobdellids are probably not as prolific nor do they pro­ vide any parental care. These two extremes regarding parental care could affect the abundance and distribution of these three families of leeches. One might expect that a higher mortality rate would occur among the cocoons and young of the hirudids and erpobdellids. The cocoons, however, possess relatively the venters of the adults other ecological factors become involved, such as transport by turtles, frogs, and pos­ sibly waterfowl, movement to areas of abundant food, etc. These cause a dispersion of glossiphonids throughout a body of water and also to other waters. Consequently, most members of this family are abundant and widely distributed in Michigan waters. The hirudids and erpobdellids, although probably not as prolific as the glossiphonids, possess individual charac­ teristics which enable most of them to survive in substantial numbers and become widely distributed in Michigan. CHAPTER V SUMMARY AND CONCLUSIONS Summary A study attempting to correlate several environmen­ tal factors with the abundance and distribution of leeches in Michigan was conducted from 1961 through 1966. A few observations made in 1967 and 1968 were included. Fifty bodies of water were examined in Marquette County in the Upper Peninsula of Michigan and six each in Barry and Kalamazoo Counties in the Lower Peninsula. A comparison was made between the leeches found in the two peninsulas. A comparison was also made between the leech fauna of Michigan and that of Ohio, Illinois, Minnesota, and Wisconsin. A representative number of the waters studied were tested for pH and total alkalinity and their effect on the abundance and distribution of leeches was considered. The temperature, color, depth, bottom compo­ sition, and soil type of these waters were determined and their effect on leech distribution was discussed. The effects of the feeding habits, enemies, and reproductive habits of leeches on their abundance and distribution were determined. 127 128 Conclusions 1. Nineteen species of leeches were taken from the 62 waters examined in Michigan. This brings the number of leeches known to inhabit Michigan waters to 22. 2. The leech fauna of the Upper Peninsula compares closely with that of the Lower Peninsula. Placobdella hol- lensis, Illinobdella alba, I . p unctata, Pisclcola m i l n e r i , and Nephelopsis obscura were found in the Upper Peninsula, but not in the Lower Peninsula. Placobdella p i c t a , Haemopls lateralis, and Philobdella gracilis have been found in the Lower Peninsula, but not in the Upper. The leech fauna of Michigan compares closely with that of Ohio, Illinois, Minnesota, and Wisconsin. 3. Helobdella stagnalis was the most abundant leech found during this study, while Erpobdella punctata was the most frequently encountered species. 4. Most of the leeches in the four families, Glossi- phonidae, Piscicolidae, Hirudidae, and Erpobdellidae, found in Michigan were widely distributed in the parts of the state studied. 5. The glossiphonids were most frequently encountered in waters having a total alkalinity of lOOppm. or more. The hirudids, Haemopls marmorata and H. grand!s were most frequently encountered in waters with a total alkalinity of less than lOOppm. In contrast, Macrobdella decora showed no preference in relation to total alkalinity, whereas Dina 129 fervida and Nephelopsis obscura occurred more frequently in waters with a total alkalinity of less than lOOppm. 