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AN ECOLOGICAL SURVEY 



ISLE ROYALE, LAKE SUPERIOR 



PREPARED UNDER THE DIRECTION OF 



CHAS. C'ADAMS. 



A Heport from the University of Ulchigan Wtiseum, published liy ihc State Biolojjical 
Survey, as a part of the Report of the Board of llie 
Geological Sun'cy for I oris. 



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TABLE OF CONTENTS. 



Introductory Note and AcknowledgtueDts, bj Dr. Charles C. Adams, 

PART I. ECOLOGICAL PAPrSB. 

Isle Roynte as a Biotic Environment, by Dr. Charies C. Adams 

I. Introduction , 

1. Itinerary and Peraonnet of the Party. 

2. Aims and Method of Work 

3. PKTiouB fiiolc^eal Inveatigationa upon Isle Royale 

4. Hiatorical Note 

5. Available Maps of Isle Royale 

II. The BioU Considered by Stations 

1. Location of Field Stations in 1005 

2. General Characteristics of the Stations 

III. The Evolution of the Gross Environment 

1 . Geological Succession 

2. Topography and its Origin 

8. Atmospbenc Influences and their Evohjtion..'. 

a. Climate 

b. Seiches 

c. Climatic Succession 

d. Lake Storms and their Influence 

4. Surface Currents of Lake Superior. 

5. Origin of the Habitats 

i The £cological Relations of the Invertebrate Fauna of Ue Royale, Michigan, 

by Dr. H. A, Gleason 

I. Introduction 

II. The Lake 

in. The Inland Lakes 

JV. The Tamarack Swamp and the Arbor Vitae Swamps 

V. The Gravel and Sand Beaches 

VI. The Rock Beach 

VII. The Cladonia Clearings and Jack Pine Ridges 

VIII. The Balflftm-S[inice Forest 

IX. Artificial Clearings 

X. Summary. 

3, Th« Ecological Distribution of the Birds of Isle Royale, Lake Superior, by 

Otto MdCreaiy 

I. Introduction 

II. Light^house Peninsula, (between Rock Harbor and the Head of Conglom- 
erate Bay) 

1. Lake Superior and Beach (Station I, Sub. 1.) 

2. Spruce and Balsam Forest (Station I, Sub. 2 and 3) 

3. Tamarack and Arbor Vitae Swamps (Station I, Sub. 4) 

4. Jack Pine Ridge (Station I, Sub. 5) 

5. Sphagnum and Spruce Bog (Station I, Sub. 6) 

e . Valley at the Head of Conglomerate Bay (Station I, Sub. 1) 

in. Trail to MeCargoe Cove 

1. Ransom Clearing (Station 11, Sub. 1) 

2. Benson Brook (Station II, Sub, 1) 

3. Spruce and Tamarack Swamps (Station II, Sub. 2 and 5) 

4. Rock Ridge Clearings (Station II, Sub. 3) 



207309 



IV. WeBteni End o! Rock Harbor and Trail to Sumaer Lake 

1. Harbor. (Vicinity ot Station III, Sub. 2) 

2. Small Islands (Station III, Sub. 1) 

3. Bulrush Zone and Delta (Station III, Sub. 3) 

4. Trail to Sumner Lake (SUtion III, Sub. 4) 

a. Birch Forest 

b. Birch and Coniferous Forest 

5. Sumner Lake {Station III, Sub. 5) 

V. Siskowit Lake Reiion 

1. Siskowit Bay and Shore (Station V, Sub. 1) 

2. Trail to Siskowit Lake (Station V, Sub. 4) 

3. Siskowit Lake (Station V, Sub. 6 and vicinity) 

4. Burning West of Outlet to Siskowit ha.ke (Station V, Wdnity of Sub. 9), 

5. Long and Menagerie Islands (Station V, Sub. 10) 

VI. Summary. 

1. Water Birds 

2. Shore Birds 

3. Birds Frequenting Swamps 

4. Birds of Clearings and Partial Clearings 

5. ffirds Frequenting the Forests 

4. The Fall Migration of Birds at Washington Harbor, Isle Royale, Lake Superior, 

by Max Minor Peet 

I. Introduction .' 

II. The Environment 

1. Tho Clearing 

2. The Forest 

3. Food 

III. The Weather Conditions and Migrants 

1 . Weather Conditions 

2, The Bird Migrants. 

a. Warblers 



b. Spar 

c. Hawl 



Hawks 108 

d. Owls 104 

e. Thrushes 104 

t. Other Birds 104 

IV. Large Bird Waves 105 

1. First Wave 106 

2. Second Wave 108 

3. Third Wave 107 

4. Fourth Wave 107 

5. Fifth Wave 109 

6. Sixth Wave 109 

V. The ReUtlon of Weather to Migration lOB 

1. Influence of Wind '. 110 

2. Influence of Temperature Ill 

3. Influence of Barometric Pressure Ill 

4. Condition of the Sky Ill 

5. Summary and Conclusion Ill 

VI. The Routes of Migration 112 

VII. The Perils of Migration 113 

1. Fatigue 113 

2. Natural Enemies 115 

3. Blunders and Fatalities 116 

. The Ecological Succession of Birds, by Dr. Chas. C. Adams 121 

I. Introduction 121 

II. Representative Literature on Habitats and Succession 123 

1. Habitat Preference 123 

2. Succession 126 

III. The Major Avian Environments 128 

IV. Minor Avian Environments and their Associations 133 

V. Avian Suecession 134 

1 . General Remarks 134 

2. Succession on Isle Royale 134 

a. The Aquatic Association and Habitat 136 



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LETTER OF TRANSMITTAL. 



To the Honorable flte Boai-d of Cieological Survey of tlie 8tHte of Micli- 
igau: 

Gov. Fred SI. Wanier, Tresideut. 
Hod. T*. 51. Forry, Jr., Viceri-esident, 
Hon. L. L. IVi-iglit, Secretary. 

<5eutlemeii : — I b^ to present berewitli for priutiug, a report by Dr. 
Cbiis. C AdauiB on the ecology, tkat it* the natural history, of I»)e Boyale. 
Thin conies to us with the approval of I>r. A. G. Kuthi'eii, our Chief Field 
Xatoi-alist. and oiir Board of Scientific AdviBers, and is u continuation 
of the work published in our annual report for 1005. 

This contribution to the Biological KuiTey of the State, which the 
leKittlatni'e authorized nie to Kupen'ise by Act No. 250 of the session of 
11105, comes fi-oni the ITuivei-sity Museum. The explorations were made 
without expense to the State Survey by means of contributions from 
frien<38 of the Museum. As tliis wovk is in hamiony with the aims of 
the niological Hui'\'ey we ai-e fortunate in securing such co-operation. 
The i^jwrts on the Porcupine Monntains and Isle Royale at the north 
eod of the state complement the work on Walnut I-ake, Oakland county, 
and ^hat in Huron and Tuscola counties. 

I trust that the present reiwrt will be of service to the schools of the 
state. 

Vei-y respect fullv, 

AI.FRED C. LANE, 

State Geologist. 



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ILLUSTRATIONS. 



Fig. 1. The Light-house at Rock Harbor, lele Roy^e 

2. Cliffs between Tonkin and Conglomenite Bays. 

3. Summer storm waves upon the be&ch (I. 1) nt the liead of Tonkin Bay, 

south of the Lkht-house 

4. Sand beach at the head of Conglomerate Bay (1, 1) 

5. Beach pool (1, 1) near Tonkin Bay 

6. Transition from the beach (I, 1) to rock clearing (I, 2), south ot the light- 

7. Natural rock opening (I, 2) or avenue, farther up tbe elope than in Fi|. 6. 

8. Natural rock clearing or opening (1, 2) north ot the Light-house at Rock 
Harbor 

9. Natural rock clearioK (t, 2) south of the Light-house 

10. Aibor Vitae b<^ (I, 4) near Tonkin Bay 

11. View from the Jack Pine ridge (I, S), lookmg toward the head of Con- 
glomerate Bay 

12. &cond growth ot White Birch on the Trail to the Jack Pine Ridge (I, 5), 
Conglomerate Bay 

13. Jack Pine ridge (I, 5) Conglomerate Bay 

14. Sphagnum-Black Spruce bog (I, 6) near the Jack Pme ridge 

15. Small island near the head of Rock Harbor {III, 1) 

16. Buhwh >one and delto at the bead of Rock Haihor (lU, 3) 

17. Exposed section of spit formed as the water level has lowered in Rock Har- 
bor, near the bonnning of the trail to Sumner Lake (III, 4) 

18. Sumner Lake (in, 5) 

19. Western end of Sumner Lake (III, 5} 

20. Northeastern margin of Sumner Lake (III, 6) 

21. Southeastern comer of Sumner Lake (III, 5) 

22. Western end of Sumner Lake (IIL 5) 

23. Northern shore of Sumner Lake (III, 5) 

24. Rock opening about Camp on Siskowit Bay (V, 3) 

26. Rock opening at the Siskowit camp (V, 3) 

26. Rock opening on Siskowit Bay (V, 3) 

27. Border of the miening about the Siskowit Camp (V, 3), near the beginning 
of the trail to Siskowit Lake (V, 4) 

28. Ant nest in the opening at the Siskowit Camp (V, 3).. 

29. Genera] character of the south shore, near the eastern entrance to Siskowit 
Bay (V, 2) 

30. Rock pool on the beach (V, 2), where a variety of invertebrates was 
secured 

31. Saxifmga aizoon on beach (V, 2) 

32. General view along the shore at V, 2 

33. Farther up the same slope as Fitr. 32 and adjacent to it 

34. Still farther up the slope and adjacent to Fig. S3 

35. Looking up the slope on the western portion of Station V, 2 

36. Upper portion of western part of Station V, 2 

37. Detail of western part of Station V, 2 

38. Character of ground cover in parts of the Balsam-Spruce forest (V, 4) 

39. Open space in the Balsam-Birch forest (V, 4) 

40. Open space in the Balsam-Bircb forest (V, 4) 

41. Tamarack Swamp (V. 5) 

42. Spruce margin of Station V. 6 »olS 



43. £fiaok Spruce margin of Station V, 5. . . 



ILLUSTRATIONS. 

Page. 

44. Bog margLn of Station V, 5 16 

45. Long Island gull rookeiy, (V, 10) 16 

46. Pond in Tamarack— Black Spruce swamp (V, 11) 16 

47. Margin ot Lily pond (V, 11) 16 

4«. Black Spruce in Cassandra i one of Station (V, 11) I« 

49. Maple forest on the Desor trail (III, '04) 16 

50. Forest along Washington Brook (IV, ' 04) 16 

51. Showing origin of the glacial Great Lakes, their relation to the ice sheet 
and their Mississippi drainage. (After Taylor and Leverett). Cham- 
berlin and Salisbury, Geology, III, p. 396. Fie. 516 33 

5'2, Showing the Algonquin stafe of the Great Lakea; a wal«r barrier to north- 
ward dispersal ot the land biota. (After Taylor and Leverett). Cham- 
berlain and Salisbury Geology, III, p. 401. Fig. 521 34 

53. Contour Map of Isle Koyale, Michigan. Contour interval 100 feet. Pre- 

fared by Dr. A. C. Lane 34 
he Nipissing Great Lakes; showing the fresh-water highway or barrier 

in the west and the sea barrier in the east. (After Taylor). Chamber- 36 

lin and Salisbury, Geology, III, p. 404. Fie. 522 

55. Surface currents of Lake Superior. To show their possible influence on 

the origin of the biota. (Drawn by Hall, after Harrineton.) 56 

56. "The Wetidigo Road", from the clearing at the ciub-house to Wendigo, 
Washington Harbor 56 

57. LonglHland(V, 10), Siskowit Bay, looking toward late RoyaleLight-HoUBe. 56 

58. Gulfrookery on Long Island (V, 10) 56 

59. Gull rookery on Long lahind (V, 10) 5ft 

60. Eagle nest at Tobin Harbor (IV, 8) 56 

61. Variations in the shell width of Potygyra aUmlahrit 298 

62. Variation in the shell height of Pol^gyra aSn>labria 298 

63. Lymnaea slagnalU varieties from Isle Royale 298 



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, CONTENi-S. vii 

Pagp. 

b. The Shore and. Marsh Association and Habitat 138 

c. Bc^forest Association and Habitat 139 

d. Aspen-birch Association and Habitat 139 

e. The Climax Association or Formation and Habitat 141 

3. Interna! Factors 142 

4. Environmental Factors 143 

5. Environmental and Associational Convei^nce 144 

6. Succession and Environmental Evolution 145 

7. The Relation of Succession to Organic Evolution 145 

VT. Some Principles of Succession 146 

VII. Some AdvantBj^eg of a Knowledge of the Laws of Succession 149 

. The Coleoptera of Isle Royale, Lake Superior, and their Relation to the North 

American Centers of Dispersal, by Dr. Chas. C. Adams 157 

I. Introductory Note 167 

II. Notes on the Habitat Relations of Beetles. 158 

III. The Succession of Beetle Associations 160 

1. The Lake Shore and its Beetle Associations. 160 

2. Rock Openings and Associated Beetles 160 

3. Lake, Fond and Bog Habitats and Associations 161 

4. The Forests and their Beetle Associations 161 

IV. The General Characteristics of the North American Beetle Fauna 163 

1. Compiled Generaliiations on the Fauna 163 

2, Comments on the Preceding Generalizations, and on the Literature of 
Gec^raphic Distribution 182 

V. The Present Centers of Dispersal of the Beetle Fauna 183 

VI. The General Characteristics and Affinities of the Isle Royale Fauna 190 

1. Faunal Characteristics 190 

2. MisceLaneouB Notes on the Fauna 191 

VII. Lists of Isle Royale Beetles 192 

1. List of Species Collected in 1905 192 

2. SupplemenUry List of Isle Royrie Beetles, by A. B. Wolcott 204 



. Notes on the Vwetation of Isle Royrie, Michi^n, by W. P. Holt 217 

I. General Observations on the Plant Sooeties 217 



2. Shore Vegetation 222 

3. Forests 224 

4. Burnings 225 

II. Annotated List of Plants 227 

2, Annotated List of Certain Isle Royale Invertebrates, by Dr. Chas. C. Adams — 249 

3. Annotated List of the Mollusc* of Isle Royale, Michigan, by Biyant Walker 281 

I . Introduction 281 

II. Faunai Affinities 281 

UI. Annotated List 283 

4.. Report on the Isle Royale Orthoptera of the 1905 Expedition to Isle Royale, 

by A. P-Morse 299 

I, General Remarks 299 

II. Annotated List of Species 302 

III. Station List, 1905 Collections 

5. Neuropteniid Insects from Isle Royale, Michigan, by Dr. James G. Needham 30S 

6. Diplera of the 1905 University Miiseum Expedition to Isle Royale, by Prof. 

James S. Hine 308 

7. AnnoUted List of Isle Royale Hymenoptera. by E. S. Titus 317 

8. The Ants of Isle Royale, Michigan, by Dr. William Morton Wheeler.. 325 

9. The Cold Blooded Vertebrates of Isle Royale, by Dr. A. G. Ruthven 329 

10. Annotated List of the Birds of Isle Royale, by Max Minor Peet 337 

I, Introduction 337 

II. Classified List of Birds Observed in 1905 339 

1. SiunmeT Residents 339 

2. Migrants 339 

3. Wmter Residents (migrants from the north) 340 

4. Permanent Residents j^., . . . . ,340 

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viii CONTENTS. 

P««e. 

III. Annotated List 340 

11. Noit«8 on Isle Royale Hammals and their Ecological Relations, by I>r. Chas. C. 

Ad&nu 389 

I. Introduction 389 

n. Mammal Succeasiona 390 

1. Irftke-Pond-Swamp Seriea 391 

2. The Land Series 392 

III. Faunal Affinities and MisrationB 393 

1. The Geographic Affinities of tbe Fauna 393 

2. Post-Glacial Origin of the Fauna 394 

IV. Annotated List 396 



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INTRODUCTORY NOTE. 



Throngli the genei-OMity of Mr. Bryant Walker, of DetFoit, Hon. Peter 
White* and Mr. H. M. Kaufman of Marquette, tlie Univereily Mu»eum of 
the University of Michigan wan able, in tlie summer of 1904, to »end a 
party to the Porcupine Mountains and to Isle Royale, Mivliigan. The 
aim of the party was to collect specimens for the museum and to make an 
ecological snrvej' of the regions visited. The party wan only able to 
spend a few weeks on Isle Eoyale at that time, but through the continued 
generosity of Mr. White and Mr, Walker, the survey was continued dur- 
ing the summer of 1905. The present volume on the natural liistory of 
the island has resulted from these surveys. 

To Mr. White and Mr. Walker the Museum is under special obligations 
for their hearty and Hul)stantial support, not only in the funds provided, 
but also for their aid in securing the transportation of the party. Many 
other individuals also ns-sisted in various ways. Those who aided the 
party in the matter of trauxportation wei"e: Mr. Henry Kussel, of the 
Michigan Central Railway; Mr. Geo. T. Arnold, of the Union Ticket 
Office and Dock of Mackinac Island; Mr. U, H. Brigham, of the U. 8. 
and Dominion Ti-aunportation Company ("Booth Line'') ; Mr. Henry 
Meyering, of the Graham and Morton Line; Mr, M. Adson, of the Duluth, 
South Shore and Atlantic Railway. The survey is furthermore indebted 
to Section Director C. F. Schneider of the Michigan Section of the Clim- 
atological Sen'ice of the U. S. Weather Bureau, for the loan of meteoro- 
logical instruments; to Major Lansing H. Beach, J>etroit, of the Light 
House Establishment, for permission and suggestions as to camping in 
the abandoned Ligbt-houee at Kock Harbor; to Mr. Geo. C Stone, Sec- 
retary of the Washington Club of Duluth, Minn., for the use of their 
grounds and many fnvora from their care-takers, Mr. Olias. Preulx and 
Mr. Michael Hollinger; to Mr. K. Keutson, of Park Place ("Seutson's 
Hesort"), Rock Harlmr, Isle Royale, for many favors during the stay 
upon the island; to .Mr. J, H. Malone, Keei)er of the Isle Eoyale Light, 
and to his sons, particularly to the Assistant Keeper, Mr. J. A. Maloue. 
for many favors and for their hospitality. It is a pleasure to have this 
opportunity of thanking these persons for their cooperation. 

On the return of the party from the field, work was at ouce begun upon 
the collections, and in this a large number of specialists have aided by 
the determination of the specimens. Acknowledgements are made to 
such persons throughout the report and will not be repeated here. Those 
who were not members of the party, hut who have prepared papera ai'e : 
Mr. Bryant Walker, of Detroit, Michigan, Dr. W, M. W'heeler, American 
Museum of Natural History; Mr. A. P. Morse. Research Assistant of tho 
Carnegie Institution, and Wellesley College; Dr. Jas. G. Needham, Cor- 
^Recenllf deceascrl. 



xiv INTRODUCTION. 

nell University; Prof. J. S. Hine, Ohio State University; Prof. E. S. 
Titas, Utah Agricultural Experiment Station ; Dr. A. G. Ruthven, Univer- 
sity Museum, UniverBity of Itfichtgan, and Mr. A. B. Wolcott, Field 
Museum of Katural History. 

The volunteer members of the Museum party should be mentioned in 
this connection: Dr. B, A. Brown, l>r, H. A. Gleason, Mr. W. P. Holt, 
Mr, Max Minor Peet, Mr. Otto McCreary, and the w-riter. It will be 
evident that the volunteer work of thia report comprises the major part 
of it. 

Personally the writer wishes to expi-ess his appreciation of the assist- 
ance of Mr. Walker and Mr. White; of the cooperation of the members 
of the party and the many specialists who have examined the specimens; 
and of the valuable suggestions and assistance of : Mr. Nonnan B. Conger, 
Inspector U. 6, Weather Bureau, Detroit; Dr. Glover M. Allen, Boston 
Society of Natural History; Mr. Frank Leverett and Mr. P. B. Taylor, 
of the U. 8. Geological hurvey; Prof. H. F. Wickham, State University 
of Iowa ; and to Mr. A, B. \^'olcott, of the Field Museum of Natural 
History. Also to Dr. A. C Lnne of the Afichigan Geological Survey for 
many favors and courtesies, including the preparation of the topographic 
map, and to Dr. A. O. Buthven, Chief Field Naturalist of the Survey, 
for assistance in the publication of the report. 

The shortcomings of this report will he no more evident to any one 
than to the writer. If, however, with its defects, it preserves some "van- 
ishing data," and i)reBentR suggestions for the improvement of such 
ecological survej-s, it will have served the purpose for which it was 
intended. 

CHAS. C. ADAMS. 
July 23. Ift08. 

Hull Zoological Laboratory, 
University of Chicago. 



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ERRATA ISLE ROYALE REPORT. 



Pni^e 11. lli>« <3. '»' torgtr resd large. 

l>)tge ts, line 11, for White Spruce read Blact Sjir-urr. 

I*aK« I4> llDe T, (or liit been read had been. 

rn^e lO. line 16, for Cicaila reBU TlWrrrii. 

I'ase le, line 40, tor ohM-Hoiv read aiit-llnii. 

Page l^t line SS, for hardioood rpad linnlicr.oai. 

Page 21. line 24, for HibblKcan rpBd Hipjilgcug land elgenhere In Ihe rerort). 

^BKe 21, line 25, (or ccTjicolor rend p(e*frliip(. 

Page 21. line' 48, for I.imnaea rwd Lj/iHiiofu (Had elsewliere la the report). 

I>BK« 22. line 28, for SF read iH. 

£*age 22, line 43. for Acchna read j1c»c'iho. 

I*aKe 20, line 21. (or Orupharria read G//iopflnfna. 

r>flK« 2T. line 2. (or Mlloir read blUOKii, 

page 20. line 14, (or Fly. ^ read r"'!?!. ii oriJ (U. 

Paiee -40. Ilo^ 21, for FAe bcarfnif of Ihi latter read tfiirlr. 

rsfCR •IT, line 46, for c read J. 

I-aK» -48. line 14, (or Fig, S3 read Fly. Sj, 



rage i 



f 33, for r read J. 



[•age 01> llitc 28. dele (Fig. SSI. 

Page G3, line 10, dele Ftff. X: 

Page 03, line IS, for Lalie read Lakfi. 

I'age «-4. line a, (or Fig. U read Fig. }S. 

Page 41^, llUM 48 and 40, for Formica ailamall read F 

I'OKB GS. lines 50 and 01, dele !fo. lU. 

j-Bge ''''t '!"* ■^"' '"'■ O'Tp'iocna read agrojiharna. 

Page I>8. line 23, tor XI read F/, 

Page HO, line C, (or coniKHoH of ircafAT rend ooiiilff 

I'age 13S, line 31, (or fotterfd read Jorcultd. 

I>aee l!t2, line 20. foe BuriM, F. «. read Uurag. F. I.. 

page 158, line 43, tranapoHp B»prf«((rf« and Trichlai. 

page ISO. line -9, for Op-opftofmn rend Ourophariia. 

Page 161. line 44, for Tftefr ' " ' 



I'age 



18B, line 2S, dele "William. 



I>age 200, line 46, for Balitobma read JloUtoblut. 

j>a^e 20B, line 46, for A'cy Jeraeu read -VriP Jcrie/i. 

PblS^ 257, line 17. re&d SaH(ci'Ia'-=l((Wor. 

page 201. line 43, (or /aasfilaeae read Jni*f<fnc. 

I>il^e 284. line 28. for St-6i read f7r)». K(-(W. 

page 30rt. line 28, dele 1. 

page 306, Hue 26, add ». 

j>age 3O0. tine 30. add 8. t'oao"*". 

page 342, line 25. (or Fig. *l rend fig. JT, 

paee 3S0. line .15, add Pig. 60. 

Page 3G4, line IT, add Flo. n ; line 20, dele Fig. n. 

Page 8©3. line 13, for (ojlMfHce* read liifccfiicrii. 

pg^^ S07. line T, for Canton read CotON. 

I'age 40T, Hoe 26, (or /, .V. itaionc read J. ft. Jfnioic 



I'age 



419. line 14, (or Hoopt read Baofim. 



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ISLE KOYALK AS A BIOTIC KSVIROSMEKT. 