6. The abundance and distribution of the leeches studied appeared to be restricted by waters with low pH. Glossiphonia complanata, Placobdella parasitica, Placobdel­ la hollensis, Haemopls grandls, Erpobdella punctata, and Nephelopsis obscura were found in waters with a pH below 6.0; however, they were not abundant in these waters. Waters with pH readings as high as 9.3 have no apparent effect on either the abundance or distribution of the leeches studied. 7. Water current is a major limiting factor to the abundance and distribution of the leeches studied. were found in extremely fast moving waters. None The glossi­ phonids are best able to withstand moderate currents. Haemopls grandls, H. marmorata, and Dina parva were common­ ly found in lotic waters, but primarily in the eddies. 8. Nephelopsis obscura seems to be restricted in its distribution by high water temperature, whereas Philobdella gracilis ture. is possibly restricted by low water tempera­ All the other leeches encountered were found over a wide range of temperatures and seemed unaffected. 9. depths The leeches studied were most abundant at ranging from .6-1.2 meters. present beyond this depth. They are undoubtedly They are probably restricted at great depths, however, by a lack of vegetation, absence of substratum for attachment, and low oxygen content. 1 3 0 10. Most of the' leeches studied were found along a variety of bottoms, with no preference for any particular bottom shown. The important requirement for their presence seems to be the existence of submerged objects. 11. The soils surrounding a body of water may indirectly influence the abundance and distribution of leeches by altering the chemical contents of the water. 12. The reproductive habits of the leeches studied undoubtedly affect their abundance and distribution. The glossiphonids are very prolific and exhibit parental care and as a group were the most abundant leeches found during this study. 13. Parasites undoubtedly have no effect on the abundance and distribution of leeches. Predators may exert a minor effect especially in regard to the erpobdel­ lids. 14. The feeding habits of most leeches undoubtedly exert the greatest effect on their abundance and distribu­ tion. It is conceivable that all of the other factors con­ sidered have only an indirect effect in that they influence the abundance and distribution of the food organisms of leeches. SELECTED BIBLIOGRAPHY Bennike, S. A. B. 19**3. Contributions to the ecology biology of Danish freshwater leeches. Folia limnol. scand. 2:1-109. Bere, R. 1931. Wisconsin. Leeches from the lakes of northeastern Trans. Wise. Acad. Sci. Arts Lett. 26:437- ^ 0 . Fassett, Norman C. I960. A Manual of Aquatic Plants . The University of Wisconsin Press. Keith, M. M. 195*1. A survey of the leeches (Hirudinea) of the Duluth area. Proc. M i n n . A c a d . S c i . 22:91-92. ___________ . 1955. Notes on some leeches (Hirudinea) from the Yukon Territory, Canada, and Alaska. Proc. Minn. A c a d . S c i . 23:103-10*1. . I960. A simplified key to the leeches of Minnesota. P r o c . Minn. Acad. S c i . 27:190-199. Leverett, F. and F. Taylor. 1915. The Pleistocene of Indiana and Michigan and the History of the Great Lakes. Government Printing Office, Washington, D.C. ___________ . 1929. Moraines and Shorelines of the Lake Superior Basin. Government Printing Office, Washington, D.C. Mann, K. H. 1955* The ecology of the British freshwater leeches. J. A n i m . E c o l . 2*1:98-119. ___________ . 1957a. A study of a population of the leech Glossiphonia complanata. J. Anim. E c o l . 26:99-111. ___________ . 1957b. The breeding, growth and age structure of a population of the leech Helobdella 'stagnails. J. A n i m . E c o l . 26:171-177. ___________ . 1962. Leeches (Hirudinea): Physiology, Ecology and Embryology. London and New York. Their Structure, Pergamon Press, _____ . 1-96*1. ,A Key to the British Freshwater Leeches with Notes on Their Ecology. Freshwater Biological Assoc. Sci. Publ. N o T iPT 131 132 Mather, C. K. 1963. Haemopls latero-maculatum, new species. Amer. M i d . N a t u r . 70:168-174. Meyer, M. C. Quebec. 1937* Notes on some leeches from Ontario and Can. Field Natur. 51:117-118. 1940. A revision of the leeches (Piscicolidae) living on freshwater fishes of North America. Trans. Am. Micro. S o c . 59:354-376. ____ . 1946. Further notes on the leeches (Piscicolid a e ) living on freshwater fishes of North America. T r a n s . Am. Micro. Soc. 65:237-249. __________ . 1954. The larger animal parasites of the freshwater fishes of Maine. Maine Fishery Research and Management Bulletin 1:36-37* Meyer, M.C. and J. P. Moore. 1954. Notes on Canadian leeches. Wasmann J. B i o l . 12:63-96. Miller, J. A. 1929. The Leeches of O h i o . Ohio State University, the Franz Theodore Stone Laboratory, Contribution No. 2. __________ . 1937. A study of the leeches of Michigan, with key to orders, suborders and species. The Ohio Jour. Sci. 37(2):85-90. Moore, J. E. Alberta. 1964. Notes on the leeches (Hirudinea) of N a t u r . H i s t . P a p . N a t . M u s . Can. 27:1-15. _____ . 1966a. Further notes on Alberta leeches (Hirudinea). N a t u r . H i s t . P a p . N a t . M u s . C a n . 32:1-11. __________ . 1966b. New records of leeches (Hirudinea) for Saskatchewan. Can. Field Natur. 80:59-60. Moore, J. P. 1901. Lab. Nat. Hist. The Hirudinea of Illinois. 5:479-546. Bull. I l l . . 1905. Hirudinea and oligochaeta collected in the Great Lakes Region. B u l l . U. S. Fish B u r . 26:155. 1922. Southern Canada. The freshwater leeches (Hirudinea) of Can. Field Natur. 36:6-11. 1923. The control of blood-sucking leeches with an account of the leeches of Palisades Interstate Park. Roosevelt Wildlife Bull. 2:1-53. . 1924. The leeches (Hirudinea) of Lake Nipegon. Univ. Toronto Studies Biol. Ser. 25:15-31. 133 Moore, J. P. 1936. The leeches of Lake Nipissing. Field N a t u r . 50:112-114. Can. ........ 1959. Hirudinea. In ' F resh-wafer Biology, 2nd edition. John Wiley & Sons, Inc., New York, pp. 542-557. Moore, J. P., H. F. Nachtrieb and E. E. Hemingway. 1912. The leeches of Minnesota. G e o l . and N a t . H i s t . S u r v . Zool. S e r . 5:1-150. Moore, J. P. and from Alaska 9:11-17. M. C. Meyer. 1951. Leeches (Hirudinea) and adjacent waters. Wasmann J. Biol. Mullen, C. 1926. Some observations on the habits of leeches. Proc. Iowa Acad. S c i . 32:415-417. Oliver, D. R. 1958. The leeches (Hirudinea) of Saskat­ chewan. C a n . Field N a tur. 72(4):l6l-l65. Pawlowski, L. K. 1936. Zur Okologie der Hirudineen-fauna der Wigryseen. A r c h . Hydrobiol. R y b a c t . 10:1-47. Pennak, Robert. 1953. Fresh Water Invertebrates of the United States. Ronald Press, New York. Richardson, L. R. 1942. Observations on the migratory ■ behavior of leeches. C a n . Field N a t u r . 56:67-70. ___________. 1943* ?he freshwater leeches of Prince Edward Island and the problem of the distribution of leeches. ,Can. Field Natur. 57:89-91. Roelofs, E. W. 1944. Water Soils in Relation to Lake Pro­ ductivity . Tech. Bull. 190, Mich. State Agr. Exp. Sta. Ruettner, Franz. 1963. Fundamentals of Limnology. University of Toronto Press, Toronto. Rupp, R. S. and M. C. Meyer. 1954. Mortality among brook trout Salvelinus fontinalis, resulting from attacks of freshwater leeches. Copeia. 4:294-295. Sandner, H. 1951. Badania nad Fauna Pijawek. O ecol. U n i v . L o d z . 