BY im. CHARLES I'. ADAMS. 
I. INTHODtCTION. 

i. Itinerary and Feraomtel of the Parly. Tlie Univereity Mueeuin 
party left Ann Arbor, Michigan, Junf> lili, and reiifhed the abandoDed 
light-house at Rock Harbor, Isle Koyalo, on tlie morainfc of July 5, 1905. 
Th^ jMirty was coinpoeed of the foltowing: N. A. Wood, Dr. R. A. 
Brown. Dr. 11, A. (iteaaon, W. P. Holf, Otto Mcf.'reary, a camp hand, 
B. F. Silvery, and the writer. In fieueral, the duties of the various 
!Deinber» were as follows: Mr. Wood, llie Museum taxidermiHt. 
looked after the trapping of mammaln and the inTparation of bird and 
mammal skins. He wns assisted by Dr. Brown, who gave moat of his at- 
tention to the study of the bii-d life, and who remained with the party 
until July 25. In the study of bird life. Dr. Brown, Mr. Wood and Mr, 
Mi'Creary co-operated, the latter devoting his entire time to the ecolog- 
ical phase of the work. l>r. (Ileason devoted his attention to the collec- 
tion and ecological study of invertebnites. particularly molluscs and in- 
sects, and most of the photographs were taken by him. In eollecting in- 
sects about the camps, he was assisted by B. F, Savery. Mr. Holt's time 
was devoted to the study of the vegetation. The writer, who was in 
charge of the expedition, gave special attention to the environmental 
dynamics, biotic succession, and the genera! correlation of the work 
of the various members of the party. 

Dnring the stay at Kock Harbor, Fig. J, the following localities were 
examined : The shore, from the light-house south to the bead of Conglom- 
erate Bay; the region abcwit the head of Rock Harbor and Sumner 
lake: a line from the month of Benson brook to Sargent I^ake and Me- 
Carjjoe Cove.; and the vicinity of Tobin Harbor; in other words, the 
localities included in Stations I-IV, 

The party remained at Rock Hartor from July ~i to August 1, an^I 
then moved to Kiskowit Bay. Here Mr, Max M. Peet joined the party 
on August S, and devoted his attention to collecting birds ond mammals. 
He also took a numlter of the photographs. While at the Siekowit 
Camp, the bay and lake of that name were examined, and also the Hay- 
town trail and the islands near the Isle Koyale Light. All of these lo- 
calities are' included in Station V. 

On August IT the party moved to Wiishington Harbor, and was then 
partially disbanded. The members who remained made their headquar- 
ters on the grounds of the Washington Club, at the head of Washington 
Harltor. After September 3 Mr. I'eet alone remained there until the 
22nd, in order that he might continue the study of the fall migration 

..Google 



2 MICHIQAN SURVBT, 1908. 

of the birds. He returned on the last boat of tlie Reaaon for Dnliitb, 
Minn. 

Dnring the previous (190i) season, the Mnseum prfrty had made a 
general examination of the vicinity about Washington Harbor, so that it 
was now thought desirable to devote more time to other localities. With 
the exception of bird migration, no detailed work was done in this vicin- 
ity in 1905. In addition to the region about the head of the harbor, l^ke 
Desor was also reached from this point by means of the road along the 
crest of the (ireenstone Itange. 

2. The Aim and Metltodfi of Work. The field work whs couductcd oii 
the same general plan as that pursued during the pievioue season in 
the Porcupine Mounitainn and at Washingjfxin Hapboi'. Much inor** 
ground was covered, however, l>ecause it seemed imprnhable that a third 
trip could be made to the same region. It therefore seemed desirable 
to gain some idea of the biota of the island as a whole, because of its 
Canadian character. Even then, the survey was confined almost exclu- 
sively to the region south of the (ireenstono Range. 

As mentioned in the report for 1904 (Kuthven. '06, pp. 11-12). the aim 
of the work was not simply to collect 8|)ccimens, but also to study the 
relations of the plant and animal life (the biota) to their surround- 
ings. The environment as well as the biota was considered from a dyn- . 
amic standpoint, and an ctfort was made to analyse the environment 
in order that the dominant conditions and proceiwes of which it is com- 
posed might be recogni7.ed, and their laws of change be perceived and 
formulated. To i-esolve au<'h a pi-obleni as this must of necessity 
require more time and detaite<l investigation than the possibilities of a 
few months work will pei-mit, and yet it is equally evident that prelimi- 
nary work should be carried on from a genetic stand|>oint. becanst^ such 
a method determines upon what facts cni])haKis should l»e i>laced. and the 
broader and moi-e general relations, as well as the details, are equally 
subject to a genetic and dynamic treatment. In jireliniinary work of 
this character, it is considen-d of special importance to discover, if pos- 
fiible, the order of the major biotic siiccessions, becanse these succes- 
sions must be clearly i)er<'cived before their causes can undergo ade- 
quate analysis. Our knowledge of causes generally laps far behind our 
recognition of successions. 

Thus throughout the study of the Isle Royale biota a special effort 
was made to investigate the genesis or successions of events. The en- 
vironment has not lieeu considered as limited to habitats alone, bnt 
also to include that greater unit, the geographic. To ignore this is 
to overlook the real background. It is believed that certain advantages 
ai*e derived from this method of work, which, although they may be 
recognized from other points of view, are likely to be subordinated to 
other facts. It should not for a moment Ik? thonght that this method is 
considered as the only one of approach, but it apjiears to have certain 
advantages which seem to justify its adoption. Nor should it be inferred 
that the genesis of the biota and the habitat is all that should he in- 
cluded in an ecological survey. The problem of suci-essinn is only one of 
several which clearly show the intimate i*elations and i-esponses between 
organisms and their envtmnmenf ; others that remain to be investigated 
involve physiological and structural change»<. and various modifications 
of habit and behavior of Ixith plants and animals, 



ECOLOGY OF ISLB ROYALE. 3 

The ecolofcical relationa in the north are so different, in some i-cajiocta. 
from th«ffie farther south, that one inay easily form au erroneous *;ou- 
(■eption of the conditions under whicli nucli a pieliniinapj- investigation 
may be made. A very favorable condition for the work wjiu the fact thiit 
practically all the time was devoted to it, instead of only ocniHJonal 
trips being made for the purpose, as is necessary with those busy with 
other dutieH. There are also certain advantages in being able to be io 
(Im? field coutinuouely, an a certain familiarity with conditions iw ac- 
tjuired in the beginning, especially whei-e the variety of forms is limited, 
nhirh otherwise would involve, time upon each visit. Although 
tuoat of the ntenibers of our party were upon the island only during July 
nnd Angust, yet at this time those seasonal plieuomena were concen- 
trated which require much more, or several times that amount of time 
for their development farther south. The seasonal contrast is well il 
Inatrated when the siminier season at the other extreme of the State of 
Michigan — 5(10 miles away— is contrasted with that of Isle Itoyale. 8uch 
relations are further reinforced by the fact that the species and 
societies which are dominant in the various northern habitats are 
ycTv much smaller in nunil»ei- than farther south. This necessarily sim- 
plifies the problem, and to a corresponding degree i-educes the cham-es 
of error in anticipating biotic changes which are correlated with those 
"f the environment. This is a i-elation of much imiH>rtance in liie study 
t'f HMcicssida. The writer was especially impressed with the rrUitirr 
xiMpUrilu of the problem of eiivironnieutnl relations and of the biotic 
Bncressiim «|>on Isle Royale, and has received further coufiniiation of the 
"pinion that a tendency to exaggerate the complexity of the cnviifm- 
meat is prevalent. 

An iiii|M>rtant aid .in envii-onjnental analysis has l>een reieived from 
llie elTort to distinguish the major or geograpliic features of the gross 
eiiviroii.nient from the minor habitat units which make up the mosaic 
or complex, although their mutual and genetic n-lations weiv not 
"vcrlooked. Some of these relations have l)een well expressinl by 
llontgniuery in his comment on distribntion t'll(>, p. G) as follows: 
"And, as is always the case when the method has been consistent and 
wientific. the factors of distribution and the meaning of it will ulti- 
mately be stated in very simple form. These factoi-H appear to ns now 
to be enornionsly complex, but this is Wcause we hai-e hardl.i" com- 
menced to analyze them."' 

At this point it should lie mentioned that there are certain dif- 
ficulties which tend to confnse tlie field worker, which, if clearly 
understood, will often be of aid in ecological studies. In pursuing 
fleld studies, in addition to a knowledge of the species, one of the fii-st 
essentials is a familiarity with the habits and habitat preferences of 
organismB; and further, there should be the ability Io i-ecoguize bow 
the dominance of one society is transformed into that of another. 
The lack of a sufficient power of constructive imagiuation makes the 
detection of such traosformations very difflruH. ]>erhapa even impossible 
to some. This limitation almost completely i-estricts such a student 
to the purely descriptive phases of fleld ecologj', because the explanatory 
phase lies beyond his gi-asp, although there remains for him a hirgn 



4 MICHIGAN SURTET, 1908. 

field (or UBeful and valuable activity in the stndy of habitat preference, 
and the mntnal relations of the aBBOciated species in given habitats. A 
familiarity with the forms studied, under diverse circumstances, de- 
velops a certain perspective which is a great help in preventing confusion 
caused by minor and relatively insignificant details. 

The limited time spent in the present investigation did not permit 
detailed studies of the interrelations of the oi^aoisms within the habitat, 
either in their relation to the environment or to each other. In local 
studies attention is usually given to detailed life histories rather than 
to a delibCTate study of their interrelations as members of a society. 
The emphasis which is sometimes placed upon individnal life histories 
would lead one to expect that such histories could be assembled and 
would give us the same result as if they had been studied as a aocietf/. 
But the points of view are so different that such a result, although 
theoretically possible, is unlikely to be obtained. At this time we only 
wish to emjdiasize the fact that both methods should be used to secnre 
the best results. For example, in applying these principles to the study 
of birds, the life histories of the dominant species of a society might 
first be given special attention. Then the relations of the dominant 
species to others of the association and to the environraeut may be deter- 
mined and later on the subordinate kiuds considered. This will involve 
prolonged study in the field (and laboratory) of the habits of nesting- 
feeding, rearing of young, etc., as influenced not only by other mem- 
bers of the same species, but also by other species in the same habitat. 
The same general method is applicable to other groups of organisms. 

3. Previous Biological Inveatigatians upon Isle Royale. Previous to 
the investigations by the Museum party in 1904, {Huthven and others. 
'06) there seems to have been very little study of the Isle Royale biota. 
Several collections of plants and animals have been made, but very little 
has been published about them. In 1848 W. D. Whitney was "ornithologist 
and botanist" for the government geological survey parties, and he pub- 
lished a list of the plants found. (Foster and Whitney, '51, pp. 359-381). 
Incidental mention is also made in these geological reports of the collec- 
tions of animals (Foster and Whitney, '50, pp. 17, 51, 201; Jackson, '49. 
pp. 423, 440, 441,) ; but, so far as known to the writer, no detailed reports 
were published. 

So far as the vegetation is concerned, the most important source of 
information is the maps of the Ives Linear Survey. Here the general 
character of the forest, the extent of the swamps, and the underbrush 
are indicated. Mr, Henry Oilman ('73), of Detroit, made two visits 
to the island (one of which was in 1873), and his botanical and ethno- 
logical collections were presented to Columbia and Harvard Colleges- 
In 1890 Mr. F. E. Wood made a collection of plants from the vicinity 
of Rock Harbor and presented them to the herbarium of the Botanical 
Department of the University of Michigan; and in 1901 W. A. Wheeler 
('01) published a short pajwr on some plants taken on the northeast 
end of the island, ' 

The invertebrate fauna found in the deep water off Isle Royale was 
examined by Smith in 1871, and a list of Coleoptera from Isle Hoyale 
was published by Hubbard and Schwarz ('78). The writer has recent!\' 
published a paper on certain phases of the problem of succession, ns 



ECOLOaV OF ISLB ROYALE. 5 

illustrafed by the birds upon Isle Boyale. This paper, with Bome ad- 
ditions, is included in this volume. Detailed references to these papers 
will be found in the accompanying bibliographies. 

From the above remarks, is it quite evident that very little attention 
bas been given to the biolog:ical conditions of the^ island, and moch 
remains to be done. In all probability other naturalists have visited 
the Isle, but I have not learned of their results. 

4. Historical Nate. The history of Isle Royale, since its cession 
by the Chippewa Indians in 1843, is, in brief, one of prospecting, 
mining explorations, flying, summ^ resorts, and scit*ntiflc surveys of 
the topography, hydrology, geolog}-, and biology. 

A general historical account is given in Lane's report ('98) on the geol- 
ogy of the island, and need be mentioned here only in outline. There 
is abundant evidence that in prehistoric times the Indians mined copper 
«n the island. Within three or four years after the cession of the island 
to the United States, it was invaded by prospectors and explorers, so 
that by 1847, according to Lane, "the island presented perhaps as lively 
a scene as ever in its history." At this time the Linear Survey was 
made by William Ives. But this period of activity was only of short 
doration, for the decline was almost as rapid as had been the ascent , and 
by 18S5 the "island was a desert once more, with no permanent in- 
habitants." (Lane). This passive condition of affairs lasted until the 
Lake Survey engineers arrived in 1867. This survey continued until 
1871, during which interval explorations were somewhat revived, and 
continued for several years, only to be followed by another relapse and 
still another ascent in 1891, when a number of careful and detailed 
explorations were made for copper by means of the diamond drill. 
But this activity also ceased about 1892. A year later, and again in 
1895, Dr. Lane visited the island for geological investigations. 

The mineral sources are thus seen to have been the main attraction. 
The forest growth is too stunted and inaccessible to have merited the 
attenti<m of lumbermen, although several timber prospectors were pre- 
sent during the summer of 1905. During more recent years the Ash- 
ing and summer resort business have attracted some attention to the 
island, and have made it accessible during the summer through regular 
steamboat service. The climate, scenery and the fishing make the 
island very attractive as a summer resort, but it should be recognized 
that if the scenery is to be preswved the forests must be protected 
from fires, because reforestation is exceedingly slow on land with such 
a shallow soil. It is to be hoped that the geographic isolation may be a 
protection from such devastation, because the cool summer climate, 
the rocky coast, the forests, the picturesque scenery, and the 
surrounding Lake Superior, are natural features which should long 
r^natD attractive to summer visitors. If the interest in copper should 
revive permanently, the biota will become gr^tly modified, in which 
cajse some conception of the conditions upon the island in 1904 and 
1905 will be preserved by these records. 

[It may be of interest to note here that 86,000 acres of the island 
were for sale in the winter of 1908 for $150,000. Lane.] 

5. Available Maps of Tale Royale. The available maps are not 
generally known to the public, and are therefore listed here, especially 
those which are of value from a biological standpoint. >oIc 



6 MICHIGAN SURVEY, 1908. 

1. The Ives Linear Survey Maps. Because of their large size (2 
inches to the milej, and the detailH concerning the character of the 
swamps, the forest and the soil, this is the most useful map for tbe 
field. Photolithogi-aphii- copies of the township maps, of which there 
are eighteen, inav be secured for 25 cents each from the General Land 
Office at Washington, ]>. €. 

2. The U. 8. I.rfike Survey Chart of Isle Royale, (Catalog No. Sh.). 
This is very uBeful l>ecau8e it indicates the topop^phy, in part liy 
hachures, and gives the details of the coast, including soundings and 
the character of the bottom. A large tract of the interior, between 
lakes Desor and Chickenbone, is unmapped. This nuip may be secure<l 
for 25 cents from the Lake Hur\ey Oflices at Detroit and Duluth. An 
excellent cliart of the cntiif- Lake Superior basin may be secured from 
the same source. 

3. Lane's Geologicjil Map. Published hy the Michigan State Geo- 
logical Survey. It accompanies Lane's report ('98) on the geology of 
the island, and is on a scale of ■% of an inch to the mile. 

4. Passage Island Topogi-aphic Hh^t. This is the only sheet pub- 
lished by the V. S. Geological Survey which includes any part of Isle 
Royale, and it covers only the extreme northeastern end of the island. 
This may be secured from the Survey for 5 cents. The contour inter- 
val is 2» feet, and the scale one inch to the mile. 

5. An English land company is said to own much of the island, and 
has published a map on a scale of % of an inch to the mile. 'Oie agent 
for this company is R. R. Goodell, Houghton, Michigan. 

H. TUG BIOTA CONSIDEBBD BY STATIONS. 

1. The Location of Field Stations in, 1905. As a detailed survey of 
the entire island was impossible, it was necessary to select representa- 
tive localities and conditions, or habitats, and to devote to these all 
available time for study and collecting. In order to make sure that 
these conditions were re])reseutative, considerable care was necessary 
in locating these i^tations. In general a Station, in the strict sense, 
stands for a region, while a Substation refers to a particular habitat, 
usually of relatively limited extent. The character and extent of a 
Substation, (or, as it is generally called, for the sake of brevity, a 
"station,''! was determined primarily by the relatively homogeneous 
character of the conditions. Thus a "station," as the Balsam-Spruce 
forest (V, 4) for example, varied somewhat in its extent with different 
groups of organisms. In the case of birds it included a greater area 
than was necessary for many invertebrates, such as land snails, but 
in every case such a "station" is intended to enable one to determine 
wiiat organisms were dominant and chaiacteiiKtic of such a sample 
sitnation. 

Some such system of sainpliug is generally advantageous or necessary, 
and ihis is particularly essential in the case of a surveying party, in 
order to give deflniteness and co-ordinated activity to their work, parti- 
cularly if the results are to be made at all comparable. Of course some 
individual judgment is necessary in applying such a plan to different 
groups, but no more perhaps than is necessary to carry out any other 
comprehensive plan. ^- i 

^ ^ i,Cooglc 



ECOLOGY OF ISLE ROYALE. 



J. Lwntion of Field Statiattfi, I'.m. 



s^tation I. Light-houKe Peninsiila. l>etween Kwk Harbor and tlie 
iM^aa of Conglomerate Bav, Sec. ;!G and X. K. '/i Sw. 3i. T. 66 N., R. a^t 
W\ 

Snb. 1. Lake and Bay Beaches. 

Sob. 'a. Natural Rock'CIearinps, K. E. 14 Rec. 26. 

Sub. 3. Balsani-Spruce Forest, N. E. V4 Sec. 26. 

Snb. 4. Tamarack, and Arbor Vitae Swamps, See. 26. 

Snb. 5. Jack Pine Ridge. S. W. 1/4 Spc. 26 and S. E. V^ Sec. "- 

Sub. 6. Sphagnum-Spruce Bog. H. W. 14 Sec. 26 and 8. E, 

Vi Sec. 27. 
Sob. 7. Light-house Clearing, N. W. % Sec. 26. 
Stution II. Rock Harbor and McOargoe Cove Trail, Sec. 27, 22, 21, 
20, 29, 30, T. 66 X., R. 34 W.. and Sec. 25 and 26, B. 35 W., T. 66 N. 

Sab. 1. Benson Brook and Ransom Clearing (outlet of Benson 
Lake), >'. E. 14 Sec. 27 and S. E. y^ Sec. 22, T. 
66 N.. K. U \V. 
Sub. 2. Tamarack Swamp, S. W. 14 Sec. 22 and 8. E. ^ Sec. 

21. T. 66 X., R. 34 W. 
Sub. 3. Rock Ridge Clearings (burned ovei). Sec. 21 and 20, 

T. 66 >'., R. 34 W. 
Sub. 4. McChi^oc Cove, at end of Trail. X. E. y^ Sec. 2(1 

T. 66 N.. R. 33 W. 
Snb. 5. Forbes Lake, N. E. 14 Sec. 28, T. 66 N., R. 34 W. 
Station III. Western End of Bock Harbor. Sec. 28, 33 and 32, T 
66 N.. R. 34 W., and Sec. 5 and 4, T. 65 N., R. 34 W. 
Sub. 1. Small Island, S. E. y^ Sec. 32. 
Sub. 2, In Harbor at West end of Island, Sub. 1. 
Snb. 3. Bulrush Zone and Delta, Sec. 32, T. 66 N., R. 34 W. 
Sub. 4. Trail to Sunlner Lake, Sec. 33, T. 66 N., R. 34 W. 
Sub. 5. Sumner Lake, Sec. 33 and 34, T. 66 N., fl. 34 W. 
Sub. 6. Southwest Coves of Rock Harbor, Sec. 5 and 4, T. 

65 N., R. 34 W. 

Station IV. Tobin Harbor and Vicinity, T. 66 and 6T N., R. 33 W. 
Sub. 1. Scovill Point, Sec. 26 and 35, T. 67 N., R. 33 W. 
Sub. 2. Island Xo. 14, Sec. 26, T. 67 X., R. Sti W. 
Sub. 3. BaTOu, Xorth of Monnment Rock Trail, X. W. 14 l^ec. 

34, T. 67 X., R. 33 yV. 
Bnb. 4. Trail to Monument Bock, X. W. Vi Sec. 34, T. 67 X^., 

H. 33 W. 
Sub. 5. Clearing at Xeutson's Resort (Park Place), Sec. 4, T. 

66 N., R. 33 W. 

Sub. 6. Small island in Tobin Harbor. Sec. 5, T. 66 X.. R. 3;t \V. 
Sub. 7. Head of Tobin Harbor, Sec. 7, T. 66 N., R. 33 W. 
Sub. 8. Trail to Greenstone Range, Sec. 7, T. 66 X., R. 33 W., 

and Sec. 12, T. 66 N., R. 34 W. 
Sub. 9. Mountain Top, Sec. 12, T. 66 X., R. 34 W. 
Station V. Siskowit Bay, Lake and Vicinity. 

Sab. 1. The Beach, (at camp). Sec. 32. T. 65 X.. R. 35 W. 
Sub. 2. Heath Zone and Beach, Sec. 33, T. 65 X., B. 35 W. 
Sub. 3. Bock Clearing (at camp). Sec. 32, T. 65 X., R. 35 W. 



8 MICHIGAN eURVBY, 1908. 

Sub. 4. Trail throogh Baisam-Birch Foreet, Sec. 32 and 31, T. 

05 >'., R. 35 AV. 
Sab. 5. Tamarack Swamp. N. W. % Sec. 32, T. 65 N., R. 35 W- 
Snb. 6. South Shore of Siskowit Lake. Sec. 31 and 32, T. 65 

>'., R. 35 W. 
Snb. 7. Havtown Trail, from Siakowit Ijabe, West Line of Sec. 
24, aoi-oss Sec. 13, T. G5 N., R. 3(1 ^V., cf. Lane. '!I8. 
pi. XI. 
Sub. S. .\rlM)r Vitae Swamp, at end of Havtown Trail, X. W. 

i,i Sec. 13, T. 65 N., R. 36 W. 
Sub. 9. Ontlet of Siskowit Lake, N. W. i/l Sec. 36, T. 65 X., 

R. 36 W.. and Kec. 31, T. 65 X., R. 35 W. 
Sub. 10. Long Inland Gull Rookerv and Menagerie Island, T. 

64 X.. K. .35 W. 
Sub. 11. Taiuaiack Spruce Swamp, See. 33. T. 65 N., R. 35 W. 
The following stations were examined by tbe Museum partr during 
the season of 1904. Part of these Stations were re-examined and will 
be referred to by Station number and date, thus: Sta. I. '04. 

Station I, 'M. Clearing on the Shore of Washingfoii Harbor, Sec. 2u, 
T. 64 X., R. 38 W. 

Station II, '04. Washington Treek. Sec. 29, T. 64 X., R. 38 W. 
Station III, '04. Trail along the top of (ireenstoue Range (Desor 
Trail). T. 64 X., R. 37. 38 W. 

Station IV, '04. Washington Brook. Sees. 2S and .32, T. 64 X., R. 3S W. 
Station Y. '04. Tamarack Swamp, Sec. 20. T. 64 X.. R. 38 W. 
Station VI. '04. South of Greenstone Range, Sec. 32. T. 64 X.. R. 38 \V. 
Station TII, '04. Lake Desor, T. 64 X., R. 32 W. 

Station VIII. '04. Western end of Siskowit Bav. Wees. 27 and 2S, 
T. 64 X., E. 37 W. 

Station IX, '04, SonthwpHtern end of Minong Trap Range, Sec. 30. 
T. 64 X., R. 39 W. 

Station X. '04. Washington Harbor. T. 64 X., R. .38 W. 
2. General Charactfrhtu-s of the Statiottf*. In this swtlon, I do not 
aim to give a completely correlated account of the biota of each stn- 
tion, but to present a general idea of the main characteristics of the 
various situations examined, and some of their common and represeii- 
tati^'e plants and animals. Photographs illustrating the chamc- 
teriaticB of the various ''stations" will accompany thJK section, and 
should be consulted in connection with the text. 