4:1-50. Acta Zool. Sapkarev, J. A. 19 6 7 . The taxonomy and ecology of leeches (Hirudinea) of Lake Mendota, Wisconsin. T r ans. W i s e . Acad. Sci. Arts L e t t . 56:225-253. Scott, I.D. 1921. Inland lakes of Michigan. Biol. Surv. P u b l . 30, G e o l . S e r . 25. Mich. Geol. 134 Sooter, C. 1937. Leeches infesting young waterfowl in northwest Iowa. J o u r . Parasit. 23(1):108-109. Taube, C. M. 1966. Leeches. Michigan Dept. Cons. Res. and Devel. Report No. 55* Thompson, D. H. 1927. An epidemic of leeches on fishes in Rock River. B u l l . N a t . Sur. 1 1 1 . 17: Art. 3. Veatch, J. 0. 1953. Soils and Land of Michigan. Michigan State College Press. Welch, Paul S. 1948. Llmnological Methods. Book Company, New York. ___________ . 1952.Limnology. New York. The McGraw-Hill McGraw-Hill Book Company, APPENDIX 135 Table Al. Class: A Taxonomic Classification of Michigan Leeches Adapted from J. P. Moore (1959). Hirudinea Lamarck, Order 1818. Rhynchobdellida Family Genus Glossiphonidae Vaillant, 1890. Glossiphonia Johnston, 1816. Glossiphonia complanata (Linnaeus, 1758). Genus Helobdella Blanchard, 1 8 7 6 . Helobdella stagnalis (Linnaeus, 1758). Helobdella fusca (Castle, 1900). Genus Theromyzon Philippi, 1867. Theromyzon rude (Baird, 1863). Genus Placobdella Blanchard, 1893- Placobdella montifera Moore, 1912. Placobdella hollensis (Whitman, 1872). Placobdella parasitica (Say, 1824). Placobdella picta (Verrill, 1872). Placobdella rugosa (Verrill, 1872). Family Genus Piscicolidae Johnston, 1 8 6 5 . Illinobdella Meyer, 1940. Illinobdella alba Meyer, 1940. Illinobdella punctata Meyer, 1940. Genus Piscicola Blainville, 1818. Piscicola milneri (Verrill, 1871). 136 ^ Order Table A1 (cont’d.) Gnathobdellida Family Genus Hirudidae Macrobdella Verrill, 1872. Macrobdella decora (Say, 1824). Genus Haemopls Savlgny, 1820. Haemopls grandls (Verrill, 1874). Haemopls plumbeus Moore, 1912. Haemopls marmorata (Say, 1824). Haemopls lateralis Genus (Say, 1824). Phllobdella Verrill, 1874. Phllobdella gracilis Moore, 1901. Order Pharyngobdellida Family Genus Erpobdellidae Erpobdella Blainville, 1818. Erpobdella punctata (Leidy, 1870). Genus Dina Blanchard, 1892. Dina parva Moore, 1912. Dina fervlda (Verrill, 1874). Genus Nephelopsis Verrill, 1872. Nephelopsis obscura Verrill, 1872. 137 Table A2. 1. A Key to the Leeches of Michigan. Mouth a small pore in the anterior sucker, from which a muscular proboscis may be protruded; jaws absent. Order Rhynchobdellida, 3 Mouth large occupying entire cavity of the anterior sucker, no proboscis 2. ............................ 2 Five pairs of eyes arranged in a submarginal arch (Fig.Al A); jaws present or absent. Order Gnathobdellida Hirudidae, 1*1 Three or four pairs of eyes in separate labial and buccal groups (Fig.Al B,C,D); jaws absent. Order Pharyngobdellida Erpobdellidae, 3. Body flattened and not divided 19 into anterior and posterior regions, anterior sucker confluent with the body .................. Glossiphonidae, 4 Body cylindrical, usually divided into anterior and posterior regions, anterior sucker distinctly separated from the body (Fig.A2 A) Pipcicolidae, 4. One or more pairs of eyes well (Fig.Al E,F,G) separated ................................. One pair of eyes close together (Fig.Al H) 5. One pair of eyes ........ ..................................... Three or four pairs of eyes 12 ........................ 5 8 6 7 138 Table A2 (cont'd.) 6. A conspicuous dorsal horny scute (Fig.A2 B); no distinct dorsal papillae Helobdella stagnalis No dorsal scute, three dorsal longitudinal rows of papillae 7. (Fig.A2 C) Three pairs of eyes ......... Helobdella fusca (Fig.Al E); two longitudinal dorsal and ventral b a n d s . Glossiphonia complanata Four pairs of eyes (Fig.Al G) 8. .... Theromyzon rude Anterior segments widened to form a head-like structure, three dorsal longitudinal keels (Fig.A2 D) ............. Placobdella montifera Anterior segments not widened, no dorsal keels 9. .... 