Station I, Substatifni 1. The Lake and Bay Biachix. This "station" 
includes the shore line from Rock Harbor, near the light-bouse, Fig. 1, 
to the head of Conglomerate Bay. The entii-e shore was not studied in 
detail, as most of the time nas devoted to the lieachcs which ai-e l>eing 
formed at the heads of the coves and bays. Quite a variety of condi- 
tions are repre8ent<Hl along this shoi-e. due not only to the degree of 
exposure to the waves of Lake Superior, but also to the character of 
the rocky coast itself. All degrees of shore and beach are developed, from 
overhanging and vertical '■liffs, Fig. 2, with bases strewn with large 
blocks lowered by sappiug, to a shore line with a low angle strewn with 
shingle and gravel, and a sandy beach, as found at the head of Conglom- 
erate Bay. In harmony with the dip of the i-ocks and the effect of the 

D, _, i..C0CH^Ic 



ECOLOGY OP ISLE KOYALE. » 

glacial ice movement upon tbe valley alapes, which tend to be gentle 
on the soQtheaetem aide, the corresponding shores of the bays and 
coves are nsually at a low angle, except possibly where faulting has 
taken place, or a wave cut terrace has been developed. The northern 
Bides of tlie bays are comparatively abrupt, and there is thus a tendency 
for the cliffs to occur mainly upon tbe northern slopes and shores. The 
larger bays are the submerged portions of the valleys, marie the 
location of the less resistant rocks, and are inherited topographic fea- 
tures; but many of tbe minor cnvcs and the rocky headlands have been 
carved by the activity of the present lake. The beaches are only de- 
veloped at the beads of the coves and bays, and are very largely com- 
posed of shingle and gravel. The only extensive sand beach seen was 
at the head of Conglouievate Bay. The character of the material com- 
posing these beaches clearly shows its local origin, and emphasizes the 
isolation which prevents long shore transportation of sncli material. Thus 
only floating material is liable to extensive long shore dispersal, a signifi- 
cant fact that beOTS npoo the dispersal of the snail life along the shore. 

During severe storms, the wave action upon this coast is quite in- 
tense and even the waves of the summer storms are quite active, aa may 
be »een by referring to Fig. 3. The blue deep lake water comes close 
np to the shore, so that generally no breaker line is developed off shore. 
In several places there arc numerous reefs or islands (usually the iso- 
lated continuations of the rock ridges), which tend to break the force 
of the waves rolling in from the opeu lake. 

>'o effort was made to study the life of the open lake, only the shallow 
irater of tbe bays and coves being examined. The major environmental 
features of tbe coast are tbe Ijower, Middle and Upper Beaches; but these 
are only differentiated clearly at the heads of the coves and harbors. 
The Ijower and Middle Beaches are only seasonal expressions of the same 
phenomena, but ecologically they are fairly distinct. 

Tlic Ijoicer Bench. This beach extends from the shallow water to 
the upper limit of the summer waves. The submerged portion is not 
sharply defined above on account of tbe changes in level of the water 
surface, dne to waves, the pariodical and seasonal fluctuations, and the 
atmospheric pressure fseichesl. In time there has been a downward 
migration of the entire beach zone, a tendency which is in part counter- 
acted by the northward elevation of the land. This is the zone domi- 
nated by water, ice, and wave action. It is certainly a sharpiy defined 
tension line upon an exposed coast, which clearly suggests that it is not 
probable that many forms of animals have made tbe transition from 
fresh water to the land under such conditions. If we consider the 
shore habitats as including all stages from a rock cliff to the sand beach, 
the lower beach and the protected shores are the most favorable aquatic 
habitats upon sncb shores. 

Upon the sloping rock, shingle, gravel and sand beaches is found a 
varied fauna. lu winter, when the bays are frozen over, a calm is pro- 
duced which must be favorable to the pi-eservation of the aquatic life 
upon this stormy coast. 

The general character of the sandy beach at tbe head of Conglomerate 
Bay is shown in Ft;/. .{. The life of the submerged portion of the shore 
ie quite limited, except on the beaches and protected portions. The vege- 



10 MICHIGAN SURVEY, I90S. 

tation consists of al^e, which grown in moderate abim<^aDce. though 
not luxuriantly, as found about the Gull Rookery (V, 10), or at the 
fishermen's cainp at Rock Harbor, a fact which suggests that the abun- 
daofl^ of suitable nitrogenous material is mucli greater in such places 
tbau in the open lake water. With the development of the fall storms. 
Mr. J. A. Maloue states that the** rocks (V. 10) are washed free of 
the algae, thus evidently necesuitating a repopulation of these surfaces 
each season. 

The characteristic fauna secured in the shallow water shore margins 
were the snails, Litnnaca stagnnHs. L. cmarijinata. and Pht/sa sai/ii. A 
small fish, the Miller's ^'humh, VranitU'a frnnklini. is also fairly abun- 
dant and characteristic of this shore. 

Upon low rocky shores beach pools, Fig. 5, are occasionally found 
which, when favorably located, art supplied with water by the ordinary 
summer waives, otherwise by storm waves and rains. The precarious 
existence of life in such places is indicated by the general type 
of the fanna, which shows exceptional power of locomotion, usually 
coupled with a short life cycle. The immature stages of insects 
are rather characteristic, as shown by nymphs of the water boat- 
men, Ctyrixa, dragonflies and Caddis fly larvae. Water beetles were 
represented by Rhantus J)inotatu8, and the snails by Limnaea emarginata 
and Planorbia "parvus. The Gulls and Spotted Sandpipers should be 
mentioned as birds which frequent these conditions. 

The Middle Beach.' This beach occupies the strip of shore over wliicii 
the winter waves retreat as they fall to the upper summer storm limit. It 
is thus seen that the Middle Beach is only a temporary or summer aban- 
donment of part of the upper shore, which is repeatedly claimed by the 
winter wares. In summer this strip is exposed to denudation; in the 
fali and early winter, to the fury of the waves, and, later, it is coverefl 
with ice. Driftwood and debris tend to lodge here and to accumulate. 
It is an important region of biotic invasion for laud forms. Beach 
pools are also developed in this area, upon the abandoned wave cut ter- 
races of earlier lake levels. Upon the cliff faces, sloping rock shores and 
shingle beaches, little is found that is favorable to life, but upon the 
protected sand of the Middle Beach, relatively favorable conditions for 
many organisms are found during its period of exposure. The character 
of the substratum of the Middle Beach varies from rock to shingle, 
gravel and sand. 

The characteristic features of the vegetation, where the wave action 
is not too severe, are the fruits which are washed ashore by the waves, 
together with certain annuals and lichens. The fauna varies with the 
chara<-ter of the conditions. The open character of this beach and the 
relative abundance of animal food makes such situations favorable for 
spiders of the genus Pardosa. The same open character makes the 
shores a favorable patrol for certain butterflies, particularly Bagilai- 
chia aHhcmis. Insects and snails washed ashore by the waves also 
characterize this habitat. 

The Upper Beach. This part of the beach is beyond the reach of the 
waves, and forms the transition between the open beach area and the 
Inland forests. The width of this belt varies greatly with the gradient of 
the shore. Where the beach is continuous with a more or less bare rock 



ECOLOGY OF ISLE ROYALE. 11 

ridge, tbis hnbitnt may be rattier extpusire and ill defined, as at the ridge 
south of the lighthouse (I. 2(. but when it bordei-s a depresfiion. as iit 
tlie head of the rockboiind coveH, or where a beaeh is well de^'eloped, 
this transitional zone is more clearly defined and limited. When thia 
beach is wide and grades into the rovk o|>eningit, rr in F'lfiurrs fl and 7. 
the cruataceous and fqliaceous licheim grow niton the rorks; but if 
sM>il accnmulates, as is shown in Fifj. 6, the Cladonin — Jlearlierry society 
becomes established, and includes some annuals, such as SoHdario. A 
limited variety of insects, especially ants, characterize such conditions. 
When adjacent to the forests, in depressions, this tteacli is generaliy bor- 
dered by alders, soine aspens and young trees. 

The fauna consists largely of insecls, such as bntlerflies, certain dra- 
gonflies and Hymenoptc-ra, which frequent the open places on wing. 

Station I, Substation ~. yatural Rock Clearings. This Station con- 
sists of two sniall rock opeuings. one just north of the light-house, and 
the other south of it, on the north side of the entrance to Tonkin Buy, 
only a short distance from the light-honse. They were both park-like ave- 
nues extending along the ridges, largely bordered by the Balsam-Spruce 
forest. 

The north ridge will first Ite considered. The general character of the 
opening is well shown in Fiff. S. The White (Spruce, Balsam, Paper 
Birch and Arbor Vitae bound the rid^e on either side, within which 
there ie a distinct heath zone of Bearberry and patches of Clmlonia, while 
along the central aisle there is a shallow residual and humic soil on the 
almost bare rock. The south slope is rather gradual, but the north 
slope and the end of the ridge at the shore form a ciiflf. 

The fauna of this location was limited, Snails were found among 
the CUidonia, such as ^'crtigo, Zonitoiilrs arborcu and Pyramidvla 
cnmkhcitei anthoiii/i. This was also a rnnway for Hares, 

The south opening or clearing is situated on a low sandstone ridge 
which slopes down to the heach. and is thus in marked contrast to the 
north clearing, which ended in a cliff. This gradual slope beautifully 
illustrates the transition from the bare rock beach, through the moss 
and lichen zone, to tlie Cladonia. Itearberry and Solidago fiora, (Fiffn. 
6 and 7), and on to the crest of the ridge. Fig. 9, with its dominance 
of Cladonia and Bearl>erry. The severity of the conditions is furthered 
b.y the weathering of the sandstone into thin scale like layers, about '/i 
of an inch thick, which l)ecome loosened and slide down the slo|)e. Thus 
a vegetation may become fixed to the rock surface, but not permanently 
to the slope. These scale like fi-agments are shown in Fig. G. That a 
greater amount of vegetation would grow here, if the soil were allowed 
to accumulate, is shown in Fig. (1, where such conditions have been 
produced by the presence of a larger boulder. The (7?fldo»i.ia -Bearberry 
aVenue extends along the crest of the ridge, Fig. 9. This is bounded 
by large Jack Fines near the beach, and farther from the shore by the 
^Isam-Bircb forest. 

The zonal distribution on the ridges is quite marked; the central strip 
is composed of Cladonia, Bearberry, Solidago, and lAnnea borealis: 
while this is bordered by a shrub zone composed of Junipcrtis nana. 
alder, Arbor Vitae and young Balsams, and a bordering ti'ee zone is com- 
posed primarily of Balsam. When once the shade of the forest, es- 



12 MICHIGAN SURVEY. 1908. 

specially that of the Balsams, encroaches upon the Cladonia society, 
the Bearberry first becomes reduced in number, and is then replaced 
by Aster macrophyllus, and a moss from the forest floor. The former 
is perhaps the most striking and characteristic shade plant upon Isle 
Boyale. The succession, or order of invasion on the ridge, from the Cla- 
donia to the Juniper and into the Balsam forest, is thus briefly shown 
in the transverse section from the central ridge to its margin. This zonal 
phenomenon, as will be seen later, is only an expression of the relative 
rates of invasion, and is not a phenomenon separate from the normal 
sacceaaion. 

The soil upon the top of the ridge is about two inches deep. It is 
residual, supplemented by the faiunus from a now extinct emstaceous 
lichen society (that of the dadonfa-Bearberry), and at its margins 
by the Juniper, Balsam, Birch and Ja^k Pine leaves and debris and 
further, to an important degree, by the excrement of the numerous 
Varying Hares which frequent the rock ridges. 

In the case of rock ridges which entend down to the beach and are 
thus in direct communication with the shore drift, conditions exist 
which show how such ridj^es may have been invaded by lichens from 
two sources — the shore drift and the ex^wsed beach itself — because of 
the continuity of the rock habitat. Of course possibly another origin 
is to be found in the fact that this ridge was itself once a beach. Ants, 
^^si^oppers and a few other insects characterize this fauna, which 
is limited in variety, but fairly abundant in individuals. The Hares 
are abundant and form distinct paths or runways, as shown in Fig. 9. 

Station I, Substation 3. Balaam^White Spruce Forest. This station 
included the forest traversed by a blazed trait from near the south- 
eastern part of Sta, I, 2, and extended northward to the clearing about 
the light-house (I, 7), and beyond it to the north rock clearing (I, 2). 
Most of the region occupied by the forest is of low relief, with an occa- 
sional low rock ridge or hill. The dominant tree was the Balsam Fir, 
with much Paper Birch and White Spruce. Where the forest was very 
dense, especially if due to the number of Balsams, the ground was densely 
shaded and there was almost no herbaceous ground cover; but wherever 
there was a small opening, due to a fallen tree, or where one had been 
cut down, there was an abundant growth of Large-leaved Aster and 
White-flowering Raspberry ; and it ■was in the midst of such conditions 
that young Balsams abounded. These were very characteristic plants in 
such conditions. In most cases at thick layer of humus covered the 
ground, but the tree growth was of small size. The common size of the 
Balsam was about 4 inches, the larger ones reaching 8 to 10 inches. The 
Birches averaged larger, usually about 6 inches, yio evidence of bums 
were seen, but probably many trees have been cut from this vicinitj, 
because of its proximity to the light-house, and the former Indian camp- 
ground now occupied by the fishermen. The Balsam appeared to become 
dominant at this place, as more young trees of this species were seen 
than of any other. 

The fauna found in this forest was rather limited, and doubtless 
great numbers of the insects which were taken in the clearing about the 
light-house (I. 7), bred in the adjacent forests. This is particulai-ly true 
of the Cerambycids and other wood infesting beetles, the wood-boring 



ECOLOOY OP ISIJJ ROYALE. 18 

Hymenoptera (Urocerus), and tlieir parasites. In addition to such spe- 
cies as feed upon Balsam, White Spruce and Paper Birch and their asso- 
ciated vegetation, there were those animals which are dependent npmi 
the shade, moisture, soil, decaying logs and other features aesociated 
with forests. To this class Ijelong certain insects which frequent decay- 
ing timber or the fungi growing apon them, and the earthworms of the 
soil, "the ground beetles or Carabidn, and the ground-inhabiting spiders, 
Lycosids. Some of the birds found were: Chickadee, Red-breasted 
Nnthatch. Golden-crowned Kinglet, Whitewinged Crossbill and Purple 
Finch. 

Station I, 4. Tamarack and Arbor Vitae or Whito Spruce Sicampg. 
This swamp Is located in one of ibe valleys near the head of Tonkin 
Bay, and extends back from the bay about one-fourth of a mile. It 
be^oB just back of the beach and is bordered by a strip of Alders, 
Paper Birch, Mountain Ash, young Balsams and White Spruces. The 
rock walls of this valley are abont 75 or 100 feet apart and are well 
shaded and covered by lichens and mosses, the south surface largely by 
lichens alone. Back of the marginaJ beach strip above mentioned, comes 
the dense growth of very large Arbor Vitae trees, intermingled with nu- 
merous large fallen trunks, partially decayed and covered with a dense 
growth of mosses. In the dryer places the ground is covered with a 
dense litter, and a thick damp or wet mass of mosses, but no pools of 
water. The undergrowth is composed of young Balsams, Birch and 
Oround Hemlock, Fig. 10. 

Proceeding farther np the valley, the Arbor Vitae is replaced by 
Balsams and Paper Birch; the forest is more open, and the amount of 
mosB on the ground is greatly reduced, and is replaced by a growth of 
Large-leaved Aster and large quantities of Ground Hemlock— all of 
this vegetation being indicative of mesophytic conditions. In this re- 
gion there are scattered pockets or small pools of water containing 
dogwoods. Still farther up the valley the Balsams and Arbor Vitae oon- 
tinue and Tamaracks are added, but no stnnding water was found. The 
valley turns, and returns to the bay on the north side of the ridge which 
bounds the Arbor Vitae swamp on the north; the entire basin is thus 
somewhat horseshoe shaped. The returning section becomes almost pure 
Tamarack and contains numerous small pools of water. The conspicu- 
ous feature of this environment is its jnngle-like character, the rapid ac- 
cumulation of litter and liumus, and the damp substratum. 

The fauna of such a bog is surprisingly limited in variety and amount. 
A few shells were found, as Pymmtdula (^onkkeitei anikoni/i and, in 
the small pools, Pisidium. The large numbers of Mosquitoes and Black 
Flies made up for all deficiencies, and were almost intolerable. The 
birds frequenting this forest were the Red-breasted >.'uthatch. Black- 
throated Green Warbler and Chickadee. 

Station I, 5. The Jack Pine Ridge. This ridge is located near the 
mouth of Conglomerate Bay. on the north shore. Some general idea 
of the location is given in Fiff. 11. which is a view looking toward the 
head of Conglomerate Bay. Just bock of the beadi, on an outcrop of 
conglomerate, was a small rock clearing, with Cladonia. Juniperus nana. 
and a wild rose. From here the trail extended through n narrow strip 
of forest, composed of Balsams. White Spruce and Arbor Vitae, with an 



14 MICHIGAN SURVEY, 1908. 

undergrowth of Balsam, Mountain Alder, and a ground co\~er of Lat^e- 
leaved Aster, and passed on through a belt of young growth of Birch, 
with the usual White-flowering Raspberry and Large-leaved Aster, Fiff. 
12, and up the face of an escarpment to the crest of the ridge, which 
had a height of about IflO feet above the lake level. Proui the nliundauce 
and characteristic growth of Jack Pines on this ridge, the station tak^ 
its name. Part of the ridge has been burneil over, as was shown by the 
burned and fallen timber, but the part to which our attention was given 
was apparently an oi'iginal growth. The Jack Pine was scattered, and 
largely occupied the depressions and the larger crevices. The ridge is 
fairly flat topped, but is occasionally broken by transverse gullies, which 
contain As|>enB, Birches, etc. The surface of the lava has weathered 
but little in some places, the original roche nioiitontt's surface 
being very clearly preserved, and the planed glacial surface but little 
eroded, Xear the escarpment, however, disintegration and decomposi- 
tion have been much more active, probably influenced in part by Jake 
waves at former lewis, thereby developing a talus slope, composed of an- 
gular blocks, and in sonic places foi-ming a stony soil. Alt intermediate 
stages are found between these two extremes. In addition to the large 
amount of bare rock surface, and that covered by only a thin layer of soil 
and vegetation, the shallowness of the soil is further evidenced by over- 
turned trees. Fig. 13. This soil is of residual and oi^anic origin, the 
crustaceous lichens and the ('ladonia-Uearlierry society, and later the 
Jack Pines, having contributed much to its foruiati<»n. The ex<-reinent of 
the Hares has also l>eeu an important factor in soil formation, and that 
of the Lynx also, though 1o a nmch less degree. 

The process of weathering must be relatively rapid on this ridge, 
because it is exposed to tlie winds at all seasons of the year, aud to the 
marked seasonal and daily changes of temperatui-e. The heat of the 
noonday sun is excessive, and the radiation from the nearly bare rock 
must be rapid, as it also is at night, so that the various influmioes con- 
sequent to temperature changes are allowed full play. Weathering i»- 
further favored by the irregularities of the surface, and the crevices, 
which allow the accumulation and downward conduction of this moisture, 
thus permitting the prying action of ice. 

In general, the succession of plant swieties on this ridge appears 
to be about as follows: Lichens are the jnoneers on the rock anrface, 
and these may be of several species. UinbiJicaria, and the crustaceous 
and foliaceous forms. As a soil dei-elops in the crevices or on the sur- 
faces, these are followed bv Cladonui. Hearberry, Sibhaldiopsia tridentata, 
Solidago, Diervilla dicrcilla (Bush Honeysuckle); and later, when the 
soil Itecomes deei)er. by Amelanchivr, Pniinin pennxylffinica (probably 
dispersed to these ridges by birds) and Junipents nana. The presence 
of the Small-toothed Aspen, willow aud an occasional Bireh pi*ol«ibly in- 
dicates the next society. In the shade of the Birches and Jack Pines 
dolidfiffo and Aster niairophyllun occur, if sufllcient soil is developed. 
From the character of the vegetation in the ravines which traversed the 
ridge, and ui>on the latus slope toward the bay. it is apparent that the 
next society lends to !»■ that of Birch and Aspen with some Balsam. 
Pennsylvania Cherry, yit. Alder; and a ground cover of Large-leaved 
Aster, Lai^-flowering Haspberr.x', (irotind Cornel and Lf/copodhnii. It 



ECOLOGT OF IBLB ROTALE. 15 

is clearly seen tbat nniODg these there are several elemeats ot the 
Balsam, White Spruce and Birch forest eociety, which tends to ulti- 
mately possess the ridge. 

The fauna of the ridge ib quite diversified, and there is a general 
faiinal correlation corresponding vith these successions of the regetation. 
Thus during the Lichen-Heath stage, ants and spiders, certain shells, 
and grasshoppers are abundant. As the soil becomes thicker or the 
creviees deepen, a subtraranean fauna, consisting of myriapodB, earth- 
worms, etc., develops. As shmbs and trees encroach in patches, the 
animals frequenting the open tend to perpetuate themselves mainly at 
the open margins. From this condition on, so far as the fauna is con- 
cerned, it is largely a question of an "opening" or a forest environment. 
Ko long as this habitat remains open, the grasshoppers, ants, spiders, 
butterflies, flies, and certain Hymenoptera, Hares and Bats are character- 
istic, and this condition tends to nmtinue as long as the trees are 
scattered. The Cicada is very characteristic of the Jack Pine stage, 
and although it occurs elsewhere in young Birches it is not so character- 
istic a.s on these hot ridges. With the advent of the Balsam-Birch society, 
which is slowly encroaching upon the ridges, the forms frequenting the 
open will disappear, or linger in the tfpen siHtts where local conditions 
have retarded the advance of the forest. (Inly a few birds were seen 
here, but Hares hud been numerous, as was ahcuvu by the large amount 
of excrement, and there was similar evidence of the occurrence of the 
Lynx. A bat was flushed frimi under a stone at the edge of the escarp- 
ment. 

Station I, f). Taniamck-Spiuci! Bog. This is a very small bog located 
at the base of the north slope of the -Jack Pine Ridge (I, 5), and roughly 
estimated aa about -30 by 300 feet in extent. The central part is 
covered with sphagnum, Cassandra, and a scattered growth of Labrador 
Tea. Widely scattered throughout the bog occur Tamaracks and Black 
Spruces, small Birches, Dwarf Cranberry, Cotton Grass and alders. No 
standing open water was found in this area, nor was the bottom quaking. 
Bordering the sphagnum zone is one of alders, willows, and a tall grass 
which merged into n zone of Tamaracks, willows, alders, Cassandra, and 
Balsam; Fig. 1). Along tlie western end a narrow strip of water, a 
few inches deep, was found, which flowed through a ravine across the 
ridge. Along this outlet the deeper soil and moisture has permitted 
the development of Balsam, Birch, Kmall-toothed Aspen, Mt. Maple, 
Ground Hemlock, Ground Cornel, Lai-ge-leaved Aster, and a few Black 
Ash trees. 

The fauna, like the vegetation, was not studied in detail, but the fol- 
lowing general relations were observed. In the open central Sphagnum- 
Cassandra society were numerous large ajit nests. A Toad \vas ob- 
Herved here: and the following birds: Golden-crowned Kinglet, White- 
throated Sjmrrow, Cedar Waxwing, and Black-throated Green War- 
bler. 

Station J. 7. Light-house Clearing. This was a small clearing which 
has been made about the Lij^t-house: it connects by a path to the fish- 
ing camp on Rock Harbor. It covers about half an acre, and was orip- 
nally, in all probability, a Balsam and Sprace forest like the surround- 
ing forest. A sod covered much of the gi-ound, and there were numerous 



16 MICHIGAN SURVEY, 1908. 

weeds, of whicL the Cow Parsnip nmbels furnished excellent places for 
collecting Syrphid flies, Cerambrcid beetles and Hymenoptera. 

The fauna of this clearing consisted largely of insects which fre- 
quent flowers, and butterflies which fly in open places ; but a few animals 
were found about the Light-house itself. The Chipping Sparrow bred 
in this clearing. Fig. 1. 

Station II. This station included the clearing at the mouth of the 
stream which drained Lake Benson, and which we called Benson Brook, 
and followed the biased trail to Sargent Lake, and on to McCargoe Gm'e. 
The clearing at the beginning of the trail at Rock Harbor marks the 
site of the former settlement called Ransom on the old maps. 

Station II, Substation 1. Ransom Clearing and Betigon Brook. The 
clearing was occupied by scattered Small-toothed Aspens and Birches, 
and was well sodded with grass and Bed Glover. Our attention was 
called to this locality because of the great number of Garter Snakes 
(Thamnophis sirtalia) which were found there. These snakes were veri' 
abundant in a small area east of the mouth of the brook, in a rauk 
growth of grass and among some mils. 

The brook contained but little life, although it was carefully examined 
near its mouth and farther back where the trail ci-osses the bi-ook. 
Only a few dead Physa were found, and a louug fish, at the mouth of tlie 
brook. 

Station II, Substation 2. Tamarack Swamp. This is a long swamp 
which is crossed by the trail, and which contains a scattered tree 
growth of Tamaracks, Black Spruces and Arbor Vitae, a dense shrub 
growth of Cassandra and Labrador Tea, and a ground cover of Sphagnum 
and Pitcher Plants. While no water was seen on the surface, it was 
a wet swamp. 

This locality was only examined for birds and mammals. 

Station II, Substation 3. Rock Ridges. This station number is given 
to the open rock ridges which were crossed by the trail between II, 2 
and Sargent Lake. These ridges have been burned over and are largely 
destitute of soil and the Cladonia growth usually found on other rock 
ridges. Small-toothed A8i)en8 generally border these ridges which have 
a northeasterly southwesterly direction. The heat during the middle 
of the day is excessive. The scant vegetation which grows in some 
crevices and depressions in the rock leaves an open area which is decided- 
ly favorable for grasshoppers. In some places they were exceedingly 
abundant and many ridges were examined almost solely for their grass- 
hopper fauna. In the dry soil on one ridge an anti-lion laiTa was 
found in the dust at the. base of its funnel, and a large (iaiter Snake 
was taken on another. The grasslioppers found liei-e were Vliloealtin 
conspi'rsa and abdominalis. t'ircotcttix ven-ucttUiliis. MvlanopUiS 
aiaskaiius and faaciatus. 

Station II, Substation -J. McCargoe Core. This station simply marks 
the location of the end of (he trail, and the cove where a few molluscs 
were found. There were dead shells of Anod<nita grandis footiana, 
which were abundant at the edge of the water. Here upon the low 
rocky shore 'were also found specimens of Limnaca stagnaUs. 

Station II, Substation 5. Forbes Lake. The examination of this 
small lake was mainly confined to the north shore, as the south shore 



Oeologlcal Burvejr of MkhlgaD, 



I Report for 1008. 



PIG. 1. TUB LIGUT-II0U8I-: AT ROCK HARBOR, I8LK ROYALE. 



FIO, 2, CLIFFS BETWEEN TONKIN AND CONGIX>MBKATE 



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G«ol(«leal Survey of Michigan. 



Anniuil Report (or IDOS. 



AT THE HEAD OF 



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PIG. 4. SAND BEACH AT THE HEAD OF CONGLOMEHATE BAY (1, 1). 



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0«al(tfnl Survey of Michigan. 



Annual Report for 1608. 



FIG. 5. BEACH POOL (I, 1) NEAR TONKIN BAY. 



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Gcologfe*! Snn'e)' of Ulcblgaa. 



Anuusl Reirart tor IBOS. 



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Qeologleal Survej' af MlcblgBD. 



, 2} NORTH OF THK LIGHT- 



B' THE LIOHT-HOU8B, ADJACEWT 



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Geoiosiol Surve? o( Hiehlgan, 



ADDual Keport tm 1S06. 



FlU. 10. ARBOR VITAR BOG <I, 41 NICAR TONKIN BAY. 



VIEW FROM THE JACK PINE RIDGR (I. r>). LOOKING TOWARD THE HRA1> 
OP COSttLOMERATK BAY. 



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Otologlcal Survey ot Micblgan. 



I Report for 1)>08. 



NE KIDOK (I, 5) rOXOLnMERATK BAY. 



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a«oI«|lc«l Survey of HIchlsan. 



1 Report for 1908. 



14. SPHAGNUM-BLACK SPRUCE BOG (I, '() NEAR THE JACK PINE RIDGE. 




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Geological Surve; of Michigan. Annual Report for 1908. 



-I.RI!SII ZONE AND DELTA AT THE HEAD OP ROCK HARBOR (III, 3). 






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Qtoloflo] Barv«y c 



jal Report for 1908. 



*'"'■ IT. GXPOBSD SECTION OF SPIT FORMED AS THE WATER LEVEL HAS 

LOWERED IN ROCK SAItBOR, NEAR THE BEGINNING OF TUE 

TRAIL TO BDMNER LAKE (III, 4). 



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GMlogtrsl Survey of UtcblKan. 



I Report for 1908. 



FIG. 18. SUMXKll LAKE (III. .".I. KASTERN KMI. 



PIG. 19. WESTERN END OF SLM.NER LAKE (III, B). 



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Geological Survey of Mlcblgan. 



Annual Report for 1908. 



MAUCIN OP SUMNER LAKE (III, 0). 



FIO. 21. 80UTBBA8TBKN 



OF SUHNRR LAKE C 



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G«i>1oKlcal Survey o( Michigan. 



I Report lor 1908. 



FIG. 22. WESTERN END OF SUMKBB LAKE (III, 6). 



PIG. 23. KOBTUERN SHORE OF RTjMNEB IJ.KB (III, 



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Oeoli«lcal Sairej ot HkbtgaD. 



FIG. 24. ROCK OPENING ABOirr CAMP ON 



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FIG. 26. ROCK OPENINO AT 8ISK0WIT CAMP (V, 3). 



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Geoloiilcal Hurvey of Mlchigaa 



FIO. 26. ROfK OPKMNG ON SISKOWIT I 



! (V, 4)-^ I 



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Oeoloilral Surrey of Mlcblgin. 



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FIG. 28. ANT NEST IN TUB OrBNlNO AT THE 8I8K0WIT CAMP (V. 



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I. 88. FABTHEB UP THE SAME SLOPE AS IN FIG. 32 AND ADJACENT TO IT. 



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FIG. 85. LOOKING t'P THE 8LOPE OX THK WESTEBN POBTION OF STATION V. 2. 



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FIG. 39. OPEN SPACE IN TOE BAI.»AM-BIKCII FOEEST (V. 4). 



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FIG. 40. OPEX 1 



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FIG. 40. MAl'LE K0HK5T ON THE DBSOR TRAIL (III, Ol). 



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ECOLOGY OF ISLE ROTALB. 17 

ie rocky and eteep, with Birehee and other trecB growing down to the 
water. The north shore has been largely burned over, and is being re- 
placed by Birches and Small-toothed Aspens, whioh are now dominant; 
the nnde^Towth consists of alders and the abundant Large-leaved Ast^. 
The water in the lake is brownish. At the western end there are White 
Waterlilies, near the shore Yellow Waterlilies, Calthn pahistrig, Equiae- 
turn, and farther back Cassandra and alders, Tamarack, Arbor Vitae, 
and Black Sprnce. 

On the north shore a rocky point projects into the water, and east of 
this along the shore is a floating sphagnum bog, ranging in width from 
aboDt 40 to 100 feet and containing Pitcher Plants, Low Cranberry, 
Buckbean, scattered sedges and Blue Flags, and a shrub growth of 
Cawandra, Labrador Tea and Wild Rosemary. Scattered trees of 
Tamaracks, Arbor Yitae, and Black Spruce grow to tiie edge of the 
water. Water stninds in the small depressions over this b<^. 

The fauna was not studied in detail, but the forms collected were 
as follows; The spider, Pardosn (jIaciaJis, with" egg masses, was found 
mniung about over the wet sphagnum; a dragonfly, Aeschna, was seen 
on wing; two species of grasshoppers were found in the wet Sphagnnm; 
Melanoplus extremv8 and, in the wetter places, nymphs of Mecoatethus 
lineatua were quite abundant. There were also great numbers of 
mosquitoes and Black Flies, Upou some driftwood near the end of 
the lake was found PJii/m gyrina (Ko. 71 A.). Yellow Perch were so 
abandant in this lake that locally it is called Perch Lake. A Canada 
Jay was seen in the top of a tree. 

titatioii III. ^Vrlltn■}l■ tJnd of Itocl- Harbor. This station was in- 
tended to include those localities near the western end of Bock Harbor. 

Station HI, Substation 1 and 2. fimaHl Island. The general character 
of this island is shown in Fig. 15. This is a small, rocky, wooded island, 
the trees consisting of one large White Pine, about 14 inches in diameter, 
Arbor Vitae, Birch, Balsam, and White Spruce, the dominant ones being 
the Balsam, Arbor Vitae and Birch, with a shrub growth of Mt. Alder, 
Willow, Nine-bark, Monntain Ash, A mcUinchier alnifoUo. Upon the rock 
occurred Cladonia, Bearberry, and IjOw Juniper, and toward the western 
end of the island, where the trees shade the ground, grew Lycopodium 
complanatvm, mosses and Clintonia borcalis. 

Gf the fauna, the bird life only was examined; Cedar Birds and a 
Song sparrow nested here, the former being quite abundant. 

The submerged western end of this island formed Station III, 2. 
The bottom was composed of sand and angular rocks. In the shallower 
water Anodonta grandis footiana valves were found, and live animals 
in water about 18 Inches deep. These rocks also furnished a number 
of Limnaea atagnaUs, and a dead specimen of Planorbis bicarinatus. A 
few scattered rushes (i^eirpus) grew at this place. 

Station III. Substation S and S. Head of Rock Harbor. These 
stations include the delta at the month of the largest stream flowing into 
the Harbor, Station 3, and the sandy and rocky shallow water zone 
extending from III, 2 around the head of the Harbor, Station 6, 

The genera] character of the delta, III, 3, region is shown in Fig. 16. 
This small delta had been formed by a small sluggish brown-stained 
brook, 15 or 20 feet wide, which enters the Harbor at this point The 



hyCOOgIC 



18 MICHIGAN SURVEY. 1908. 

chanDjel contained a growth of ValHaneria 8}}irali8, Potanwgeton orispui, 
and the banks supported a growth of sedges, Lycopodium- complanatum, 
Clintonia boreaUs, alders and Mountain Ash. The surface of the delta 
is strewn with driftwood and other plant remalDS, upon a clean sandy 
bottom. Nearer the shore, upon a muddy bottom, were found an abund- 
ance of Amphipod crustaceans, HyaUlla knickerbockeri, Gammerus 
Imnaeua, and the small bivalve molluscs, Pisidmm. The fiesh water 
sponge, kpongiUa lacmtrix, was found here, and water striders, Gerris 
remigia. were found on the surface. Individuals were abundant, bo that 
the fauna is relatively varied. 

Substation 6 included the southwestern coves of the Harbor, The 
bottom was rocky, and covered in places with much sand ; the water was 
shallow and contained, near the shore, many patches of rushes, Scirpue 
and Kquiaetum. Anoilnnta grandis footiana, Limiiaca xtagnalis and 
Piaidium were the characteristic molluscs, and a few flsh were found. 

The protected character of the shore is noteworthy, as no beach is 
developed, because "the coves are protected from the hea>7 lake waves. 
Another characteristic feature is the sand bottom. Tliis sand is carried 
toward the head of the Harbor by the currents. Even at higher Lake 
levels, this Harbor was sandy, as is shown by the sand banks on the 
north shore, and these are being re-worted by currents and waves 
and carried up the Harbor. The spit developing from the south 
phore. Fig. 17, illustrates this. 

Station III, Subatationa 4 and 5. Sumner Lake attd Tifiil. The trail 
to Sumner Lake (III, 4), begins on the south shore of Koek Harhoi- 
and extends south about one-half mile to Sumner Lake. It passes) 
through a second growth of Birch and Aspen (which has followed a 
burn), a small Arbor Yitae swamp, over a rock ridge to the north shore 
of the lake, where there are a few large Norway Pines, from 12 to 
15 inches in diameter, and a few White Pines. But little attention was 
given to the life along the trail, although a few observations on the bird 
life were made, and some mammals were trapped. However, Sumner 
Lake proved to be such an intei-esting locality that attention was given to 
it more especially than to the trail. This lake has many of the charac- 
teristics of a large lily pond, because the White Wateriilies and Pota- 
mogetons form such a wide belt around the lake. Pigs. lS-^3. In pass- 
ing from the interior of the lake toward the shore, the following zones 
of v^etation are fouud: The bulrush zone, which is well developed, 
with its denser growth about the eastern end ; then the Yellow Water- 
lilies, followed by the dense sedge zone which produces a substratum. 
In the eastern and western ends of the lake the water gradually shal- 
lows; but on the sides the change is more abrupt, thns interrupting 
the shallow water zone of sedges, as shown on the north shore, 
Fig. 23. This encroachment of vegetation upon each end of the lake 
is very marked, and is much more extensive at the eastern end, 
where the lake is drained into the bead of Conglomerate Bay by a small 
brook. The encroachment at the western end of the lake is well shown 
in Fig. 19. A partial view of the eastern end of the lake is given in Fuf. 
18. The sedge zone contains a variety of plants, including several or- 
chids. Iris, Pitcher Plants, Buckbean, scattered Eriophorum and Sphag- 
num, Cassandra end Andromeda. The substratum is quaking and sinks 



:byCOOgIC 



KCOLOQY OF ISLB ROYALE. 10 

several inches below the water level with the weight of one's body; occa- 
sionally small but deep holes are found through this subetratum, and 
care must be taken to avoid them. This zone is very broad and contains 
an abundance of life. Outside the sedge zone occur alders and Tama- 
rncks, which border the forests at the base of the slopes. 

The fauna of the open Waterlily, Bulrush and Potamogeton zone con- 
sists of insects flying over the water, such as the dragonfly, Aeschna 
and the leaf beetles Donada, which abound, especially about the Yellow 
Waterlilies. On the surface film were water stridera, Oetris margma- 
ius. and whirligig beetles, Gyrinidae. Sticklebacks were abundant, and 
are quite characteristic of such waters, as is another small ilsh. Loons 
were frequently seen here, and also a Hooded Merganser. Toward the 
onter margin of this zone "whei-e the lilies are often closely matted on 
the surface, the insect life and the surface film fauna are the most 
abundant. A live mussel, Anodonta grandis footiana, was found on the 
bottom; and the snails, Planorhis campanulaius and parvus, were found 
in small pools in this sedge zone. The bottom in this vicinity, and that 
bordering the water margin of the ridges, is covered with a mass of 
partly floating debris, the appearance of which suggested to Wood, who 
first observed it, that something had exploded and scattered the strands 
of debris about the surface. It is not improbable that the formation 
of marsb gases will adequately explain this phenomenon, (Cf. Peuhailow, 
Science Vol. 22, 1905, pp. 794-7%). 

The dragonfiics were Enallagma hageni, Acschna, Somatocblora 8hurt- 
h/fi. and Lacorhina proximo, the last being very abundant. 

Where the sedge zone wan absent, as at our raft landing at the end of 
the trail, an abundance of needles, leaves and twigs from the over- 
hanging conifers and hardwoood had accumulated at the shore, and were 
stained almost black. The water of the lake ia brownish. At this 
point a number of invertebrates wei-e taken, including shells, leeches, 
insects, etc. 

Station IT. Tobin Harbor and Vicinitj/. As very little time was 
spent at this station, the description will be correspondingly brief. Tobin 
Harbor is a deep, nari-ow, protected bay, similar to that at the bead of 
Rock Harbor, but narrower. The adjacent hills are forest covered, 
largely with Aspen and White Birch. In the vicinity of Neutson's Resort 
there is a large, cleai-ed area. Mattson's resort is located on an island in 
this Harbor. The most marked scenic feature of the Island, Monument 
Rock, is on the north side of Tobin Harbor. 

It is a noticeable fact that many of the low islands in Tobin Harbor, 
and especially those near its eastern end, are clothed with vegetation 
close to the edge of the water. They are thus in marked contrast with 
the islands along the southern shore, and to the various points of rocks 
which project into the water. 

Upon a small island, Number 14 on the Land Office map, were found 
small rock beach pools, just above or near the height of the usual quiet 
weather waves. The water in one was about a foot in depth and con- 
tained a very small amount of algal growth. In this pool were found 
water stridens, Qerris remigis, a few other insects (No. 30), and small tad- 
poles. In another small pool about 10 inches above lake level, and with a 
temperature of 77"^ (the Harbor water having a temperature at the. 



20 MICHIGAN SURVEY, 1908. 

time of 50° F,), were many Bpecies of lAmmiea cataacopium and a 
few adult Physa, vVlgae were oiily aeen in tlie creviceB. 

Station IT, Substations 1 and 2. Rock Pools arui Scovill Point. 
BcoTill Point ib an almost bai'e, glacially planed, narrow and low rock 
ridge, projecting out into the lake. Xumerous small faults occur on 
the sloping southern side, and these, supplemented bv the waves, etc.. 
have produced rock poolB. In one case a long row of pools occurred 
along the line of the fault. The presence of tadpoles about an inch 
long wonid suggest that these pools have some duration. The higher 
Rummer waves might also reach many of these pools. Water stridei-s. 
Oerria, are abundant upon the surface, and a large deep ravine, nenr 
the lake level, contained Sticklebacks, but no shells were observed in any 
of these pools. 

Station IV, Substation 3. Bayou East of the Monunvent Rock Trail. 
This is a very Bmall pond which is connected with the Harbor by 
a small stream just large enough to admit a row boat. It illustrates the 
last stage of separation of the valleys from the Harbor, as only a very 
slight fall of the lake level would completely isolate it. In this particular 
ease the outlet is on the nouth side, and not at one end as is iisnally 
the case. The central part of the pond is open water and is surrounded 
by an almost complete zone of Yellow Waterlilies, and a sedge zone con- 
taining several low shrubs. The Waterlilies were badly infested by a 
small le^ beetle, Galcrucella- nifinpkaea; larvae, pupae, and freshly 
emerged beetles were taken. A few dead shells of Anodonta marginata 
and one of Limnaea wegasonia (the only spe<nnien taken upon the 
island) were secured here. An extensive suspended flocculent mud cov- 
ered the bottom, so that molluscs could not obtain a foothold. At the west- 
ern end of this pond innumerable small tadpoles formed an almost 
compact pavement upon (he bottom at the edge of the watci'. A few 
dragonfliea were seen, but were not captured. 

Station IV, Substation 4, S and 9. Forest on tlie Greenstone Range. 
These three stations are combined Itecause they are related to the forest 
occupying the Gi-eeustone range. The trail to Monument Rock (IV, 4) 
begins on the north shore of Tobin Harbor and extends northwest about 
one-half mile to Monument Rock. The forest is dense and is apparently 
a second growth of Balsam, White Spruce, Birch and Aspen, with un- 
derbrush of Mountain Alder, Mountain Ash, Ground Hemlock, and a 
ground cover of Few-flowered Cranberry, Clintonia boreaUs, IJnnea 
borealis, and AVild Sarsaparllln. In the moist places was found Ground 
Cornel, Aster macrophyllus, an Equisetum. Lycopodium. and, in wet 
places of the swamp traversed, the Buckbean and Bkunk Cabbage. For 
some distance on the slope down from the base of Monument Rock occur 
large blocks which are covered by a dense mat of mosses, and the 
ground is covered with a thick layer of humus, so that the general 
appearance of the vegetation is that of a mesophytic forest. 

The trail up the Greenstone (IV, 8), begins at the mouth of a small 
brook at the head of Tobin Harbor, and follows the crest of an open 
burned over ridge southeast for about half a mile. This ridge contains 
a scanty growth- of Amelanchier oligocarpa and alnlfolta, Prunus penn- 
ayl&tnica, Jack Pine, wild rose, SoUdago, Bearberry and Yarrow. From 
the end of this ridge a valley crosses to the north and contains large 



ECOLOGY OP ISLE ROYALB. . 21 

.Vspens, Tamarack, Norway aud White Fine, and an underbrush of 
Speckled Alder and Ground Hemlock. After croBsing this depreeston, 
the trail aecendg the elope and croBses tlie burned ridges where there 
is a growth of Birch and ^Vapeu. The slope increases more abruptly 
as the crest of the Greenstone is approached. This is the vicinity of an 
old signal station and has an elevation of about 4G0 feet, according to 
the I^ke Survey. This forest along the crest comprised Station IV, 9. 
The large trees stand above the snrrounding second growth, on the 
burned area, and can be seen for some distance. The Balsam, Birch 
and Quaking Aspen are the dominant trees, the Balsams reaching a 
diameter of about 10 to 12 inches, and the Birches and Aspens abont 12 
to 15 inches. The shrub growth is composed of Mountain Maple, Ground 
Ilemlock, and the Few flowered Cranberry, the ground cover of Dier- 
riUa diervtlla, I^rge-flowering Raspberry, Aster macrophyllua, CUntonia 
bort-alis, Lmnea horcalis, Lycopodiutn, Wild Sarsaparilla and Brake 
Fern; the White and Black iSpruces being only occasionally seen. This 
forest produced dense shade. Fallen timber is abundant in places, but 
no signs of fire were observed. This ridge was bounded on the north 
by a cliff of perhaps 20 to 30 feet, below which was a, long talus slope 
covered with Birch, Aspen and Balsam. From the top of this ridge there 
is a splendid view to the north. The crest was followed west to a small 
open burned area where DiervUla diertdlla and Large-flowering Baspberry ' 
were abundant. The leaves of the latter were badly perforated by the 
abundant grasshoppers, Hibbincim tvicrmilatiis and Metanophis alaa- 
kanvs. In the deep wood a Ti-ee Toad. Sj/la versicolor, was found, and 
Varying Hares and Red Squirrels were seen. 

Station IV, Substation 5. Clearing, and Vicinity of -Veu(«on'« Resort 
(Park Place). There is a rather extensive clearing at Neutson's Resort, 
so that very little collecting was done in this virinity. A collection of 
grasshoppers was made here by Brown and Wood, nud the following 
list of plants was made by the former from the same vicinity: White 
Spruce, Birch, Aspen, Mountain Alder. Junipcrua nana, Wild Bed 
or Pennsylvania Cherry, Red and White Clover, Bush Honeysuckle, Fra- 
garia vesoa, Cow Parsnip, and lAjcopodium complanatum. 

The grasshoppers were: CMoenltis ahdominalis, Camnula pelludda. 
Hippixcut tiibcreiilatns, Circotettix verruculatus, Melanoplus alatkamm. 
and hnrord. Two butterflies, Argynnis at)antis and Pyrameia cardui, 
and the dragonfly Lcstes unguinilatus were also taken here. In Rock 
Harbor, at Neutson's, leech egg capsules of Nephalopsis obscure, and 
Physa were taken. On a small island across the Harbor to the south, 
in a Sphagnum, I'itcher Plant and Tamarack swamp, a number of Wood 
Frogs, i liana cmUabrigcnsis) were taken. 

A Red-bellied Snake (Storeria occipitonia^data) was reported to have 
been killed in the clearing, during July. . 

Station IT, Substation 6. Small Island in Tobin Harbor. This sta- 
tion includes the sedges and shallow water at the west end of a small 
island in Tobin Harbor. The bottom was covered with sand and large 
angular blocks of rock. Limnaea stagnalis was very abundant and oc- 
corred in water with a depth of about three feet. The shells are very 
fra^le. Upon the rocks Physa occurred in limited numbers and was wide- 
ly scattered, but they were abundant on the stems of the sedges not far 



22 MICHtGAN 8URVBY. 1908. 

below the surface. The yoang of L. atagnalia occurred with the Phifsa. 
A specimen of Qordiua aqualicus was secured here. 

Station IV, Substation 7. Hca4 of Tobin Harbor. A small brook 
empties into the head of the Harbor, and its brown water brings into the 
bay quantities of vegetable remains and flocculent debris. These cover 
the bottom near the mouth of the stream, and although carefully ex- 
amined, were found to be singularly free of animals. Upon patches of 
Potamogetoti perfoHatum occurred a few Phyaa, and in the shallow 
water a few flsh were found. (No, 35.) 

Station V. 8i»lto\cit Bay, Lake, and Vicinity. This station includes 
all the localities in the vicinity of Hiskowit Bay and Siskowit Lake, ex- 
cept that of VIII, '04, which is at the head of Siskowit Bay. 

Station V, Svitstation 1 and 3. The Bay Beach at Camp, and the 
Rock Clearing. The beach (V, 1), is protected from the waves of the 
open lake and bay by large off-shore islands. The rock composing it is 
vesicular lava, and slopes lo the water at an angle of ."ibout 10 degrees. 