9 Papillae prominent, abundant, rough, and pointed. Placobdella rugosa Papillae small, few, smooth, and rounded 10. Accessory eyes present .......... 10 (Fig.Al I). Placobdella hollensis No accessory eyes present 11. ........................... Color variable, usually dark greenish-brown, ventrally striped, no rows of semicircular orange spots along the margin. Placobdella parasitica Dark greenish-brown, not ventrally striped, a row of semicircular orange spots along the lateral margins 12. .................... Placobdella picta Suckers distinctly marked off from the body, 11 139 Table A2 (cont'd.) pulsatile vesicles on the sides of the body (Fig.A2 E) ................. Piscicola milnerl Suckers not distinctly marked off from the body, pulsatile vesicles absent 13. .................... 13 Body divided into two regions of different width, a narrow anterior region and a wide posterior region (Fig.A2 F) Illinobdella punctata Body not divided into regions of different width. Illinobdella alba 14. Large jaws with 35 or 65 teeth on each jaw, not arranged in pairs ............................. Jaws small or absent, teeth arranged 15. in pairs .... 15 16 Thirty-five teeth on each jaw, dorsal surface brown with a continuous median orange stripe, ventral surface orange . Phllobdella gracilis Sixty-five teeth on each jaw, dorsal surface green with a median row of orange spots and lateral black spots, ventral surface orange. Macrobdella decora 16. Jaws small with few teeth Jaws and teeth absent 17. .......................... 17 ......................... 18 With 10-16 pairs of teeth on each jaw,coloration variable but generally black with gray flecks. Haemopls marmorata With 20-25 pairs of teeth on each jaw, color dark olive green, a dark median dorsal 140 Table A2 (cont'd.) longitudinal stripe ...... Haemopis lateralis 18. Lip narrow and arched, grayish in color with dark blotches, up to 300mm. long Haemopis grandis Lip broad and flat, grayish in color with few or no dark blotches, with a reddish or orange band along lateral margins 19. Haemopis plumbeus Three pairs of eyes, first pair largest (Fig.Al B); rows of longitudinal black spots on the dorsum Erpobdella punctata Pour pairs of eyes, no longitudinal black spots on the dorsum ..................................... 20. 20 Anterior and posterior two pairs of eyes arranged in parallel (Fig.Al C); coloration gray, spotted with black, up to 100mm. long. Nephelopsis obscura Lateral eyes of each pair arranged slightly posterior to the medial eyes (Pig.Al D); reddish in color, up to 50mm. long .......... 21. Gonophores separated by 3 to 3*5 annuli, small, may attain a length of 25mm........ Dina parva Gonophores separated by two annuli (Fig.A2 G); may attain a length of 50mm.......... Dina fervida 21 141 C B Hirudidae Nephelopsis obscura Erpobdella punctata F Dina spp. Glossiphonia complanata Helobdella spp. acc. eyes H Theromyzon rude Fig. Al. Placobdella spp. Placobdella hollensis The eye patterns of several leeches. 0% E Fig. A2. Structure of leeches. G A, separated sucker of piscicolids; B, horny scute of Helobdella stagn a lis; C, dorsal papillae of Helobdella f u s c a ; D, head-like structure of Placobdella montifera; E, pulsatile vesicles of Piscicola milneri; F, narrow anterior region of Illinobdella punctata; G, gonophores and annuli of Dina fervida. Table A3. Marquette County Stations, Their Location, Dates Sampled, Chemistry, Species, and Number Collected. 