The Rock Clearing aboot Camp (V, 3), begins at the beach and ex- 
tends up the slope backward ta the Balsam, Spruce and Birch forest os 
a park-like opening. The soil, mainly of residual and humic origin, is 
very shallow and completely lacking in places. The surface of tbe rock 
is rough, showing that it has been eroded simee glaciation. In places 
the rocks are more or leas covered by crustaceous and foliaceous lichens, 
or, where there is more humua or soil, by Cladonia, Fig. 24. Where 
the soil is deeper is found Bearberry, Narrow-leaved Cow-wheat, Soli- 
dago, grasses and moss. A scattered shrub growth is composed of 
JunipOrus nana and AmelanchAer, Fig. 25 and 26, and the bordering 
tree growth consists of Balsam, Black and White Spruce, Birch and 
Arbor Vitae. In the sbade of these bordering trees. Aster macrophyUus 
develops in abundance. The above description also outlines the prob- 
able succession of plant societies upon this surface, all stages of which 
are now to be found witliin tliis area. Many smaller patches of this 
open condition are found scattered through the forest and are becoming 
shaded and converted into the forest as a soil develops, Fig. 27. 

The fauna of the openings is rather characteristic and abundant. 
Of course many species range over a variety of plant societies and only 
reci^nize a clearing or forest society, and not their varieties. Yet 
others are much more sensitive to smaller environmental units. This 
is well shown by certain ants. Ant nests of Formica fusca as illustrated 
b.V Fig. 28, and certain spiders, as Pardosa, are quite characteristic of 
the Cladonia-Bearberry plant society. The opening, as contrasted with 
the forest, is frequented by grasshoppers, such as Melanoplus faaciatu»; 
butterflies, as Argynnia and Bamlurchia arthemia; and the dragonflies, 
Sympctrum, which were very abundant and characteristic (Aechna 
patrols the margins of such openings). The robber fly, Asilua annttlatua, 
frequents such open sunny areas where animal food abounds. 

The vertebrate frequenters of the open were the Flickers, which were 
observed by McCreary to destroy ant nests. Several of such demolished 
nests were seen. Toward night a nocturnal association frequented the 
■open; the bats on wing; and the hares came from the forest to feed, 
having been in hiding during the day. 

No doubt the presence of this opening, in part, determined the location 

■ D,s ZK;l„COOgIC 



ECOLOGY OF ISLE ROYALB. 23 

of the log cabin used by our party. The logs were thoponghly infested 
by beetle larvae, and attracted vast numbers of parasitic Hymenoptera. 
For this reason, insect collecting abont the cabin was of an exceptional 
character, and included a considerable variety of species. From the 
adjacent forest a number of trees had been cut, but this did not material- 
ly inflaence this locality, except near the shore. 

Station V, Substation 2. Heath Zone and Beach. This locality in- 
cludes a strip of rocky coast on the south shore of the island, a short 
distance east of the entrance of Biskowit Bay, Fig. 29, and extends from 
the edge of the water back to the forest. This is an exposed section of 
the coast and is unprotected by offshore islands, so that easterly storms 
from the open lake have full sweep on this shore. The slope is a fairly 
uniform rock surface, with an upward slant of about 10 degrees, and 
is composed of amygdaloidal lava. Crevices of various dimensions, from 
a mere crack to a deep rock ravine, extend obliquely up the slope. One 
of these ravines, the only large one, divides this station into two sections 
east and west. The eastern section of the slope is covered by a Cladonia- 
Junipents procumiens society, while the western section is occupied by 
a Cladonia-Junipcrug fWrta-Hnckleberry society. Thus there are three 
fairly well defined natural divisions of this part of the coast, the beach, 
tbeprocumhcns. and the nana societies. 

1. The Beach. The low angle of the slope, and the exposed sitaation 
and deep offshore water all combine to make the beach zones quite wide 
(four or Ave paces) upon this slope. No collections were made upon the 
riubnierged beach and only a' few specimens wen; taken upon the lower. 
The characteristic species, however, were a small hemipterous shore in- 
sect, Salda ligata, a caddis fly, and ants. Above the lower beach is a 
wide upper one, characterized by a dark green moss (Grimmia) and 
crustaceoas lichens. 

A number of rock pools occupy the oblique crevices which extend 
up the beach. The largest of these is shown in Fig. SO. This is a pool 
about 4x8 feet in diameter and contains about 15 to 18 inches of 
water. On the surface of the water were fragments of insects, water 
striders, Qerris remigis, and on the bottom, dragonfly nymphs (No. 14), 
while caddis fly larvae crawled upon the sides and bottom. No algal 
growth was visible. The character of this insect life suggests a pool 
of some duration, but the absence of shells suggests a lack of permanent 
water. Numerous baein-like depressions, a few inches in depth, occur 
on the lower beach and on the foliaceons lichen-covered portions of the 
middle beach. The sharp angles of some of the pools show that these 
are occasionally produced by the removal of small blocks of rock. Most 
of the pools, however, occur in crevices. From one of the large pools 
a frog, Rana clamitans (No. 120), was taken, clearly showing how tad- 
poles may reach, such pools. 

In the crevices and behind angular rock projections occur Harebells, 
Yarrow, Ninebark, and an interesting succulent Saxifrage, Sasifraga 
aizoon, Fig. SI, and some grasses. In the crustaceous lichen zone is 
a greenish moss, Orimmia, and in the crevices are Bearberry, Juniperua 
procunibena, and Arbor Vitae. 

2. The CUtdonia-Juniperua procunibcna Society of the Eastern Section, t ^ 
About ten paces farthei- up the slope, Fig. 33, this crevice society spread^''- 



24 MICHIGAN SURVEY, 1S108. 

out, and, with the addition of Cladonia and some Jumperus nana, fomw 
internipted patches or streaks, Fig. 33, which farther up the slope fuse 
and form a solid mat, completely covering the surface of the rock, Figs. 
S3 and 34. The dominant forms are Juniperus procumbens and certain 
species 0/ Cladonia. This was the only place where J. procumbens was 
found growing on such an extensive scale, or associated in abundance 
with Cladonia. This formed a novel and beautiful sight, the light-colored 
patches of the Reindeer-lichens in places intermingled with the bright 
green of the procanibeng to form a variegated mat. The beauty of color 
and pattern is lost in a general view, as in Fig. Si. The White Spruce 
invades the slope in crevices, just in advance of the solid mat formation, 
but the soil is so shallow that it may be blown over, as is shown in 
Fig. 33. Procumbens grows bo densely and close to the ground that 
it greatly favors the formation and retention of the soil, and it appai-- 
ently precedes, on this slope, the Cladonia. In the large crevices within 
this zone grow patches of White Pine, Balsam, Mountain Alder, Spruce, 
Birch and Arbor Vite. The general relations of this slope can easily 
be seen by a comparison of Figs. 32, 33 and 34, in which is shown the 
transition from the bare wave washed beach, the flat growing lichens, 
the pioneers of the mat formation invading the crevices, and the domi- 
nance of the J. procumbens -Cladonia society with its scattered trees, up 
the slope into the Balsam-Bpruce forest. This same order probably also 
expresses the succession of plant societies at this place. As previously 
mentioned, the fauna of the lower beach is quite limited, the greater 
variety occurring in the pools; but farther up the slope appear various 
forms which frequent the open. On the scattered part of the Cladonia- 
procunibens zone occurred the spiders, Pardosa glaoialia and 8temali8, 
the grasshoppers, Circotctiix verrucuJatus, Melanoplus alaskantts and 
faadatus. A ground beetle, Pterostichua femoraUs was found under 
CUtdonta, and under similar conditions were found an abundance of 
shells, Aoanthinula harpa, Strobilopa virgo, Vertigo tridentata, Vitrina 
limpida, binneyana, Euconulua fulvus, Euconulua cher»inus polygyratws, 
Zonitoidea arborea milium, AgrioHmaiB oampeatria, Pyramidula cronk- 
heitei anthonyi, Helicodicua parallelus, and Cochlicopa lubrica. The 
number of these shells which have a distinctly boreal range is particular- 
ly noteworthy, suggesting that such a habitat has some of the character- 
istics of a "boreal island." 

3- I'A« Cladonior Juniper us nana Society of the Weatern Section. 
Here, as at the eastern section of this slope, the bare lake beach bounds 
this area shoreward. A general view up this slope is shown in Fig. 
35. The bare wave-washed lower beach is in the foreground, and the 
green moss and light colored lichen zone is a broad belt above it, followed 
in tnm by foliaceous lichens, and in the crevices by Aspen. The rock 
surface is considerably router than that of the east beach. In general 
appearance this beach is much more like that about the camp at Siskowit 
Bay (V, 3) than the Cladoniaprocumbcna section, and contains more 
of the Low Janiper rather than the Procumbent Juniper. There is 
also much more exposed rock, and a miich more diversified flora. In 
places the Low Huckleberries are very abundant, while they are not 
at all conspicuoQB on the eastern section. To get an idea of the general 
appearances Figures 3-i and 36 should be cmnpared. ■ 



ECOLOGY OF ISLE ROYALE, 25 

Figure 34 shows the marked dominance of proctniibcnK, which was not 
abandaut oo the west slope. The plant life is more varied, with the 
foliaceons lichens and bunches of corat-like Cladonia. ferua, grasses, Soli- 
dago, and the willows and aspens in the crevices, Fig. 37. These forms 
give a very different aspect to the pioneer society from that of one 
composed of a Cladonia-procumhena mat. The open or patch like char- 
acter of this society suggests' that the retarded development of the 
Tegetatiooal cover may be related in some way to the scarcity of J. 
procumbens, which is such an excellent agent in soil formation. But 
■why this shrub should not thrive here is not known. 

Associated with these conditions wei-e the snail Polygyra alboUibris, 
and the grasshopper MeUinoplus faaoiatus. The absence of the dense 
mat, and less soil, greatly reduced the variety of animals frequenting 
snch conditions. ' 

Tal:ing the station as a whole (V, 2), it is one of the most interesting 
places seen on the island. The beauty of the variegated Cladonia mat, 
the extensive area of the open habitat, the boreal character of the lichens, 
the Saxifrage and many of the shells, the apparent completeness of the 
lueservatioa of the stages in the transformation fi-om the lower beach 
back to the forest, all combine to make this situation one of the most 
interesting and important of those examined.* 

Statioji T, Substation J. Trail through BaUamrSprucc Forest. This 
station begins at the opening about camp (V, 3) and extends northwest 
to the south shore of Siskowit Lake, opposite the eastern end of an 
elongated island. The topography of the region traversed is one of low 
relief, with only occasional low rocks, hills, or ridges, la to 20 feet in 
height, and a few shallow and moist ravines. A thick layer of humus 
covers the surface, except on the ridges. The trail first passes through 
a forest of White Spruce, Balsam, Birch and scattered Tamaracks. 
Among these trees are many fresh windfalls, due to the winds and the 
shallow soil. In the more shaded portions the ground cover consists 
of a dense growth of mosses, liverworts, Fig. 38, with Aster macrophylliis , 
in the less shaded portions. There arc open patches 10 to 15 feet in 
diameter scattered about through the forest, especially on low rock 
ridges, which contain a growth of Cladonia, and illustrate the last stages 
of the decline of the openings. 

In the moist depressions was found an abundance of Round-leaved 
Cornel, alder, and also Ground Hemlock, Mountain Ash, Balsam, White 
Spruce, and the Ground Pine (abundant.) There were many fallen 
and decayed \o^. In and characterizing the more open places, such as 
were a^ociated with lar^ Birches, are the Large-flowering Rasp- 



liien mre the moat Important papers on tbJa subject, 

IMS. EcokwIeBl Distribution an Incentive to the Study of Llchena. HryoloKiat, 5, pp. 39-40, 
IMS. BomeCiHnniini Trp«a o( I.lchen tormallons. Bull. Torrey not. Club., 30, pp. i\2-il6. 
IWS. Some Talua Cladonia, Formations. Bot. Ciiu., as, pp. ie.'i-208, 
IMM. A Llclun Bodety ot Sandatone Riprap. Bot. Gaz., 3S, pp. 26S-2S1. 
CcmtributlonB to the Btudr of Lichens of Ulnneiota: 

I. Llehanj ot the Laka ot the Woods. Uinn. Bol. Stud.. 1. 18H. pp. W3-70I 
IL Uchent ot Hlnneap^ and Vidnlty. Ulnn, Bol. Stud.. I, IBH. pp. 703-735. 

III. The Rock Llchena ot Taylon Falls. Ulnn. Bot. Stud., 2. isOS. pp. 1-18 

IV. Uehcnt of tbe Lake Supeilor Redon, HInn. Bot Stud., a, 1809, pp. 21.'>-Z7e. 

V. Uchena of the Uinneaota Valley and Southwestern Ulnnewla. Ulnn, Bot. Stud., 2, 181 
_ pp. 277-820. 

TI. licbens of Northwestern Minnesota. Minn. Bot. Stud., 2. 1001, pp. SS7-709, , 

VII. Lichens of the Northern Boundary. Minn. Bot. Stud., 2. 1003, pp. lS7-23e. \a p 

4 ^'d'^ 



26 MICHIGAN SURVEY, 1908. 

berry, SarBaparilla (dominant), and CUntonia borealU, bnt Aster 
mojctxyphyllua was not as abondant here a« elsewhere. Tbe larger 
Birches averaged about 12 to 15 inches in diameter. The general 
apjiearance of the conditions is shown in Fig. 39. This patch of birches 
was near the swamp (V, 5). From this Birch colony, on to the end 
of the trail to Siskowit Lakej tbe forest was dominated by large Birch, 
with a few quaking Aspen, Balsam and White Spmce, while in the 
damper places Ground Hemlock and Dogwood were abundant. On 
the ridges there are small "island^" ot Cladonia, mosses, Bearberry and 
a ground pine. The general appearance of this forest, in an open place, 
is sbown in Fig. 40. 

While there are thus minor differences which prevent absolute homo- 
geneity in the general conditions .of the forest, yet these differences do 
not seem to particularly influence the environment as a whole. The 
general transition from the openings, as found on rock ridges like those 
about camp {V, 3), to the Balsam-Bpnice forest may be seen by a com- 
parison of Figures 25, 26, 27 and 38. 

The fauna of the forest (V, 4) is rather varied. Tbe shells are repre- 
sented by Acantkinula harpa. Zonitoidea arborea, and PyramiAuUt 
cronkheitei anthonyi; the beetles by the carabid, Calathua, the fnngns- 
inhabiting beetles, Boletobius. Tritoma and Qrophaena, and doubtless 
many of the other species which were taken about the flowers and the 
comp. The wood-boring Hymenoptera, as UrGcenis. are also character- 
istic of this kind of forest. Hares remained concealed in the forests 
during the day, but at dusk they came in large numbers into the clear- 
ings to feed. The birds had begun to migrate when this location was 
examined, so that little attention was given to their habitat preferences. 

Station V, Substation 5. Tamarack Swamp. This swamp lies between 
Siskowit Lake and the western end of the trail through the Balsam- 
Spruce forest [V, 4). This is a valley swamp bordering a small stream 
which flows through the swamp. The central open part of the swamp 
Is occupied by a small pool or pond, Fig. 4h which is invaded by Yellow 
Waterlilies, Surrounding this is a zone of Buckbean and sedge, the 
overgrowing sedge being more conspicuous. This sedge zone is quite 
wet aud quaking. The current of the stream passed through this zone 
and parted the sedges in a wet line two or three inches wide. At its 
outer border, the zone becomes invaded by small Tamaracks, 4 to 5 
feet highj alders, willow, scattered Cassandra and Pitcher Plants, and 
Wild Rosemary (common). Eiiophonim, the Blue Flag and the Purple 
(.'inquefoil occur in some of the depressions. A very few small Arbor 
Vitae also occur here. A ntrip of trees bordered the stream, while 
farther south occurred the Cassandra aud Sphagnum zone proper. The 
latter contained scattered Blue Flags, and upon dry hummocks, colonies 
of Cladonia, which seemed rather out of place. The margin of this 
area was invaded by the Tamarack, Black Bpruoe and Labrador Tea. 
The geneva! ap]jearance of this forest is sbown in Figs. -}2 -}?. Near the 
margin of the swamp, where the spruces are quite large and the ground 
well shaded, the growth of Ijabrador Tea and Sphagnum was very Inx- 
uriant. The growth of Sphagnum at this place was by far the most 
luxuriant seen upon the island. It grew in billucks over fallen trees 
and stumps, and stood considerably above the general level of the swaniflJp 



ECOLOGY OF ISLE ROYALBL 27 

Thifl grouBd cover was not limited to the awamp, but invaded 
the Baisam-Spnice forest in large billow growths, such as is 
shown in Fig. H- In other places the undergrowth and ground 
cover of the BaJsam-Spruce forest apparently invaded the swamp, as 
was Been by the interminnling of the two plant Bocieties. Here 
thei-e is a mat of the Sphagnnm and Labrador Tea intermingled 
with Ground Cornel. CUntonia horealia and young Balsams. As 
in FUf. 4i.- tl'is might also be interpreted to mean an Invasion of the 
Balsam-Sprnce forest bv the swamp ; but the vigor and dominance of the 
Balsam society' favors the interpretation that this is an Invasion of the 
swamp by the Balsam society. It is not surprising that along snch a 
tension line either society may dominate at times. 

The fauna of this bog consisted of a Garter Snake (7*. sirtalit), found 
near the small brook flowing into the western or upper end of the bog. 
Here also was fonnd Hyla pickeringii, Rami clamitana, and R. 
cantabrigensia, and a water strider, Oerria, running on the surface. In 
tbe Cassandra and Sphagnum hammock zone were found the grasshop- 
pers, Mecostethus Uneatvs, Melanoplvs extremua, and Stenobothrus 
curtipennis. Nearer the central lilypond, among the sedges and Cas- 
sandra, were found the dragon flies, Tetregoneutia spinigera, Aeaohna, 
t/eucorlUnia hudaonica and Sympetrum obtrusum, and the spider Epcira 
patagiata. Through the central area of the bog the stream was' only 
indicated by the parting of the sedges, but at the lower or eastern 
end it again became well defined, and contained the small Stickle- 
back, Eucalia inconatans. Beetles taken ffom this bog were Haliplut 
ruficollia, Eydroptyrua triatia and Agabtis congener. The molluscs were 
represented by tbe small bivalves, Pi8i4ium. 

Station V, Subatation 6. South Shore of Siakoicit Lake. This sit- 
uation is simply the end of the trail through the forest, and marks the 
location of some collecting in the lake. The shore is rocky, with rather 
low and ovei^own banks. 

Station V, Subatation, 7. The Hat/town Trail. This trail begins al- 
most directly opposite the outlet of Siskowit lake, where a large White 
Pine has been marked "36 W, 65 X., 19 E." This area has been burneil 
over, but farther inland the blaze on the older trees enables one to follow 
the trail. The course is shown by Lane, ('98, PI, XI), but we examined 
it only to abont the point where it is crossed by the outlet of Hatchet 
lake, at which place there was a Tamarack swamp with very large trees 
(V, 8). After crossing the burned area near Siskowit lake, this trail 
passed through dense Arbor Vitae bogs and a large area of Balsam-Birch 
• forest. In general the area traversed was ratlier deeply covered with 
soil and contained very few rock exposures, those observed probably 
being due to Area. 

Tbe general character of the upland forest, of mixed conifer and 
hardwood, is indicated by the following list: Balsams. Birch and 
Aspens {all about 10 inches in diameter), scattered Arbor Vitae (10 to 
15 inches), a few scattered White Pines (about 3 feet in diameter), and 
a few Hard Maples (some S inches). It is thus seen that the largest 
abundant trees are the White Pine, Arbor Vitae, Birch, Hard Maple and 



byCoogle 



28 MICHIGAN SUHVEY, 1908. 

Aepen, The lar^e amount of hardwood present was an unexpected feature. 
This forest ma; be considered transitional betveen the Balsam, White 
Spruce forest, iind the Hard Maple, Yellow Birch, Balsam and Arbor 
Yitse foi"eat as found on the Desor Trail (HI, .'04). 

The undei^owth consisted of Ground Hemlock in abundance, Moan- 
tain Maple, Beaked Hazel, .young Balsams and Birches. The ground 
cover was composed of Clititonia borealis, Lycopodmm lecidulum, Cla- 
bonia on dry rotten wood, Aster macrophyllus, Large-flowering Rasp- 
berry and Wild Sarsaparilla. 

Mountain Maples and young Balsams sliowed a marked tendency to 
take possession of the trail. The lack of an undergrowth in the dense 
swamps was particularly noticeable, and the clearly defined old trail 
through such places indicated relatively stable conditions. 

A very marked characteristic of this trail was its limited fanna. Very 
few birds were seen, and Red Squirrels were not at all abundant. Sev- 
eral times we saw the remains of Balsam cones where a Bed Squirrel 
had taken a meal. Almost uo effort was made to collect invertebrates. 

Station V, Substation 8. Arbor Tilac Stoamp. This swamp marked 
the end of owr Haytown Trail, and bordered on a Bmall stream. The 
dark colored soil contained much humus and was soft and apongy. The 
vegetation bordering this stream was composed of Speckled Alder, Skunk 
Cabbage, Marsh Marigold, Clintonia borealis, scattered Bine Flags, 
and Ground Cornel ; in the moderate shade, CopUs trifolia, Mountain Ash, 
young Arbor Vitae and Balsams, Tivayblade, and, in the damp places, 
away from the stream, Mitella nuda. 

In the dense and apparently well drained swamp there was a firm 
humic soil covered by a thick layer of leaves, conifer needles and twigs. 
The ground cover was composed of Ground Cornel, Clintonia borealifi 
and Wild Sarsaparilla, with an undergrowth of Mountain Maple, Ciliated 
Honeysuckle, numerous young Balsams, young Arbor Vitae, Ground 
Hemlock, Mountain Ash, and Beaked Hazel. The large Tamaracks were 
about 3 feet in diameter, and the Arbor Vitae about 2 feet, others about 
20 inches in diameter were abnudant. It is thus seen that this was 
an old and mature swamp with some very large trees, under which the 
ground was quite open. With better drainage, the young or suppressed 
imdei^-owth would succeed the Tamarack society. 

The old trail through this swamp was remarkably well preserved and 
distinct because the dominance of the large shade-producing trees pre- 
vented the derelopment of an undergrowth. No animals were collected. 

Station V, Substation 9. Outlet of Siskowit Lake. A trail or path 
ran from the head of the outlet of Siskowit Lake south to the Siskowit 
Bay beach, a distance less than a quarter of a mile. The area traversed 
had l>een burned, and second growth had developed, the best of which 
was in the depressions where the soil is deeper. The open ridges near 
Siskowit Lake, where the soil is thin, have an open growth near the 
head of the trail and support Priums pennsylixinica. Birch, Dierville dier- 
ville, Mountain Asb, Mountain Maple, Amflanohier oUgocarpa, Ground 
Cornel, Everlasting, Wild Rose, White Clover, Fire Weed and mosses. 
In the depressions among the underbrush in the deeper soil and in shade 



3y Google 



ECOLXXSY OF ISLE ROYALE. 29 

is found Dif^villa diervilla. Fire WenA, Primus pcnmylvattica, Mountain 
Maple, Mountain Aah, Birch, Mountain Alder, Willow, Sarsaparilla, 
Wood Equisetum, Oak and I^adiee' Fern. Lai-ge-flowering Raspberry, 
Comua stolonifera and CUntonia borealis. 

The open arid character of much of this station is reflected in the 
grasshopper fauna, as follows : Hibbieaig tuierciiluttift, Melanoplus 
alaskanus, fasciatus and Circotcttix v^ruciiUitua. The butterfly, Basil- 
archia arthemia was also taken here. The rare dragonfly, Ophiogomphva 
eolumbrinus was also taken at this station, probably because of the 
proximity of the rapid flowing outlet, which forms a trout stream. 

Station V. Siitstation JO. Long Island Gull Rooko-y and Mctutgerie 
Talaml. This station was examined solely for its bird life, which will 
be discussed elsewhere in this report. This large Gull rookery is on Long 
Island, Fig. ^5, which lies about one mile west of Menagerie Island 
where Isle Royale Liglit is located. This bird clearly breeds upon the 
middle and upper baches. Long Island is formed by the upturned 
edges of red sandstone and is exposed to the full sweep of the lake 
waves, as is dearly evidenced by the bniv rocks. The vegetation on the 
island was not examined, as the time thiit could he devoted to the exam- 
ination of the rookery was limited. But mention should be made of the 
abundance of algae in the lake bordering the rookery, and of their 
abundance in the rock pools on the beach. 

Station V, Subitfation 11. TamaracU-Spntcc Stcamp. This station 
includes a waterlily pond surrounded by zones of sedge, heath shrubs, 
Tamarack and Black Spruce. A general view of the pond is shown in 
Pig. ^0'. It was located nlmost due wej^t of the western end of Station 
V, 2, and only a short distance northwest of the boat landing. 

In the pond, submerged, was Utricularia. Yellow Waterlily. 
Potamogeton. and Broaema (Water Shield) ; practically all the open 
water was occnpied by the Yellow Water Lily. At tlie edge of the 
water grow the sedges, Fig. Ift, which fonu a distinct zone, and the 
Backbean. The sedge zone also includes the Egiiisctvm. Purple Cinijue- 
foil. Comaruiii pahtftre, a willow. Miipirinnn. Water Hemlock (Cictita 
bulbifei-a), and the White Bog Orchid, Beyond the sedge zone comes 
Sphagnum, AiidrOmeda polifolia, Cassiindra, patches of Labrador Tea. 
Oxycoccits oxyvoceus (Small Cranberry, abundant), Alder, Chiogcne^ 
iiispidula {Creeping Snowberry), Bunch Berry or Dwarf Gomel iComus 
canadensis), Kalmut glauoa. Pitcher Plants, Drosera intermedia. The 
trees do not extend to the inner limit of the Cassandra zone. Cladonia 
grew upon dry hummocks in this zone. In a few places, in depressions in 
the tree zone, Eriophornm was found. The trees are Tamarack. Black 
Spruce, and small Arbor Vitae. The general appearance of the Cassan- 
dra and tree zones are shown in Fig. .'18. 

The invertebrate fauna of this station was abnndant and varied, but 
the vertebrates were more limited. In the Water Lily and Pota- 
mogeton zone the Stickleback, Fucalia inconatans, was taken, and at 
the sedge margin, Rawi clamitann. Water bugs arc represented by licl- 
ostoma and Corixa nymphs, and on the surface bv the Water Stridcr, 
Qerris rvfosciftrthtits. A small shell, Phyaa aplcvtoidca. was found in 



D,3-ZBcbyCOO<^Ie 



30 MICHIGAN SURVEY, 1908. 

Biuall foot-print like pools in the outer pai-t of tbe -Btickbean and sedge 
zone. In the (.Jassandi-a zone wei-e taken the dragontlies Enallagma ha- 
geni, Aeachna, Leucorhinia proxima, the grasshopper llelanoplua alas- 
kanus, and the butterfly Pyramaia cardtii. From this Im^ were also taken 
the Arachnids, Laciniua ohiocngc, Draaatia ncglectua and Pardoaa gla- 
cialis.^ 

The stations at Washington Harbor were not examined in 1905 in as 
mnch detail as were other localities, and the descriptions will therefore 
be correspondingly brief, and will be supplementary to a similar account 
by Ruthven, '06. pp. 48-52. 

Station 7, '0^, Cleoring at the Head of WaaMngtoii Harbor. This 
clearing is the property owned bj the Washington Club of Duluth. Minn. 
Mnch of it is sodded, pastured and under some form of cultivation. 
These conditions were particularly favorable for gi-asshoppers, which 
occur in great numbers. The following species were taken: Stcnobo- 
thrua curtipennia, Chloealtia abdominalia, C. conaperaa, Mecoatethus 
Uneatiis, CatiinuJa pelliicidxi, and Melanopiua alaakanus. This area ap- 
peared to be a favorable resort for migrating birdR, as shown in the ac- 
companying report by Peet. 

Station fl. 'Q-'f. Waahington Creek. This is the small titiut streaiu 
which flows into the head of Washington Harbor. 

Station III. '0.j. Trail aUyng the Top of (fte Orcriistone Range, the 
"Desor Ti-aiV This trail follows the road which has been opened from 
the Club House (I, '04) to Lake Desor (VII, '04). At the western end 
this road traverses a forest which varies considerably in its eom- 
position. In places it is dense and apparently oiiginnl. bnt at 
one place it has been bnraed and replaced by an abundant growth of 
Bircb. The original forest is dense and composed of large trees, and 
the proportion of hardwoods is surprising, since the Balsam-Spruce 
forest is so prevalent elsewhere upon the island. The hardwoods are 
really dominant. The forest Fig. -}.9, is composed of Yellow HIitIi, 
Balsam, Arbor Vitae, and a few Hugar Staples, and the undergrowth of 
Mountain Maple and Ground Hemlock. Farther out on the trail, toward 
I^ke Desor, the Maple becomes dominant and forms an almost pure 
stand, so dense that in places there is almost no undergi-owth, and tbe 
forest appears quite open with a scattered ground cover. A loose thick 
layer of leaves and twigs covers the forest floor. In the more open 
places the ground cover is composed of Large-flowering Raspberry. Wild 
Sarsaparilla, CUntonia ioreaiia, Lycopodium, mosses, Ground Cornel, and 
the shrubs. Mountain Ma]>le, lieaked Hazel, Round-leaved Cornel, Moun- 
tain Ash iiud Red Cheirv. The Yellow Birch is a large ti-ee. with 
a diameter of about - feet; White Pine is very rai-e, but the ti-ees are 
large, even about 3 feet in diaiueter; Arbor Vitae reaches about 2 feet. 
A few Large-toothed Aspens, Black Oak and Black Ash were seen, the 
Aspens about 20 inches in diameter and the Maples 10 to 15 inches. 

Red K(]nin-el8 were seen in the foiest, the body of a Lynx was fonnd 
banging on a tree where it had been left by a trapper, and several Toads 
were seen. Invertebrate life was abundant. In an Arbor Vitae stump, 
galleries of an ant^. Caniimnotua lierculeaiitis lohymperi (140 A), were 



DyGoogle 



ECOLOGY OF ISLE ROYALB. 31 

fouud ID both the Heaeoned and the decayed wood. A few beetles were 
takea alon^ the trail; Quediua ful(fideg, Tachinu* memnomiui and 
Qeotrupea blackburnii. t^helle were abnuduDt: Strobilopg virgo, Titrea 
hinneyana, Eticonuhia cherainua itolj/ffyratm, Zonitoidca arborca, Z. exi- 
gua. Pallifera dorsali'*, Pyramiduta alternala and I'. cronkJicitci 
anthonyi. 

Station IV, '04. Waahingtoii Brook, Cf. Huthven, '06, p. 50. This 
atation was examined on the elope back of the Club-houee. It is a 
swamp forest along the boi*der of a very small sti-ejuii. Fig. 50. Part of 
the forest ie being cleared. 

Station V, '04. Tamarack Swamp. This swamp was not Tieited in 
1905. Cf. Ruthven, M)0. p. 50. 

Station VI, 'O4. North Slope of Oreenstotie Range. Cf. Buthven, '06, 
p. 49. 

Station VII, '04. Lake Deaor. Cf. Ruthven, '06, p. 51. A few addi- 
tional records are: The dragonfly, Ktiollagma oxaulana, the water Btrider, 
Oerris remigis, the flah, Coregonus artedi. At the end of the trail (III, 
'Oi) at Deaor, tiie beetle, Melanotua paradoxua, and the spider Dolomedva 
idoneus, were taken. 

Station VIII, 'O4. Wcatern End of Siskowit Bay, The large clearing 
and bnrned area at the head of Siakowit Bay marka the site of a formei" 
town, the county seat. A well-defined graded road leada from near the 
north shore of the Bay westivard and north to au old mining camj). 
This road is being invaded in places by Birches and Aepens. This exten- 
sive clearing was overgrown with many introduced plants and wae 
given only a cursory examination. The following vertebratea were ob- 
served: Bharp-tailed Grouse, (of which several were seen), the Hare, 
Toad and Garter Snake. The snails, Polygyra albolabris and Pyramidula 
altemata, and the grasshopper Stenobothrus cwrtipcnnia were found 
here. The limits of this station were changed somewhat fromthoae 
given in 1904. 

Station IX, 'O4. Sottthiccatcrn End of Minong Trap Range. Cf. Ruth- 
ven, '06, p. 51-52, Xo additional collections were made here in 1905. 

Station X, 'O4. Washington Harbor. Cf. Ruthven, ,'00, p. 52. No 
additional collections were made at this station in 1905. 

III. TUB EVOLUTION- OF TUB UR088 ENVIRONMENT. 

1. Geological SiicceaaiOH. In bis i-eport on the Porcupine ^fountains. 
Batbven ('06) has summarily outlined the general geological history 
of the Lake Superior region. It is only necessary, therefore, for our 
purpose, to repeat some of this history and to entai-ge ufton thoao 
phases peculiar to Isle Royale. The structural geology of Isle Boyale 
has been studied in detail by Lane ('98) and is relatively simple. The 
different rock formations are in narrow strips nearly parallel with the 
long axis of the island, while the dip of the rocks is toward "the basin 
of Lake Superior. The rocks north of Siskowit Bay consist of the trun- 
cated beds of ancient lava flows, interrupted by a small amount of inter- 
bedded sedimentary rocks. Although these tilted and truncated beds are 



3y Google 



32 MICHIGAN aURVET. 1908. 

inclined at a higli angle, this vras not their original position ; in all 
probability they were formed in a nearly horizontal position by flsBure 
eruptions under the sea, because the lavas are interbedded with shales, 
sandstoneB and conglomerates. In thickness these lava beds vary 
from a few inches to hundreds of feet. The narrow beds often show 
upper and lower surfaces filled with small cavities (amygdules) in 
contrast with the denser central part. These cavities were formed by 
gas or vapor while tlie lava was hot, and leave such ai rock porouK 
and less resistant to disintegrating agencies and to erosion. The same 
principles also hold for the thicker beds of lava; the outer parts are 
more porous and softer than the central part. This structural difference 
is clearly shown in the topography of the island; the ridges mark the 
central or more resistant parts of the truncated lava beds, while the 
valleys, in general, hafe been worn into the softer outer parts of the 
lava and into the interbedded sedimentary rocks. These beds are of 
Keweenawan or pre-Cambrian age; their formation ceased with an ele- 
vation of the land from the sea and their destruction was begun by 
the agents of sobaerial erosion. These processes continued until the 
titled strata were truncated and reduced to a base level. Again the 
region was depressed and upon this eroded surface were deposited un- 
conformably those red sandntoues and congloniei-ates which now char- 
acterize the Siskowit Bay region and to the southward, and are of 
Cambrian age. Once more the region was elevated, titled and subjected 
to prolonged erosion and the strata truncated as had been done with 
the Keweenawan. Similar processes continued until the marked eleva- 
tion of the land, which took place at the close of the Tertiary, and 
which initiated the repeated glaciations of the Ice Age. 

With the extension of the last or Wisconsin ice sheet in the Superior 
basin, Isle Koyale was completely overridden by the movement of an 
ice sheet from the northeast that moved almost parallel to the ridges, 
but was somewhat more inclined from the east (Lane '98, p. 183). For 
this reason there was a tendency to plane down the 'southeastern slopes 
and to presence the steeper ones which had been formed on the north- 
western side (Foster & Whitney, '50, \>. 202). As the island has a 
topography which indicates snbaerial rather than marine erosion, it 
must have had at one time a residiml soil, which, unless it had been 
swept away by a former ice invasion or the waves of some body of water, 
was probably removed at this time with the minor inequalities of the 
surface. In this manner the Superior lobe burled the island under 
several thousand feet of ice and continued its movement far to the 
southward, leaving a glacial desert in its wake. This condition of affairs 
lasted until the return movement broke up the great ice sheet into lake 
basin lobes and brought the receding ice front into the Superior basin. 
As soon, however, as this lobe wasted away from the margin of this 
basic, the water from the melting ice accumulated before it and formed 
a lake which, overflowing the rim, found its way through the St. Croix 
valley to the Misnissippi river, as indicated in Fig. ol. But, as the ice 



3y Google 



ECOLOGY OF ISLE ROYALE. 



wall continued to reti-eat toward the northeast, these ponded waters 
increased in area and formed the highest beach lines now preaerved on 
the north shore of Lake Superior. The evidence for this is fonnd on 
Mt. Josephine, located on this shore just north of the southwestern end 
of Isle Royale, which reaches an elevation of about 800 feet above the 
Lake. Far up em its slopes, according to Lawson ('93, p. 253), evidence 
of beach lines are fonnd, about which he says : "These two terraces at 
585 and 607 feet are remarkable for being the highest strand lines which 




D Ihe Ice sheet and theic Mlu- 

have thuB far been observed on the coast of Lake Superior." Under such 
conditions it seems that Isle Royale would undoubtedly have been sub- 
Tnerped. The ice retreat continued and finally Isle Royale, freed from 
the ice and in part from the lake waters, emerp^ed as a narrow rocky 
ridfi^e — the crest of the Oreenstone Range which today rises, at the 
northeastern end of the island, to a maximum elevation of about 550 
feet. As the ice wall retreated the Michigan and Huron basins became 
confluent, and an outlet to the east (Trent valley, Ontario) nt first, 
later the Port Huron and possibly the Chicago outlet ((ioldthwait, '09, 
p. 65) became available. Fig. 53, and at almut this time the (ilacial (Jreat 



34 



MICHIQAN SURVEY. 1908. 



Ltikea became iBolated from the Missiesippi drainage. The lake level 
wna lowered, and it was perhaps at this level that the beach lines were 
formed on the north side of I^ate Superior, which are now 400-500 feet 
above the present lake level (Taylor, '97, p. 126)! Similar evidences of 
ancient beaches have been recognized by Lane ('98, pp. 188-191) upcm 
Isle Eoyale, but he is inclined to place the level of this Glacial Laka 
Algonquin at about 485 feet. It is probable that more field 
work will be necessary before adequate correlations of these beaches can, 
be made. 

Seine general idea of the extent of the island at this stage mai 
gained by reference to the 160 foot contour on the accompanying mai>, 




Fig. 53. At this time, Fig. 5Z. the ice sheets had retreated far enough 
the northeast that the climate of the Superior basin must have be«i t 
greatly ameliorated that animal life could have lived in its water. Thit 
infei-ence seems probable because fossil shells have been found in the beacl^ 
line's of the same lake farther to the south by Lane and Walker (Laiw-° 
'00. pp. 248-252) , and at Port Huron, Michigan by the writer in conipauft 
with Dr. J. W. Goldthwait and Dr. A. G. Huth'ven (Goldthwait, '07, i ' 
118). Here wei-e found an abundaJice of Qomohasis Uvcscens, octtf 
sional valves of tipluierinm xti-iathnum Lam. and Unionid fragments 
fauna like that of the present beaches. It is therefore not improbahlig 
that this fauna invaded the Great Lakes drainage from the MississipiU 
during the early stages of the great glacial lakes, when they still ove^ 
flowed into the Mississippi drainage. ^ 



ECOLOGY OF ISLE ROYALE. 

. invaaioD and the Boura 



i.^oAS.-ij^-' ■ i.--^^ 



4''- 



-Sife. 



m^^ 



34 

Luk. 
waa 
forr 
abo" 
one 
l8U 
Alg 
vox 
be 

e 

gai 



MICHIGAN aURVEY, 
.ted f rom the UiBeiBBippi draitM 






II 



EcoLoay OF isle royale. as 

The time of invasion and the sources of the Glacial and post-Glacial 
sapply of life which invaded the northern land and waters presentB 
several interesting problems. At this point the origin of the aquatic 
biota la of special interest. From what is known of the fresh-water 
ac|uatic biota of the far north, it is very probable that the Glacial Great 
Lakes, at least in their later stages, were not utterly devoid of animal 
life. So it seems reasonable, to infer that such forms as lived daring 
Glacial times near the ice margin were among the first to succeed the 
retreating ice sheet. 

As the lake fauna is one of the most characteristic features of the 
life of northeastern North America, the question naturally arises as to 
where this fauna was preserved during the Ice Age. Today there is 
no extensive development of lake fauna south of the glaciated region. 
Where then was such a fauna preserved? On the west were the arid 
plains, and to the south au old land area of mature drainage aud very 
few lakes. Such relations as these suggest that this fauna must have 
occupied the lakes formed on the rejuvenated glacial topography or 
in the adjacent streams. Had there been extensive lake areas to the south 
to act as regions of preservation, it seems probable that the present fauna 
of the Great Lakes would have been much richer. Undoubtedly 
the most important fresh-water preserve was the Mississippi system, 
on account of its direct communication with the glacial drainage, thus 
allowing a southward escape into more favorable climatic conditions. 
This was also a water communication of considerable duration, lasting 
even into early post-Glacial times, and one which has greatly influenced 
the origin of the present fauna of the Great Lakes. In all probability ■ 
it was this Glacial and post-Glacial water connection and barrier 
that retarded the northern extension of so many land species, and at the 
same time favored the extension of certain aquatic animals. The later de- 
velopment of the eastern outlets did not open up snch a favorable 
source of supply as occurred farther west. 

So much for this phase of the problem. Now let us continue with 
the history of the Superior basin since Algonquin times. The fall in 
the lake level did not take place suddenly, since a series of bench 
lines are preserved which show that it halted for some time at different 
levels, but none were of any remarkable duration until it reached a 
level marked by a very extensive aeries of beach lines now preserved 
at about 60 feet above the level of r>ake Superior. This well defined 
beach represents the shore of the Nipissing Gi-ent T^akes, poat-CJIacial 
lakes whose general outline was much like that of the present lakes 
in the same l>asins, as is indicated in Fitj. '>.i. The low outlet of this 
Jake was to the east through the Ottawa, valley into the Champlain 
Sea, and is of special interest in that it is suggestive of how certain 
Great Lake animals of marine affinities (Mi/six, Pontoponna, Trifflopsis) 
might have invaded the upper lakes in post-Glacial times. At one 
time it was thought that there had been a Glacial salt water com- 
niuuication between Ijike Superior and the Hudson Bay region, but 
this view has been abi)ndoned (cf. Taylor, '97, pp. 127-12S; *!)G, pp. '2'i'>- 
256. and Coleman, '06. pp. li):!. IDS-lilil). It is definitely known that 
the land was depressed to the north of Lake Superior, but this period 
of depression was at a time when it was covered by the ice shMh. 



MICHIGAN SURVEY. 1908. 




"^ 



Dis !,= i„Cooglc 



ECOLOGY OP 18LE ROYALE. 37 

and thus the salt water was excluded. An altei'native hypothesis is 
that these animals are adapted to a constant and low temperature 
rather than to fresh or salt water, and that during Glacial times they 
were dispersed far to the south in fresh water and have only been 
preserved in restricted favorable localities. The low temperature of 
Glacial" times would be a period especially favorable for the acclimatiza- 
tion of marine forms to fresh water on arfount of the favorable con- 
ditions which accompany the slow rate of changes at low temperatures. 

The long duration of the Jfipissing (Sreat Lakes is well attested by 
the character of the beach. As Taylor ('90, p. 398) remarks: "It is 
altogether the most remarkable littoral feature of the Great Lake 
region. It is a shore line well advanced towards old age. All other 

beaches of the lakes are youthful in comparison Instead 

of the slender spits and barrier bars of the Algonquin and other beaches, 
the Xipissing beach has what may be called barrier plains, made up 
of many, sometimes forty or fifty, massive beach ridges laid one against 
the other. Many bays were entirely filled by these beach plains and 
others were cut off, so as to form small littoral lakes. Some of these 
plains are a mile to a mile and a half wide. In some instances the old 
deltas of other beaches are large and conspicuous, but the constmctive 
products of wave action have no comparison to those of the Nipissing 
beach." From a biological standpoint these facts are of special signifi- 
cance. The maturity of the beach line is a condition decidedly favor- 
able to the development of a littoral biota. The sandy shore, spits, 
bars, beach pools, cut-off ponds and lakes furnish a variety of favorable 
habitats in marked contrast with the poverty stricken character of 
life frequenting an exposed and topographically youthful lake shon*. 
Such an old beach is both qualitatively and quantitatively favorable 
to the biota, and not only favors an abundant supply but also its dis- 
persion along shore and by currents throughout such a body of water. 
The long daration of such conditions is of evident advantage to an 
extensive dispersal of such life. 

As the basin of the Nipissing Great Lakes In the Superior basin was 
so much like that of Lake Superior, it is not improbable that the lake 
currents were much the same in both lakes, so that our knowledge of 
the present lake currents should aid in the interpretation of those of 
the Nipissing Great Lakes. Such relations as these suggest that at the 
Nipissing stage, and perhaps even earlier, the lake- currents tended to 
people Isle Boyale with north shore drift. By this time the island 
was quite large, though smaller than the present island by the subtraction 
of the area below the 60-foot contour. At this time the climate of the 
region must have become greatly ameliorated so that the north shore 
of Lake Superior was perhaps repopulated from the south, largely 
around the western end of the lake. With the advent of an abundance 
and diversity of plant and animal life, a new element enters the environ- 
ment, whose influence is far reaching. The vegetation tends to blanket 
the surface with a humus layer and thus to bind the soil so that it 
retards erosion and becomes a geological agent. The influence of 
animal life is also far reaching and may be couspieuous if beavers are 
abundant. But these influences will only be mentioned here. 

The development of the Nipissing beach upon Isle Boyale hac 



"".^R?' 



38 MICHIGAN SURVEY, 1908. 

been eo clearly recognized as elsewhere. Thus Lane ('98, p. 187) con- 
Biders the present beach as the most distinct of any found npon the 
island. In a way this is not surprisinG; when we recall the fact that 
at former lake levels the small area of the island did not permit of 
an extensive stream development, hence the limited quantities of sand, 
gravel and bonldera. Thus the overriding of the ice, the isolation of 
the island in deep water, and the steep shores of resistent rocks are 
<'onditions unfavorable for supplying tools with which the waves could 
work. All of these conditions would tend to preserve the youthful 
topographic features and exaggerate the apparent relative rate at which 
the island emerged from the waves and the small time during which 
the waves beat at any particular level. The materials available to the 
present waves have therefore been cumulative. Lane ('98, pp. 188-189) 
has recognized several evidences of a fiO foot level. 

After the formation of the Nipissing beach there was an uplift towartl 
the north, as shown by Taylor's ('97, p. 127) study of this beach on 
the Canadian shore north of Isle RoyaJe. In the vicinity of Port Arthur 
this beach is at 60 feet; at Nipigon !)0 feet, and 110 to 115 at Peninsula 
Harbor, Such an assumed variation or tilting near Isle Royale sug- 
gests the necessity of great caution in attempting to correlate thR 
various beaches and emphasizes the desirability of further field work 
upon this subject. Lane ('98, p. 192) suggests that this northward tilt- 
ing has tended to pond the northeastward flowing streams and to drain 
the ones flowing in the opposite direction. Such tilting as this would 
have considerable influence upon the biota. Even in an uplift of a few 
feet per mile, in the case of Isle Royale 45 miles long, would be sufficient 
to have a marked influence upon the swamp environment, which is one 
of the most characteristic features of the island. In this manner ii 
swamp and its biota might migrate several miles, become a pond or 
'ake or even become drained, and other fates are suggested for ponds, 
lakes and other environments when such a distinct trend or dynamic 
tendency is present in a given region. 

The change from the Nipissing to the present lake level was not ii 
sudden one, as Lane ('98, p. 191) has recognized beaches at various 
levels showing its gradual character: the 30 and 15 foot levels are, 
however, the most distinct. A few observations were made ui)on two 
of these abandoned beaches, but their height was not determined. One 
was located just south of the mouth of Conglomerate Bay in a small 
cove about CO or 70 feet wide. There was an abundance of fresh drift 
wood a few feet from the edge of the water, back of this a zone of 
weathered and decayed drift, and beyond this a high boulder beach 
containing disintegrated boulders with foliaceous lichens, while back of 
the lichen zone came ^^'ild <:horry, I'aper Birch, Bear-berry, Wild Rose, 
Jack Pine, Alders and Columbine. The back sloj)e then declined into 
a Jack Pine growth. This beach is interestiug because it illustrates 
the various stages from wave-waslied, clean sand and gravel back into 
the forest growth. Lane ("98, p. 185) refers to a lichen covered beach 
on Sec. 10, T. (i5, R. :!4. The secniid of the benches mentioned is located on 
the south shoi-e near the eastern end of Siskowit Bay (Sec. 26, T. G5 >'., 
E. 35 W.). The present beach is locally known as the "Greenstone 
beach" and forms a good boat landing. 

i.,C0CH^Ic 



ECOLOGY OF ISLB ROTALB. 39 

2. The Topography and its Origin. The raoat oonBpicuous and char- 
acteristic topographic features of the island are its parallel flat-topped 
rock ridgea with the intervening valleya and numerous enampfi. These 
ridges project far out from the main body of the island and form the 
narrow rock ridges bounding the harbors, and forming a vast number 
of small islands and low rocky reefs. The tilting, faulting and trunca*- 
tion of these narrow beds clearly shows that the dependence of the 
topography upon rock structure is one of the most characteristic features 
of the island. 