4> 4> i—i CO co 4.5 O Xa rH U <; co cj rH CO Stations S w a o a H Dates 1 Redberry Pond T48N, R25W, S8 9/23/61 11/11/61 7.4 24 2 Harlow Lake T49N, R25W, S19 9/23/61 4/25/63 9/22/63 7.3 60 3 Waldo Pond T48N, R25W, S8 9/23/61 7/ 6/63 4/22/64 5/10/64 7.3 23 4 Sauxhead Lake T50N, R26W, S20 10/1/61 10/10/65 7.3 48 5 Independence L. T51N, R27W, S22 10/ 6/61 9/22/63 10/10/65 7.6 65 6 Kawbawgam Lake T47N, R23W, S18 10/13/61 7.4 CO CO •H 4J (3 CO CO O s rH 45 CO rH a rH 3 t I a •s CO 43 CO § o o a rH rH U 1 cO 3 •H cO rH r—1 CO rH s 3 CO ex •o o CO 43 0] 44 o 3 CO rH MH 0) 0) x PC CO CO cO a 3 rH •H rH iH rH l— 1 CO rH 4-1rH O N 3 3 at iH 3 fr •3 a> •3 CO *3 S a> 41 rH 43 3 43 5 O rH O >H O s 3 a) 45 H O O U 3 CO 45 c0 a rH P* H Oh a 3 rH (U 3 rH rH 3 rH rH 3 3 X) 3 •3 3 43 43 O O 3 O « s 43 3 3 •H rH tH rH 3 rH rH H rH M 3 3 rH rH 44 3 •H 3 rH |H 3 3 3 44 o 3 *3 V4 •H U o 3 43 O a s •H rH O a O 3 3 U •H a 3 e a *3 1 iH 3 PH 3 X 3 •H 3 3 3 3 tH 3 XJ •H 3 •3 •H 3 •3 O 3 O > rH O >1 3 3 3 O s 43 3 .+*1 M a 3 O 3 31 3| 3 3 45 45 SI Ol Sl«w <3 O « 3 6 3 •HI -H 3 3 w p| a 55 3 44 3 3 3 rH 44 +4 rH 3 O 3 44 a X 13 10 a O Table A3 (Cont'd.) 4J 3 CO O U <0 X o. u 6 o. o* V-/ Stations Dates 7 Road 510 Pond T49N, R27W, S23 10/14/61 6.9 8 Mangum Rd. Ditch 10/16/61 6.5 T47N, R24W, S22 9 Bagdad Pond T48N, R26W, S24 10/22/61 4/18/64 6/25/64 7.3 105 10 Park Cem. Pond T48N, R25W, S14 11/ 5/61 7.0 65 11 Rush Lake T52N, R29W, S24 5/ 6/62 7.3 29 12 Howe Lake T52N, R29W, S23 5/6/62 7.1 24 13 Mine Shaft Pond T47N, R29W, S32 5/13/62 cd to 3 Cd u tn cd o m rH o a o o ' • o| 3 «4H pi | pp| CO iH CO s 4 ex nil Si x\ i —i H o X h | u •H HI •H CO CO Vl CO a h | co CO o at 3 Vl h | CO rO 1—1 CO m | O HCI co HI CJ 3 3 a m | •H Vi ai 3 a 3 Vi O CJ 0J *3 H| SI tH •H CO •H •3 3 3 Vi at tc| 3 HI 3 Vi O s c 3 H3I H3I CJ 3 3 a ECI M| 0 3| > M 3 Oj 3 * 3l i> Vi 3 H 3 H 3 a CO o Q| Q| 55| 2 62 2 10 1 M -Cr VJ1 29 8 2 1 6 1 1 8 6 1 4 18 1 Table A3 (cont'd.) •H p CO •H I— I ««-N « CO AS o 23 P. rH U ■< (0 u tH CO Q ■W P. Stations o p. H Dates 19 Small Ponds 4/27/63 T48N, R25W, Sll 20 Vernal Pond 5/ 4/63 T47N, R27W, S18 21 Small Creek 4/26/63 T51N, R27W, S36 22 7.5 13 Cooper Lake 8/28/63 T48N, R27W, S32 7.7 65 23 Bancroft Lake T47N, R27W, S3 7.7 95 24 Boston Lake 9/ 1/63 T48N, R28W, S32 25 Tilden Lake 9/ 5/63 T47N, R27W, S23 7/ 5/64 8/28/63 •U CO p CO r—I CO •H rH Cfl P 8 03 CJ 4J CO CO u CO S3 <4H 3 M CO •H CO p CD rH tH o .P Ol «| 5C| H| PX| P ex co Q) CJ •H 4J •H CO CO M CO P CO CO o CX 3 M CO ,P rH CO CO ■U CO 4J CJ P 3 P H| M| H| tH M 0) P iH •H e cfl M O u | Ml P p| PS3| EC| W| Ol •o ex o s o CO -H > M CD CM p M 3 CJ CO JO Q| 53| l-J -Cr G\ 55 6.9 30 o 1 1 Table A3 ( c o n t 'd .) >> in PC cu Stations Dates rH <3 m O CJ co O C O a ■u p . rH cfl q cfl rH O s O O CO •H rH CO B 6C cO 4J CO 7.3 57 27 Ogden Lake 9/ 5/63 T47N, R27W, S13 10/ 5/65 7.1 47 28 Fish Lake T47N, R29W, S5 9/ 8/63 50 29 Bacon Lake T47N, R27W, S3 9/ 8/63 70 30 Morgan Creek T48N, R26W, S26 9/14/63 7.6 90 31 Ramseth Pond T48N, R25W, S20 9/14/63 7/17/64 5/ 8/64 7.6 85 32 Pond near Humboldt T48N, R28W, S31 9/15/63 61 3 u -i • 10 Schoolhouse L. 9/5/62 T47N, R27W, S23 10/ 5/65 cd o 10 PB| ffi| 26 CO • H 3 c0 IH to iH H co U s -\ H S •H r—I co A! B a> (U T3 rH rH Vi Xi H | H | 3 •H Vl a) B c0 co O c 3>4 o N CO Vi rH CO P n| Mh | B Ph | SI iH •H •o B cO Vi o CJ CU •3 M M| CO CO Vi 0( 331 3 3 1 1 CO B IH co IH CJ B 3 B 33*| W| Q| 3 1 CO Vi o p Vi cO s fervlda •H Q| 55| 1 5 2 Table A3 (cont'd.) c CO ft CO ft cd tH ft 0 O u as •H tH CO ft CX CO 4J CO Cd o CO ft m ft •ft ft ft H "2? . o