The main ridge, the Greenstone Range, is a divide which extends the 
entire length of the island, and is from about 400 to 500 feet high, 
with a maximum height of about 550 feet at the northeastern end of 
the island. At only one place does a drainage line cross the Green- 
stone. This is a small stream heading in Sec. 17, T. 64 N., B. 37 \V. 
and a tributary to Washington Eiver. This ridge is a truncated lava 
bed whose outer softer part has been eroded, thus throwing into promi- 
nence the compact resistant central core. Thus erosion, faulting and 
the dip of the rocks have combined to produce a northwestward facing 
escarpment nearly throughout its extent. The fairly Sat topped 
truncated ridges of the island clearly show that their origin must be 
due to a period of baseleveling and is no doubt related to those exten- 
sive processes which have produced the Laurentian peneplain (cf. 
Bnthven, '06, p. 45) of the Superior region. The ridge of second im- 
portance is the Minong Trap Range, which lies parallel with the Green- 
stone, about a mile to the northwest, and reaches n height of about 400 
feet. Between these ranges lies a valley containing five fairly large 
lakes, all of which drain across this range to the northward, and tbc 
probable faults indicated by Lane ('98, pl.'l) at Todd Harbor and 
McCargoe Cove are su^estive as to how the ridge has been broken 
through. Faults seem to have influenced the location of several lakes, 
such as Angleworm, Lesage, Livermore, Chickenbone, Feldtmann and 
also the outlet of I^ke Hichie into Chippewa Harbor. In addition to 
these main ranges there are great numbers of lower ones whose heights 
range from 100 to about 300 feet. East of Lake Feldtmann there is a 
bold escarpment 130 feet high, which was said by Mclntyre (Foster, 
'50, p. 506) to afford the "finest view that I have seen on the island." 

The drainage of the island presents some interesting features. At 
each end of the island the drainage is mainly along the valleys into 
the harbors at their ends. Between these two extremes, roughly marked 
by the area between lakes Desor and Sargent, the drainage, although it 
may follow the vallej's for some distance, is yet to a marked degree 
across the strata or ridges. Taken as a whole the drainage is very 
imperfectly developed. Although the island is not extensive, it con- 
talus numerous small independent streams which drain into the lakes 
or directly into Lake Superior, but it has no master stream. It seems 
probable that this is also related to faulting, as also in the case of 
the stream, which may be called Malone Creek, that flows into the head 
of Si^owit Bay. The probable influence of faulting upon the location 
of lakes has previously been mentioned, and combined with its influence 
upon streams reinforces the idea of the dominance of structure ujion 
the topography and consequently upon the drainage. But when in the 



40 MICHIGAN SURVEY, 1908. 

field the tno^t conapicuous features of this imperfect drainage are the 
vast strips of swamp land found in the valleys and bordering the lakes 
and streams. The rock bound character of the banins and the south- 
ward tilting of the surface must greatly influence the form and extent 
of these strips. The stream channels have not cut deeply but are lai^ely 
bordered by swamps, and tbe 'divides between many of them are very 
low or inar even be swamps, so that the drainage from either end of 
a swamp niay be into a different drainage line. Such imperfection of 
the drainage means that evaporation rather than mn off is the pro- 
nounced feature, and this condition, combined with the insular loca- 
tion, must greatly influence the relative hnmidity of the atmosphere. 
The brownish waters of even the largest lake upon the island, Siskowit, 
54 feet above Lake Superior, clearly shows the influence of the imper- 
fect drainage and the extensive swamps of its drainage basin. 

The general character of the soil was indicated by Ives on the Linear 
Survey map. This is as a rule shallow, the deeper being at the south- 
western end (T. 64 N., El 38 W.) and is characterized ae "sandy loam 
and stony, second rate sufficiently deep for cultivation." At the head 
of Siakowit Bay (T. 63 N., R. 37 W.) he records soil "stony, 2nd and 
3rd rate land. Soil varies from a few inches to 3 or 4 feet in depth," 
And near McCargoe Cove (T. 66 N., B. 35 W.) the soil is from 1 to 10 
feet deep. The soil then in general may be said to be shallow, second 
and third rate stony, sandy loam. In the swamps and valleys there 
is a large amount of vef^table debris, although it is prolrable that this 
is generally not deep. S'o bog lime or marl has been observed. Lai^e 
strips of the ridges are destitute of soil, especially those which have 
been bamed. No morainic materials were recognized, although the ice 
overrode the island, and glacial boulders are abundant in places, as 
about the head of Washington Harbor. Dr. Lane writes me that there 
is some till, "especially on the lee end near Washingtoa Harbor." 

The origin of these soils appears to be relatively clear as there 
are only a few possibilities available. Some of the pre-Glacial residual 
soil may have been preserved but it has not been recognized. As above 
mentioned there are some Qlacial boulders and till. The post-Glacial 
disintegration and decay of the rocks has been the most important 
source, supplemented by organic remains, from the vegetation in par- 
ticular. A fourth source is the lake deposits of sand and clay as the 
waves have worked over the entire surface. These are best preserved 
in what were once harbors or places protected from the waves. In 
many localities the origin of the soil is diverse, several different pro- 
cesses having contributed a part. 

From the above topographic relations it is seen that the flat-topped 
ridges and depressions are due to the strncture of the rock, the influence 
of base leveling processes and probably also to faulting. The present 
drainage is not sufficient to explain the primary ridges and valleys; 
these must therefore have been inherited from past conditions. The 
present drainage is therefore consequent and in its infancy, hence 
its im]>erfection. From a biological standpoint these facts are signifi- 
cant because such conditions favor isolation of small streams, swamp 
and lake habitats affect the relative humidity and produce a prominent 
zonal and linear arrangement of the habitats along the ridges and 



ECOLOGY OF ISLE ROYALE. 



valleys. The absence, residual, or organic character of tlie soil is also 
an important factor of the environment. 

3. The Atnwspkeric Influences an4 their Evolution, a. Climate. 
Unfortunately there has been no continuous series of eliniatological 
records made on Isle Royale. A few records were made by our party with 
instruments loaned by Mr. C. F. Schneider of the Michigan Weather 
Service, that, while very imperfect, are su^estive. The mean tempera- 
ture for 26 (lays in July is 58° F., the minimum record is 46", and 
the maximum 79°. From August 2 to 17 the mean is 59°; the mean 
maximum is 71° and the mean minimum is 47°. For the same period 
the maximum is 80° and the minimum 30°. There was but little rain 
although it rained all day on July 15. 

Very fortunately, however, these meager records may be ■ supple- 
mented by those from Port Arthur, about 25 miles distant on the 
Canadian shore.' This data has been kindly furnished by Mr. B. C. 
Webber of the Canadian Meteorological Service. The records cover the 
decade of 1896 to 1905, and show the mean monthly and annual tem- 
peratures, maximum and minimum temperatures, and the precipitation 
for the same period. 

The table of temperature. Table 1, shows that the decade average of 
the mean monthly temperatures for February is 7.65° P., with a maxi- 
mum during July of 62.24° and an average annual of 36.07°, The 
monthly averages of the maximum temperatures for January is 38.1° 
and for July 85.8°, with an average annual of 36.7°. The lowest 
average monthly temperature for the same period is — 27.5° for January, 
and for July 42.0°. The average minimum temperature for this ten 
years is — 30.8°. Hie monthly averages for 5 months are below zero. 



TABLE NO. 1— MEAN. M0NTH1.Y AND ANNUAL TKMPERATrREa AND AVERAGES 

FOE ID YEARS. FORT ARTHUR, 1896-1905. 

Mean Temperaturei In °F. 



V-,. 


,.. 


iWi. 


.„. 


April 


Iby. 


Jm.,. 


July. 


A* 


S^ 


Oct 


Not. 


Dk. 


i£ 




ii 


ill 

l'.S 


16 ;b 


35 .9 

II 


r, 

4T.9 

Si 

40.2 
46,7 

45.8 


si 




Si 

ai 

|; 
61 : 


in 

HI 
30:2 

S4:6 


38.S 

«:3 
40:3 


19.7 
307 

26:o 


IS.O 
14:3 

n.7 


,1 




























ISTO 




M.0 


Si S;! 
















TBlgl 


7.» 


7,66 


19 .S4 


36.19 


- 


65 82 


».« 


«,1. 


W.S 


42.58 


n.iB 


13,11 








e.E.S(khif»(MaHi)... 


Ml 


>.., 


30.8 


46.9 


„., 


67.1 


„.. 


69.1 


63.6 


»,6 


36.5 


27.0 


p.t 



' For B general eccounl o[ the Cumidtan cUinate see Stupait '08 and '05, 



jiGoogle 



MICHIGAN SURVEY, 1908. 



Highest Tempenitur 



Y»r. 


Jan. 


Feb. 


Mir. 


.\pra. 


May. 


JUM. 


My. 


A* 


Sn>t. 


'ott. 


\».. 


Dh. 


A>>D.[ 




3».0 
42.0 

3s:o 

teo 


47.0 

42:0 
MO 

38.0 
S3:o 


47 

310 

44.0 
4B0 

41:0 


G»0 

si 

87.0 
8».0 

is 


3B.0 

is 

77.0 

Is 


Bs:a 

80,0 
81.0 


8410 

8fi.a 
86 :a 
in.o 

11 


81 :a 

8J.0 

M.O 
80.0 

is 


77.0 

11 

S2.D 

il 


M.O 

08.0 

TOO 
83:0 


47:0 
81.0 
00.0 

ISO 

69:0 


40 :o 

37,0 
34:0 






r, 


ffi:::::;:::::::::;:::; 




86.0 
WO 






IS;::::.:::. :::.;;:;;;■ 


84,0 






A. r 


is.i 


33.8 


4S,- 


eg.e 


™,, 


82,, 


»,. 


82.3 


„,, 


.,.. 


.., 


... 

















n 'F 














Year- 1 J.n. 1 F.b. 


UDr. 


April. 


May. 


Jun*. i July. 


Aug. 


Sepl. 


Ort, 1 .Nor. 


I^-'t^^ 






-18.0 
-IS.O 

-17:0 
-2fl,0 

-10 :o 


8.0 
8:0 
16.0 

10,0 


31.0 

26.0 

27 :o 

2t.O 
29.0 

24:0 


38,0 

■s.o 

33,0 
3S.0 


41.0 

43:0 

42.0 

40,0 
40,0 
42.0 


34,0 

40 
41.0 

40.0 

3B,a 

I:! 


230 
2S0 

11 


,..'..» 


|i 

-2s:o 
-20:0 

-28.0 


li 


ijg... .. , , . ... 


-IS.D -2S.a 

:l!:8 31 


200 

20 :o 


-6,0 
-8.0 


I8SB 








-30.0 

-34:0 
-21,0 


:g;! 

-26,0 
-24.0 


"^■|f 






-34.0 










-W.J 


-28.4 


-14.S 


1 ?«« 


34,S 


„. 


„,. 


». 


... 


-8.8 


... 


-30.8 









The precipitation during the same period is shown in Table 2. The 
minimum average monthly i-ninfall for the period is .002 inches for 
February, wilh a maximum of 4.25 inches in July, and an annual 
total of 21.7S inches, more tlian half of which fell during the growing 
season for the vegetation — June, July and August. The snowfall 
averaged a maximum for January with 4.59 inches and an annual total 
of 25.44 inches. The deep snows of this region are thus seen not to 
be due so much to the abundant precipitation as to its pi-eservation by 
the low temperature. 



3y Google 



ECOLOGY OP ISLE ROT ALE. 
TABLE NO. Z.— PORT ARTHUR, ISM-IMie. 



Y«w. 


,„. 


F*. 


... 


A(«iL 


■>>. 


Juu. 


July. 


Auf. 


Btpt. 


Oct. 1 Kov, 


n*. 


ToUL 




O.ST 

!:1S 

o.m 

0.00 

O.DO 
0.00 

8:S 


si 

o.oo 
o.oo 


Id. 

o'oo 

0.93 
0.00 
O.DO 

O.M 
O.M 


In. 
2. 32 

0.07 

o'so 

SI 

0.00 


S 

0.38 
006 

i:37 


In. 

e.ft4 

248 

in 

if 


in. 

T.aa 


il 

203 

2:«i 


In. 

il. 

il 


S20 

11 


In. 
I.2» 

11 

o.« 
i.ai 

o:is 


i 
if 


In. 










































A'-™. 


,0S7 


.00* 


.n 


.BS 


..» 


IN 


'" 


3.12 


... 


... 


» 


.» 


„,» 


S. ESUchku 


IM 


.... 


..« 


.„ 


,„ 


3... 




i.K 


... 


,.» 


=,» 


2.03 


», 



Y(«r. 1 J«n. 

1 


Feb. 


Mur. 


April 


Hny. 


'""!* 


Am. 


Sept. 


Ort. 


Kov. 


n«. 


ToUl. 




iB. 


In. 

11 

!:! 
II 


In. 

s.t 

li 


In. 


In. 


In. i In. 


In. 


In. 


In. 


In. 
1:2 


In. 
.8 

1 

3.5 
137 


in. 




























• 






























3.3 
























11 


















0.3 










1.0 




























O.T 


















^ 


'- 


'" 


S.S7 


7.« 


0.3 










.93 


4.2s 


.,., 


M.« 














1 




















38. i 














1 









These climatic records are likely to mean little when taken by them- 
selves, but when compared with ihe conditions found in the other extreme 
of the Htate, interesting relations become apparent. Tranwau {'do b, 
pp. 3^li-3n8) has snmmarized the temperatare and precipitation nienns 
for certain localities in southeastern Michigan, and these means have 
been placed in the table with the Port Arthur data. The most striking 
difference (Table 1) is the much higher temperature throughout the 
year in southern Michigan; the mean July maxiQimn is 71.9° as coiitn>sf- 
ed with 62.24° at Port Arthur; the annual mean is 47.2° as contrasted 
with 30.07° for Port Arthur. The northern mean is l>etw('en the tem- 
perature of tlie maximum density of water (39.2°) and tlie freezing point. 
The precipitation presents almost equally striking differences. The rain- 
fall instead of being largely confined to the summer months, as at Port 



44 MICHIGAN SURVEY, 190g. 

Arthur, is mucli more generally dietributed throughout the year. The 
rainfall is also about % more in the south, the northern mean is 21.73 
inches and the aoutheru one 30.22 inches. In the north there is about 
1/5 more snow than rainfall, 21.72 as contrasted with 23.i4 inches; 
while iu the south about 14 more of the precipitation occurs as Bnow„ 
30.22 of rain as contrasted with 38.4 inches of snow. While in both 
regions the greater precipitation is in the form of snow, the longer grow- 
ing season of the plants in the south makes more of this moisture avail- 
able ; but on the other hand, on account of the higher temperature, more 
is needed. While about one-balf of the rainfall in both regions occurs 
during the growing seasou, ,iet the evaporation is much greater in the 
south so that the relative humidity is less when compared with the 
north. (Cf. Transeau, '05, a). It seems probable that the relative- 
humidity of Isle Bovate is greater than on the adjacent mainland on ac- 
count of its insular location and imperfect drainage. 

Mention should also be made of the long period of daylight in the 
north because this is of great importance to a vegetation whose period 
of growth is limited to such a short summer. 

To one accustomed to the hot summers farther south, the cool summer 
of Isle Royale is very agreeable and invigorating. Moderately heavy 
clothing is needed for comfort except during the middle of the day wheo 
the heat at times is very oppressive. This was especially the case dar- 
ing our examinations of the rock ridges. Thus on July 10 on the Jack 
Pine Hidge (III, 5} the thermometer on a mat of Cladonia recorded 
93° F. in the sun, while at the same time (2 P. M.) in the sun, but ex- 
posed to a cool breeze, it recorded 76° F. Such temperatures wonid not 
attract special attention were it not for the fact that usually the tem- 
perature is 80 much lower. The nights are very cool, and at Washiugton 
Harbor on Aug. 22 there was a frost in the valley along Washington 
Creek (II, '04). During our camp at the Light-house, when shore winds 
accompanied a storm, the temperature became so low that a fire in the 
evening was necessary for comfort. On July 15 there was a brisk east 
wind, with a mean temperature of about 50° so that the vapor of ones 
breath was visible all day. The lake breeze is at times very noticeable 
as one passes from Bock Harbor into the channel at Middle Islands. It 
is quite probable, as Jackson ('50, p. 420) suggests, that this cold lake 
air is a factor in the production of the stunted tree growth. 

The low temperature of the wet, densely foreate<i cedar swamps is 
^•orthy of special mention. As Foster remarks ( '50, p. 420) "Und» the 
shade of the crags, and among the thick evergreen swamps of white 
cedar, it not nnfrequently happens that perennial ice is found, covered 
by a layer of turf. Mr. Blake discovered a considerable area of ice thus 
preserved in midsummer, near Rock Harbor." Unfortunately our party 
did not find such conditions although such "cold islands" were kept 
in mind with the idea that under such conditions "glacial relicts" might 
be expected if these areas were of sufficient extent. 

ft. ScicliC8. The rapid and temporary changes of. the water level in the 
harbors has been the basis of much comment. This was very marked at 
Tobin Harbor and at Washington Harbor. Its influence upon Washing- 
ton Greek was quite marked, at times it would be ponded for some 
distance up streaui while on other days it would be a briskly flowin^r 



ECOLOGY OF ISLE ROYALE. 46 

-stream. Foster and Whitney ('60, p. 51) make the following comment 
-upon these fluctuations at Bock Harbor; "While at Rock Harbor, Isle 
Royale, in the summer of 1847, we witnessed the ebbing and dowing of 
the water, recurring at intervals of fltteeo or twenty minntes, durinp 
"the entire afternoon. The variation was from twelve to eighteen inches: 
and we took advantage of their recession to catch some of the small 
lake dsh which were left in the pools. The day was calm and clear 
bnt before the expiration of forty-eight hours a violent gale set in." 

This phenomenon has been investigated on the Great Lakes by Deni- 
■Bon ("98, p. 568) who states that these seiche movements are very 
marked preceding and during storms and are due to atmospheric pres- 
«Qre upon the lake. 

c. Climatie Succeaaion. From what is known of the general geologi- 
cal history of the Superior region, during Glacial and post-Glacia) times, 
it is evident that there has been a great climatic change which has been 
of the utmost biological importance. It is therefore desirable to see 
■what inferences will aid us in forming a general conception of the pos- 
sible climatic successions. It appears to be generally conceded that at tlie 
margin of the ice sheet the conditions mnst have been quite arctic in 
character, similar to that of the "barren grounds" of the far north. Such 
climatic conditions might result from a permanent atmospheric low cor- 
related with the presence of the ice she^ (Cf. Chamberlin and Salisbury, 
'06, 11. pp. 674 675 ; 111, p. 433) . The prevailing westerlies, combined with 
a permanent low to the north would favor westerly continental winds 
along the margin of the ice. Perhaps a suggestive comparison can be 
made between the seasonal transitions from the two permanent winter 
lows near the Arctic regions, into the summer condition of one low witli 
Its transitional "March weather" and that of American and European 
jjlacial lows and their transformation into the present summer arctic 
low. In connection with this subject a paper by Fassig ('99) is of spec 
ial interest. Analogies are often dangerous but the idea is of interest 
Twcause it suggests a "March weather" transformation for post-Glacial 
times. In this connection the formation and occurrence of the wind 
blown loess, with its greatest development in the west and on the east 
banks of certain streams, is of special interest, although these condi- 
tions did not develop in the north as they did farther south. The oc- 
-cnrrence of the westerly winds seems to be further supported by the west- 
■erly and southwesterly extenKion of the ire from the centers of the accutt;- 
iilation (Cf. Chamberlin and Salisbury, '06, 111. pp. 330-333). Home- 
wbat similar conditions in some respects obtained in Europe (Penck, '06, 
p. 183) but the dry winds were easterly rather than westerly as in North 
America. The European loess deposits also approached much nearer 
to the (western Europe) cosist than in America, where they remain far 
to the inferior. The Great Lake storm track mai- have bwn wider, but, 
more probably, was narrower and more intense. The northeastward re- 
treat of the ice sheet is paralleled by the northeastward migration of 
spring weather conditions (Higelow, '97. p. 48) and if this route of the 
opening of spring was initiated at this early date it must have had 
important biological consequences upon the migrating animal life of the 
Interior. The arctic and storm track tvpes of climate are perhaps the 

" ■ ■ '"IT! 



46 MICHIGAN SURVEY, 1908. 

may have, as a rule^ parsed farther south tli^n at present. If these su^- 
gestioDs are applied to the interpretation of the Glacial and post-Glacial 
Uistorj- of Isle RoyaJe, the general relations will be abont as follows: 
Succeeding the disappearance of the ice was an arctic condition with 
Khort summers and lonR winters, prevailing westerly winds, and severe 
e-tisterly or southeasterly moving storms. Such couditions as these 
would influence the direction of laie currents, wave action on the 
heaohes, and the source and movement of the lake drift, ali of which 
would greatly influence the biota- 

If the Glacial and post-Glacial adjustment of the permanent lows was 
accompanied by severe storms, this would be a factor which would cer- 
tainly influence the rate of formation and the distinctness of the beach 
lines, and it is not altogether improbable that a study of the well de- 
veloped Nipissing beach, by the development of its spits and bars, may 
furnish data regarding the lake currents and the prevailing winds. But 
in order to interpret such records it will be necessary to formulate 
criteria by means of which duration of a beach formation may be dis- 
tinguished from one of less duration but due to more severe storms and 
active currents, 

d. Th<' Ijoke Storms and their InHvenre. Tlie significance of lake 
storms is of special interest on account of the bearing of the latter upon 
the conditions of life upon the b^ch, and also upon the lake drift. Tb».t 
they must ije reckoned as an importan-t factor in the post-Glacial repopu- 
lation of Isle Boyale is evident when we recall that during the 
life of the present fauna and flora the island has never been connected 
with the mainland except by ice. Very fortunately the subject of lake 
storms has been carefully investigated by Garriott ('03) because of its 
influence upon navigation. 

The period of greatest seasonal frequency for severe storms ranges 
from September to December, with a November maximum, while 5Iarch 
contains the greatest number of such storms for the remainder of the 
year. The smallest number occur in June, July and August. 

There are several types of these storms, the most severe of which are 
those of southwestern origin and which occur between October and May. 
They are preceded by east and northeast winds which gradually become 
a gale; but when once the storm center has passed the wiad suddenly 
shifts to the northwest and is an offshore wind from Canada. Such, 
storms are frequently followed by much snow and intense cold. During 
the warmer months, storms from this direction are usually of tropical 
origin. 

Less severe storms are those coming from the middle-west. These are 
preceded by gales, first from the south and later from the east, and after 
the i>as8age of such a storm center the wind suddenly changes to the 
northwest and finally finishes with clearing weather, or if in winter, 
sometimes by a light snow. These storms are common at all seasons of 
the year, but the most severe ones occur during the cold months. 

Storms from the northwest are seldom severe; they are preceded by 
south or southwest winds, and after their passage the wind shifts to the 
west and northwest and rapidly diminishes in velocity. In winter the 
attending precipitation is generally light, in summer it is in the "form 
of thunder storms, and the high winds in squalls from the southwest 



ECOLOGY OF ISLB ROYALR 47 

at the time the center of the storm is paBaing." To this clasB belong 
the majority of lake stormB, but they are seldom severe. 

From these relations it is seen that storms whose origin is from the 
south, southwest or middle-west, are preceded by east or northeast winds 
or (middle-west) by southern winds, and followed, after the passage 
of the storm center, by northwest or west windu; while storms of north- 
western origin are preceded by south or southwest, and followed by 
weat and northwest, winds. These facts show that offshore winds from 
the eastern and southern shores of Lake Superior are the general law 
for winds preceding most storms; and that after the passage of the 
storm center all appear to be followed by west or northwest winds. These 
offshore winds are likely to be onshore winds for Isle Royale. The 
proximity of the north shore, fhe frequency and magnitude of this wind 
phenomena, clearly suggests that these factors may largely account for 
the Canadian affinities of the majority of fbe Isle Royale biota. But we 
shall see later that there are other factors to reinforce this same ten- 
dency. It may seem unnecessary to enter these details, but it should be 
remeinl)ered that the conditions under which an organism may reach the 
island is an important factor in its survival, a relation of special import- 
ance in the migration of birds. That fhe majority of these storms oocTir 
in the fall and winter, at a period of relative inactivity on the part o^f the 
Isle Royale biota, is yet a condition which would be favorable for the 
transportation of some small hibernating invertebrates. The life histories 
of these storms, especially the conditions of their termination, may be 
expected to have an important bearing upon the survival of the drift 
biota. 