l pj| pj| H| •H r—i s Cd CO AS O tH U 32 ft, Hj CO O co a jj a, rH Stations Dates 33 Lake Superior T48N, R25W, S23 9/25/63 5/ 5/66 8.1 75 34 Mountain Lake T52N, R28W, S31 9/28/63 6.9 55 35 Pine Lake T52N, R28W, S21 9/28/63 36 Goose Lake T47N, R26W, S15 1/ 2/64 1/25/64 2/15/64 37 Rod & Gun Club Pond T48N, R25W, S2 5/ 5/64 38 Wetmore Pond T49N, R26W, S31 39 West Branch Creek T46N, R28W, S20 6/28/64 as •H CO ft 0) tH >H O cd o •H ft Ti as cd ft cd ft cd as o cx ■ft ft ft| ft| CO iH cd cd u CO ft o ft ft ft •H ft 0) ft tH •H M| H| s CO ft o ft Q) •ft CO •H •ft ft CO ft cx ftl| g| P2i| cd ft cd ft o pft CO s CO ft co ft ft ft ■ft ft CO > ft col ftl ft -H > ft 0) «ft cd ft ft ft CO o P2| W| d| Q| !5| M -Cr CD 5/16/64 6.9 65 12 14 Table A3 (cont'd.) s a> ■H •H rH /*n a) cn M 03 a rH CO o c 64 CO 4J rj t> < CO rH u CO Q Stations •u a o p. H Dates 40 Tourist Park Pond T48N, R25W, S10 6/29/64 41 Dead River T49N, R25W, S9 7/ 7/64 9/16/65 7.2 Ol co 0 01 3 cn <4-1 3 J-i 33| M| H| 77 cn •w cn 3 n CO P m| M| W| 1 2 CO >4 O o 4 p CO •3 •H > M 0) M-4 a cn XI o d| Q| 55| CO p 3 -E=VO 42 Mine Shaft Pond T47N, R27W, S10 7/ 5/64 43 Carp River 7/8/64 T48N, R26W, S29 44 Dishno Creek 8/ 2/64 T49N, R29W, S29 45 Arfelin Lake 8/ 2/64 T49N, R30W, S21 46 Deer Lake 8/ T48N, R27W, S29 47 Barnhardt Creek T48N, R28W, SI 4/64 8/ 4/64 12 7.9 90 7.8 95 7.0 52 10 1 20 7 1 10 Table A3 (cont'd.) c co •H 1—I /-N cfl co .SC O flj p. rH CJ •< Cfl o tH CO 4-1 EOP. h w Stations Dates 48 Goldmine Creek T48N, R27W, S29 8/ 4/64 7.7 140 49 Middle Isle Point Bog T48N, R25W, S4 9/ 5/65 7.3 80 50 Chocolay River T47N, R24W, S6 11/ 4/65 6 p. 4-1 Cfl CO G « cfl o rH P e o o o 3 MH a) •3 3 Vi Si SC| Eh | pu| m rd| o 95 j d| cfl CD O Cfl 3 V4 cfll -“ | I rH Cfl| Cfl 4-> Cfl 4J U 3 3 P d| M| m | •H 8 cfl V4 o o 0) TJ CO •H -a c cfl Vl Cfl d| Si *1 iH )H Ul > H Barry County Stations, Their Location, Dates Sampled, Chemistry, Species, and Number Collected. CO 33 a cO •H 4J CO cn c CO o c O o 43 CO rH CO •H D o 0] e CO O CO u a o o o p. rH o H Stations Dates 1 Otis Lake T3N, R10W, S30 7/ 1/63 6/20/65 7/26/65 7.0 70 2 Pleasant Lake TIN, R9W, S8 6/22/63 7.5 45 3 Crooked Lake TIN, R10W, SI 6/29/63 7/22/65 8.5 115 4 Lawrence Lake TIN, R9W, S27 6/29/63 7/13/65 8.0 195 5 Purdy Lake TIN, R9W, S36 7/ 6/65 7/18/65 5.9 25 6 Strewins Lake TIN, R9W, S36 7/22/65 6.8 35 CO CO•H rH H i—1 CO 0) C3 •o 01 43 cd O 4J rH CO 0) 33 cd rH rH d) cd T3 o 43 CO O 3 rH M h (U 33 Cd cd CJ r—1 •H rH 4-t d) •H CO 43 CO M o u cd cd O rH cu cfl rH cd 4J cfl CO |H •H o O p o u e cd d) 6 cd PH 33 rH rH <0 CO CO 43 O O CX CJ 3 CO >H cd rH (0 H iH d> H TJ tH 43 U O CO rH M •H CX 43 PH 8 25 Erpobdella punctata Table A4. Kalamazoo County Stations, Their Location, Dates Sampled, Chemistry, Species and Number Collected. -H ts •H r—I Cfl CO as P. O rH U < <3 f—i to s 4-1 O . Stations Dates 1 Gull Lake T1S, R9W, S7 6/19/63 7/23/65 2 Augusta Creek T1S, R9W, S27 6/20/63 o a H 8.1 cO •H C O J3 p iH (0 os o I-C o CO 4J CO c CO rH a 0 o a CO cO *H i—t H rH CO a) c T> 01 X> cfl O 4-' rH CO 0) as co rH rH 0) c0 •u o CO O 3 rH <4H cu as co u •H 4-1 •H CO CO O Vl U CO CO P rl PU CO rH r-H 0) •o cO rH rH (1) •o .a o o CO H (X CO CO O W 3 Vi CO rH rH tS o a CH a Vl w co (S fervida Table A5. 177 i—1 V-CT ro 3 Kalamazoo River T2S, R10W, S23 7/27/65 7.0 200 4 Wintergreen Lake 4/10/65 4/20/65 T1S, R9W, S8 7/20/65 9.3 127 5 Gull Lake Creek T2S, R9W, S7 7/29/65 7/30/65 7.9 200 6 Marl Pond T1S, R9W, S6 7/14/63 8.5 250 12 3 17 12 153 Table A6. A. Habitats Compared In Regard to Total Alkalinity. Intermediate Standing Waters (18-59 ppm. CaCO^) Habitat ppm. Waldo Pond Howe Lake Redberry Pond Rush Lake Tilden Lake Strewlns Lake Ogden Lake Sauxhead Lake Pish Lake Boston Lake Mountain Lake Schoolhouse Lake B. Hard Standing Waters (60-99 ppm. CaCO^) Harlow Lake Horseshoe Pond Cooper Lake Independence Lake Pine Lake Bacon Lake Otis Lake 703,1 LcllCC Middle Island Point Bog Ramseth Pond Bancroft Lake Deer Lake C. 23 24 24 29 30 35 47 48 50 55 55 57 ’ 80 62 65 65 65 70 70 7^ 80 85 95 95 Hard Standing Waters (100-250 ppm. CaCO^) Bagdad Pond Crooked Lake Wintergreen Lake Gull Lake Lawrence Lake Marl Pond 195 115 127 177 195 250 154 Table A 7. Total Alkalinity Mean, Standard Deviation and Range for Eleven Species of Leeches. Both Lentic and Lotic Waters Considered. Species cr Range (ppm. CaCOg) No. of Samples Mean Glossiphonia complanata 16 88 47.5 24-200 Helobdella stagnalis 16 96 50.0 47-200 Placobdella parasitica 21 93 63.8 24-250 Placobdella rugosa 7 89 62.7 23-195 Macrobdella decora 7 88 71.1 23-250 Haemopis grandis 8 64 25.5 24-105 Haemopis marmorata 10 61 23.8 13- 95 Erpobdella punctata 20 83 52.4 23-200 Dina parva 10 73 31.8 24-140 Dina fervida 13 74 37.2 23-177 7 61 15.2 29- 80 Nephelopsis obscura ' 155 Table A8. Waters Tested and Their pH. Station pH Station pH Cranberry Bog 4.5 Rush Lake 7.3 Purdy Lake 5.9 Sauxhead Lake 7.3 Strewins Lake 6.8 Redberry Pond 7.4 Mountain Lake 6.9 Morgan Creek 7.6 Pine Lake 6.9 Ramseth Pond 7.6 Tilden Lake 6.9 Independence Lake 7.6 Otis Lake 7.0 Goldmine Creek 7.7 Kalamazoo River 7.0 Bancroft Lake 7.7 Barnhardt Creek 7.0 Cooper Lake 7.7 Horseshoe Pond 7.0 Deer Lake 7.8 Park Cemetery Pond 7.0 Gull Lake Creek 7.9 Ogden Lake 7.1 Carp River 7.9 Howe Lake 7.1 Lawrence Lake 8.0 Dead River 7.2 Gull Lake 8.1 Middle Island Point Bog 7.3 Lake Superior 8.1 Bagdad Pond 7.3 Teal Lake 8.5 Waldo Pond 7.3 Marl Pond 8.5 Harlow Lake 7.3 Crooked Lake 8.5 Schoolhouse Lake 7.3 Wintergreen Lake 9-3 Table A9. pH Mean, Standard Deviation and Range for Eleven Species of Leeches. Both Lentic and Lotic Waters Considered. Mean cr 17 7.4 .635 5.9-8.5 Helobdella stagnalis 13 7.6 .520 6 .9-8.5 Placobdella parasitica 20 7.6 .714 5.9-9.3 Placobdella rugosa 9 7.4 .413 6 .9- 8.1 Macrobdella decora 7 7-5 .497 6 .9- 8 .5 Haemopis grandis 7 7.0 1.076 4.5- 8.1 Haemopis marmorata 9 7.4 .453 6 .9-8.5 19 7.4 .934 4.5-9.3 8 7.5 .685 6 .5-8.5 11 7.5 .488 6 .9-8.5 6 7.0 2.968 4.5-8.5 Species No. of Samples Glossiphonia complanata Erpobdella punctata Dina parva Dina fervida Nephelopsis obscura Range - Table A10. The Occurrence of Eleven Species of Leeches Along Various Bottoms. —/ rN CM ' —/ i—1 / —\ CM v —* ✓—s ■=3" ---' ✓—s CM t- w • -P O / —s VO tH 3 CO -P •H G P 03 CO 6 o -p -p o P 03 3 -P •H PQ G -P 03 Q G 03 73 i—1 3 o CQ * X O o G 73 03 CO 03 I— 1 03 H O P •H G P 03 Q •» rH 03 JQ X> XI > X3 CC PG CC C5 XI 3 3 •\ 3 cd G CO 3 P to p P.cd rH 03 3 P. O (G CQ 03 3 p •H rH 03 G > cd G P 03 Q o 73 G n 0} 73 G cd CO CO CO CO Glossiphonia complanata 0 0 1 Helobdella stagnalis 0 1 Placobdella parasitica 0 3 2 0 Placobdella rugosa 0 2 0 Haemopis grandis 0 1 0 3 2 Haemopis marmorata 0 2 1 Macrobdella decora 0 0 Erpobdella punctata Dina fervida 0 0 Dina parva Nephelopsis obscura G cd 73 G cd 73 G cd 73 C cd 73 C cd CO i—I CO 3 G P P 03 •H Q P 03 G 01 3 Q P P cd a 03 03 cd a> P. O >> 03 3 G XI •H P •> cd 03 P. cd o >s •H •\ 3 O >» G cd O i—i •H (G o •> 73 G a 03 rH 3 •H G P P. i—I 3 CG p 0) Q •* X X o 3 S o 3 S 0 4 0 2 CO CO CO 3 2 0 0 0 0 0 0 1 3 2 0 0 1 0 0 0 0 0 3 2 3 0 0 1 0 0 0 0 1 2 0 2 0 1 0 0 0 0 1 0 0 1 0 1 0 0 1 0 1 1 0 0 2 2 2 0 1 1 0 0 0 0 0 1 1 1 2 0 0 0 0 0 0 0 0 1 0 2 3 2 1 1 1 2 4 2 0 0 3 0 0 0 0 0 0 1 1 0 0 0 4 2 0 0 1 2 0 1 1 1 0 1 0 0 0 0 0 0 1 1 0 1 1 3 0 2 0 0 0 0 1 1 0 0 0 3 .