There Is still another important phase of this subject, and that is 
the influence which these storms have upon the life of the shore and 
beaches. The fauna of the exposed shore of Isle Hoyale is very scanty 
and much inferior to that of the harbors, so that, generally speaking, up 
to a certain point the more protected the coast the more diversified the 
faana. This was very clearly shown by the molluscan life upon the 
shore. These storms have a powerful scouring action with the sand, 
gravel and shingle on the exposed coasts, so that a rock surface or one 
with blocks too large for disturbance by the waves is much more favor- 
able to life. 

The relation of waves to lake, currents presents a signiflcant phase 
closely related not only to the occurrence and distribution of life along 
the beach, but also to the problem of lake drift and its biolf^cal im- 
portance. A breaking wave tends to carry forward floating objects so 
that when such objects are carried along by the currents and once come 
within the range of influence of the breaking waves of shallow water, 
they tend to move with these waves into the shallow water and thus 
shoreward and are cast upon the beach in harbors, bays or about 
islands (Harrington, '95, p, VI.). 

e. The Surface Currents of Lake Superior. Mention has previously 
been made of the fact that in addition to the olTshore winds from Canada, 
which accompany certain severe storms, there are other influences 
which have a similar efl'ect upon drift — the lake cnrrents. These ai-e. 
in part, an expression of the same climatic trend and their direction 
is a resultant determined by the influence of the prevailing westerlp- 



48 MICHIGAN SURVEY, 1908. 

wiDdB, the rotation of the earth, the form and contour of the basin, 
and the positicm of the outlet. A detailed investigation of these car- 
rents was made by Harrington and Conger (Harrington, '95) wht> 
paid particular attention to the currents about Isle Royale. As these 
investigations were made during the season of aavigation, they are of 
particular interest from the standpoint of the biota, because it is dur- 
ing this same period that we must in general espect the most advantage- 
ous dispersal of plants and animals to take place. 

The simplest of these factors influencing currents are: the general 
movement toward the outlet of a lake, the prevailing westerly winds, 
the deflection to the right (or southward) of the current on account of 
the rotation of the earth. But the general form of the lake and its shore 
line, the contour of the bottom and the location of islands, introduce 
important complexities into the problem. As may be seen in Fig. 53 
Lake Superior well illustrates the influences of all these conditions. The 
small size of the outlet does not allow the escape of this vast current, 
so that there is a return along the north shore, where islands are 
encountered which produce eddies ; and in their shallow water and along 
their coasts breakers are encountered which tend to carry shoreward and 
lodge drift, 

Whea the return swirl reaches Isle Royale the problem becomes com- 
plex and is of such importance that these currents were made the sub- 
ject of a special investigation by Harrington snd Conger. In their study 
of the lake currents, bottles containing instructions were sent adrift and 
the finder was requested to communicate their recovery to the Weather 
Bureau. In this manner, supplemented by other sources of information, 
these currents were determined. The results of the investigations 
about Isle Royale are as follows : 

''Xot a single bottle has been recovered on the northwest coast of 
Lake Superior. This is not due to no bottles having been floated in that 
vicinity, as during the season of 1893 alone Mr. Conger floated 250 bot- 
tles between Duluth. Minn., and Thunder Bay, Ont. 

"This fact was deemed of such importance that the Chief of the 
Bureau, accompanied by the inspector in charge of the Lake Marine 
Service, made a special trip from Duluth, Minn., along the northwest 
coast around Isle Royale to Port Arthur, Ont, Careful note was made 
of the entire coast, all beaches examined, and observations of water 
temperature made to assist in solving the direction of the current flow 
in this region. At French River, observations were made with special 
current floats, and it was discovered that the main current was to the 
northeast from 1 to 3 miles from shore. Inside this line was found a 
current flowing to the westward. This shore current evidently begins 
farther to the east, and continues to the west end of the lake, and is 
positive at or near Duluth, as is conflrmed by investigation of the offl- 
cijiln of the city of Duluth, however, narrow and does not extend far 
into the lake. 

"Around Isle Royale there was found abundant evidence that the cur- 
rent flows to the west along the north shore of this island. Observa- 
tions of water temperature at this point are very interesting and indi- 
cate a deep stream flowing from the eastward. There appears but little 
difference in the temperature of the water at the snrface and at the 



ECOLOGY OP ISLE ROYALE. 49 

depth of 100 feet. In other localitieB to the southward there is a marked 
differeDCe between the aurface and deep water temperaturea. 

"In confirmation of this current there may be mentioned the follow- 
ing special drifts, the numbers referring to tlitme on the chart*: 
(7) Drift of the yacht Albatross in summer, during a dead calm; papei-s 
thrown overboard remained alongside of the yatch for several hoars; 
the drift was strong and uniform to the west, (8) Track of driftwood 
floated by party from the boat in a caJm off McCargoes Cove, Isle Koyale. 
(9) Drift of wreckage from the Silver Islet crib and pier which was 
washed away in a northeast storm. (10) Becord of ice floea in calm 
weather during winter of 1891 ; reported to have drifted from the north- 
east to southwest at a rate of 3 miles an hour. (11) Drift of party in 
sailboat while becalmed on July 31, 1894. (12) Steamer Cumberland, 
which went to pieces on Rock of Ages, in 1877, whose wreckage was 
distributed along the entire south shore of Isle Royale. (13) Drift of a 
champagne bottle floated by Mr. W. H. Arnold, Port Arthur, Out,, on 
October 8, 1893; and (14) the drift of a flsh barrel floated by J. H. 
Malone. keeper Menagerie Island Light, about August 27, 1885, and 
picked up twenty-six days later. The wind during this period was mostly 
from the south shore. 

"The confirmations indicate that the current between Isle Boyale and 
the Dortli shore sweeps to the west and southwest after passing the 
island and recurving rejoins the main easterly current to the south and 
west; the drift of the wreckage from the Silver Islet pier indicates that 
it recurves at some point to the southwest of Grand Marais, Minn. 

"Special attention is called to the current between Isle Boyale and 
the north shore. The great depths, the conformation of the bottom, and 
the water temperatures in this locality indicate that there is a steady 
and fairly strong current sweeping from the east through the narrow 
pathway to the west, flowing to the southwest after passing the west 
end of the island, and rejoining the main easterly current as mentioned 
above. This narrow and relatively rapid stream, like the one between 
the Manitou Islands and the Michigan mainland in Lake Michigan is 
probably the moat persistent and regular to be found in this lake, * * 

"1. Section 79. — Floated by Capt. H. 0. Jackson, steamer L. Shicka- 
luna on June 23, Ifiii'i, itt G:4o p. ni., in northwest comer. Pound by 
Charles Tiesage, Lake Linden. Mich., at entrance of McCargoea Cove, 
Isle Royale, on October 20. 1893, on the beach." 

It is thus seen that drift from the north shore of Lake Superior tends 
to be strained from the lake currents by the various harbors of Isle 
Royale. It also suggests that north shore life might also reach Kewee- 
naw Peninsula, but so far as known this has not been recognized. Drift 
waa observed in Tonkin Bay which had evidently come from a distance 
and dead birds reported by Peet, as drifting into Washington Harbor, 
probably came in part from the north shore current. The long duration 
of these currents since the Ice Age seems very probable, and undoubtedly 
they have had an important bearing upon the geographic origin of the Isle 
Royale biota, so that they cannot receive too much emphasis. 

A few words may be added concerning the probable history of the 



■ Not reproduced.on the sis 



abiGoogle 



60 MICHIGAN SURVEY. 1908. 

lake fiiirrentfl. Sinre the location of outlets, prevailing winds, topo- 
graphy of the basin and rotation of the earth all influence lake enr- 
rents, it is evident that any important change in these conditions will 
cau8e a modification in the currents. By means of these criteria theu 
we may infer what currents are likely to have existed under certain 
conditions. Some of these conditions have had a very permanent value 
in the Superior basin, because the general form of the southern shore 
(except Keweenaw Peninsula), the earth's rotational deflection to the 
right, and the iH^vailing westerly winds, made relatively deflnite condi- 
tions. Thus the early Glacial lakes in this basin, which had south- 
western outlets, must have had different currents, perhaps more or 
less against the prevailing westerly winds, and the absence of large 
islands would be favorable to uniformity. Later at the Algonquin stage. 
Fig. 52, there must have been a very complicated system of lake fxu'- 
rents, perhaps a rough outline of those of the present Great Lakes, at 
least in the deflection toward the right shores on account of the rota- 
tion of the earth, and to the eastward on account of the prevailing 
westerly winds and the eastern outlets. The broad connection between 
the Superior and the Huron bnsins perhaps also favored a north shore 
i-eturn whirl, while at the Xipissing stage, Fig. 54, in the Superior 
baiiia the currents were in general quite similar to those of the present 
lake, but more simplified in detail by the greater depth of the lake. 

If such general relations as these obtained, it will be seen that the 
north shore i-etum whirl may have been of considerable duration, and 
that the opportunity for these (^irrents to carry life from the south 
shore must have been constantly less favorable than the chances for 
them to efl'ect transportation from the north shore of the Superior basin. 
Tn this basin then it seems that the currents were flrat relatively simple, 
became quite complex at the Algonquin ^tage and were simplified at 
the Nipissing stage. A detailed study of the beach lines such as those 
of the Nipissing, might add much positive information as to these 
ancient lake currents and their biological relations. 

f. The Origin of the Habitats. Isle Royale is abont 45 miles in length, 
has. an average width of about seven or eight miles and an area of abont 
210 square miles. The Nhallow soil, rock ridges, forested swamps, lakes, 
Bmall streams, rocky coast, and harbors provide a variety of 
conditions and furnish play for such a variety of processes that many 
diverse habitats are produced. Generally speaking, the island is covered 
with a stunted coniferous forest growth. Attention has already been 
called to some of the conditions and processes "which have produced 
the major environmental i-egions and the general topography of the 
surface. If Isle Royale had high mountains and greater extent, very 
different habitats would be expected. 

As we have seen, the entire surface of the island has been beach, 
and previous to that it had been a reef in the lake, so that the beach 
represents the original land habitat upon the island. Generally speak- 
ing this habitat has migrated from the crest of the Greenstone Range 
downward for about 550 feet to the present lake level. With this 
progressive downward movement, there has been an increasing area 
exposed to subaerial processes of erosion. The origin of the harbors 
has been a part of the beach problem, but that of the protected beacb, 



lyCoogle 



ECOLOGY OP ISLE ROYALE, 51 

these with the falling of the lake have migrated outward, a^ ia sug- 
gested b; the courses of the main streams occnpying the rook valleys. 

The very immature condition of the drainage shows that during the 
present post-Olacial cycle only comparatively slight ohanges have 
modified the relief from the eondition in pre-Glacial times; it is thus 
lai^ly - an inherited topography, hence the conseqaent drainage. 
It should perhaps be added, however, that the date of the faulting is 
not definitely known; it may be very ancient, but the weight of the 
ice sheet may have had considerable influence. It thus seems probable 
that with the decline of the lake level there has been an increase and 
downward elongation of the stream environments, and that their, course 
has been determined largely by the pre-Glncial topography, supple- 
mented, of course, by the southward tilting of tbe land. The lake 
basins have had an origin similar to that of the streams and have 
tended toward extinction by tilting, inwash, organic debris and to a 
limited extent by the downcutting of outlets. On aceount of the rela- 
tively eniall amount of erosion by the ice sheet it is probable that the 
shallow swamps and the smaller sti-eams were influenced more by tbe 
ice than those features related to the greater relief of the surface; 
even moderate tilting nould considerably influence such an environ- 
ment, beoauHe within the major valleys the divider are generally low. 

The origin of certain land habitats only i-emnins to be considered. 
These have undergone a complex succefision of changes. The resistent 
lava of the Greenstone had been the least reduced by ei-osion so was the 
first to emerge from the lake le\'el. This was first a beach, and as the water 
fell from its crest the up{>er beach migrated to lower levels and the 
land habitat continued to increase in area. The beach line itself 
expanded laterally, if not in width, as the area of the island increased. 
HTien once the exposed rocks were beyond the reach 'of the naves, 
weathering and erosive processes were initiated which tended to pro- 
duce a residual soil. Plant remains from lichens were perhaps the 
first humus formers, and it ia probable that it was not until the 
period of Lake Algonquin that the lake drift which was washed ashore 
became a source of such material; but winds, birds, lake currents and 
the wavea may all have contributed pioneers of tbe higher plants. 
The harbors at the northeastern end of the island would tend to strain 
out the drift from the southwestward flowing current and the return- 
ing one along the southeastern coast of the island would tend to lodge 
drift in Washington Harbor and the Siskowit Bay region. 

As the water continued to fall to lower levels, the land biot;i 
followed down the slopes liehind the receding beach. By the 
Xipissing stage, the vegetation and many animals i\-ere probably 
well established and had begun to actively encroach upon the 
swamps and lakes and thus tended to increa.«e the land habitat. 
With the tilting that followed the fonnation of the Nipissing 
bench, a readjustment must have taken place Itetween the land 
and water habitats, hut to what degree their relative areas weit^ 
infiuenc<'d is not known. During the initial elevation ponding would 
be expected at the northeastern end of the island, but with a greater 
elevation this same area would l)e well dniined, as the divides in the 
valleys are low and the transverse drainage near the central iwirt uSfc 



63 MICHIQAN SURVEY. 1908. 

the island would tend to prevent extensive ponding, combined with 
the fact that the ralle.va extended in the Bame general direction as the 
uplift and not across it. It therefore appears that many processes 
have tended to inoreane the land habitats at the expense of the aquatic, 
such as the falling of the lake level, the encroachment of organic re- 
mains on the depressions, the perfecting of drainage lines and the 
tilting of the surface. ■ • 

With the advent of the forest a habitat dirferentiation developed 
in contrast with the natural openings. These openings were originatl.? 
doe to the lack of soil, as on the ridges, wave action, as on the beach, 
or an excess of water as in the depressions. With the accumulation 
of soil, the downward migra,tion of the wares, and tbe filling up or 
draining of tbe depressions, the range of the forest has been extending, 
and is tending to completely cover the surface. 



REFEBBNCES. 

Bigelow, F. H. 

1897. Htornis, Storm Tracks, and Weather Forecasting, U. R. Dept. 

Agric, Weather Bureau, Bull. No, 20. 
Chamberlin, T.C., and Salisbury, R. D. 

1906. Geologj-. N'ew York. 
Coleman, A. P. 

1906. Iron Ilanges of Eastern Michipicoton. Report Can. Bureau of 
Mines for 1906, 15 (I), pp. 173-199. 
Cooper, W. F. 

1906. fleological Report on Rav County. Mich. Qeol. Surv.. Ann. 

Rep. for 1905, pp. 135-426. 
Denison, F. N. 

1898. Tile tireat Lakes as a Sensitive Barometer. Rej). of the 

British Assoc. Adv. Science, 1897, pp. 567-568. 
Fassig, O. L. 

1899. Types of March Weather in the United States. Amer. Jour, 

'of Sci. (4), S, pp. 319-338. 
Foster, J. W., and Whitney, J. J). 

1850. Report on the Geology and Topography of a Portion of the 

Lake Superior Land District, in the State of Michigan. Ex. 
Doc, Ist Bess. 31st Cong.. IX, Part 1. (Copper Lands.) 

1851. Report on the Geology- of the I^ake Superior Land District. 

Part II, The Iron Region. Senate Doc. Spec'l S. 32nd Cong., 
Vol. 3. • 
Garriott, E. B. 
1903. Storms of the Great Lakes, U. S. l>ept. of Agr., Weather 
Bureau, Bull. K. 
Gilman, H. 

1873. The Caribou on Lake Superior. Amer. Nat., 7, p. 751. 
Goldthwait, J. W. 

1907. The Abandoned Shore Lines of Eastern Wisconsin. Bull. 17, 

Wis. Geol. and N.Tt. His. Survey. 

1908. The Records of the Extinct Lakes. 111. Geol. Sqrvey, Bull. 

No. 7, pp. 54-68. i„C.OO<^IC 



ECOLOGY OP ISLK ROYALE. 53 

HarringtoD, M. W. 

1895. Surface CurrestB of the Oreat Lakes. a» Deduced from the 

movements of Itottle Papers during the Beasons of 1892, 1893 
and 1894. U. S. Dept. of Agric, Weather Bureau, Bull. B, 
Revised Edition. 
Hubbard, G. H., and Sctiwarz, K. A. 

1878^ Jiist of Coleoptera in tlie Lake Superior Region. Trans. Amer. 
Phil. Soc., 17, pp. 627-C66. 
Jackson, C. T. 

1849. Geological and Mineralogical Reports. Senate Ex. Doc, No. 
1, iBt Sess. 3l8t Cong., 3, 371-935. 
Lane, A. C. 

1898. Geological Report on Isle Royale, Kficbigau. Mich. Geol. Sur- 
vey, VI, Part 1, pp. 1-281. 
1900. Geological Report on Huron County, Michigan. Geol. Surv. 
Mich., 7, Pt. II. 
liawBon, A. G. 

1893. Sketch of the Coastal Topography of the North Side of Lake 
Superior, with Special Reference to the Abandoned Strands 
of Lake Warren. Twentieth Ann. Rep. Minn. Geol. and Nat. 
Hist. Survey for 1891. 
Montgomery, T. H. 

1906. The Analysis of Racial Descent in Animals. 'Sew York. 
Penck, A. 

1906. Climatic Features of the Pleistocene Ice Age. Ge<^. Jour. 
27, 182-187. 
Buthven, A. G. 

1906. An Ecological Sur\'ey in the Porcupine Kfountains and Isle 
Boyale, Michigan. Mich. Geol. Burvev, Ann. Rep. for 1905, 
pp. 17-56. 
Buthven, A. G., and Others. 

1906. An Ecological Survey in Northern Michigan. Mich. Geol. Sur- 
vey, Ann. Kep. for 1905, pp. 1-133. 
Smith, S. I. ' 

1874. Ann, Rep. U. S. Fish Comm. for 1872 and 1873. I't. II. pp. 690- 
707. 
Spencer, J. W. 

1891. Deformation of the Algonquin Beach, and the Dirth of Lake 
Huron. Amer. Jour. Sci. (3), 41, pp. 12-21. 
Stupart, R. F. 

1898. The Climate of Canada. Scottish Geogr. Jour., 14, 73-81. 
Taylor, F. B. 

1896. Preliminarv Notes on Studies of the Great Lakes, Made in 

1895. Anier. Geol.. 17, 253-257. 

1896. The Aigoutiuin and Nipissing Beaches. Amer. Geol. 17, 397-400. 

1897. Notes on the Abandoned Beaches of the North Coast of Lake 

Superior. Alner. Geo!., 20, 111-128. 
Transeau, E. N. 

1905 a. Forest Centers of North America, Amer. Nafuralist, 39, 

879-889. 
1905 b. The Bogs nnd Bog Flora of the Huron River Vallev. Bot. 

Gazette. 40, 351-375. 



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ECOLOaY OF ISLE ROYALE. 



THE ECOLOGICAL BELATIOKS OF THE INVERTEBRATE FAUKA 
OF ISLE ROYALE, MICHIGAN. 

BY DB. HBNBr ALLAN OLEASON. 

/. Introduction. 

The most recently emerged portions of Isle Royale are the rock and 
gravel beaches which together constitute virtually the entire shore of 
the island. Animal life is found apon them almost to the edge of the 
water, and well within the limits of wave action. The phyaiographie 
succession in the island is ench that the areas originally occupied by 
beach pass through a series of changes in the physical factors, a series 
which is accompanied, sometimes hastened, sometimes retarded, by cor- 
responding vegetational successions, and which culminates in the final 
or climax plant association of balsam and spruce forest. The detail of 
this physiographic and vegetational succession is by no means uniform; 
it may proceed along either of two well-marked lines, depending on the 
immediate physical and biotic conditions, certain intermediate stages 
may be prolonged or omitted entirely, and various other deviations may 
occur. Nevertheless the final stage is always the same. Accompanying 
the changes in physiography and vegetation is a similar and dependent 
change in the fauna, so that there is a corresponding series of animal 
associations, beginning on the beaches and developing in the same di- 
rection, with the same deviations or omissions, to the final or climax 
association in the balsam-spruce forest. 

The preceding general statement rests on the assumption that the 
areas now occupied by the climax biotic associations have developed from 
the beach associations through a series of stages intermediate in time 
corresponding to those associations which now stand intermediate in 
space between the two extremes. Or briefly, as some ecologists have 
expressed it, the lateral distribntion in space recapitulates the vertical 
distribution in time Such an assumption is evidently closely akin to 
the recapitulation theory of the evolutionists, and just as that so-called 
biogenetic law has been accredited with more than its true value, so 
has this ecological dictum possibly much less importance than has been 
usually supposed. The weakness lies in too little consideration of the 
time element. It is certain that the higher land in Isle Royale has 
been subnierged. This is shown by the old beach marks now many feet 
above the present level of the lake. Consequently by the gradual emer- 
gence all of the island has passed through a beach stage. But it is un- 
warranted to conclude from this that the faunal or floral associations 
of the former beach were similar to those of the present, or that in the 
intermediate stages the biota resembled that which now occupies the 
area between the ancient beach and the present shore. While it is like- 
wise certain that with a continued subsidence of the lake level the pre- 
sent beaches will eventually be left far above the water, it must not 
therefore be assumed that their biota will show the same t 
or reach the same climax as those of the past. Changes in the tei 



88 MICHIQAN SURVEY, 1908. 

tare or rainfall may certainly keep pace with the changes in lake levels 
or even be caused by it, and in either case they would exert a profound 
inflnence on the biota. Migration of species is still taking place among 
both plants and animals, and may introduce new or even dominating 
species among the present forms. The so-called equatorial pressure of 
southern species is fully as strong now as it was directly after the close 
of the glacial epoch. Lastly, and most important of all, the influence 
of the biota itself is always to be reckoned with. Both plants and 
animals are continually becoming more plastic, adapting themselves to 
new conditions, and extending their habitats into new associations. 
They push forward more rapidly than the changes in physiography, some- 
times hastening and sometimes retarding physiographical action, and 
at all times greatly inflnencing the subsequent snccessions. 

A biotic association may develop into another by a mere re-arrange- 
ment of the interrelations, numerical or otherwise, of the component 
species, without the necessary loss of some or addition of otbers. Bnt 
such cases are rare, and the Isle Royale observationa show that no two 
associations have exactly the same species, and that with each pro- 
gression there has been an addition of certain forms which bto>me the 
most characteristic types. The first bit of beach fo^ed was occupied by 
an association possibly not unlike that of the present beaches. All the 
species must evidently have immigrated from beyond the island. When 
the soil deposits on the beach were sufficient to support a second as- 
sociation its species were derived partly from the beach itself and partly 
from new immigrants. The further development of biotic associations 
on the beach was then possible not only from immigrants, bat also from 
the two associations already present. Similarly at the present time 
each association on the island is constantly being invaded by species 
frotQ all the others, and many of them are actually able to establish 
themselves. This tends toward a homogeneity in the biota hardly in 
full accordance with the recapitulation idea. Indeed, it is very probable 
that independently of all physiographic agencies the whole surface of 
the island would eventaally be occupied by the balsam-spruce forest and 
its attendant faunal association. 

In many cases it is virtually certain that the lateral succession does 
faithfnlly repeat the vertical, and the zonation of plants around a pond 
may be taken as an example, but the filling of a pond is only a single 
step in the genetic development of the biota of an island. 

With this preliminary note of warning, the truth of this recapitulation 
theory will be assumed for the island, and the discussion of the insect 
and molluscan fauna will follow the genetic lines indicated in the first 
paragraph. 

The relationship of the various physiographic types on the island 
to each other may conveniently be expressed by a diagram (see end of 
pai)er), indicating the direction of the devolpment hy arrows. It must 
be remembered that practically any one of the intermediate etages may 
be omitted. 

//. The Lake. 

The lake (Superior) must obviously be regarded as the first stage in 
the genetic development of the faunal associations. Broadly speaking, 
the lake fauna is divisible into two main groups. The first is pelagic in 



ECOLOGY OF ISLE ROYALE. » 

character and includes those species whose distribution is entirely inde- 
pendent of the shore, for example, most of the species of fish. The 
second group is littoral; the spe^^ies occur aJong the shore in comparti- 
tively shallow water, and are to a greater or lesB extent dependent upon 
the land in its relation to the character and slope of the bottom and to 
the motion of the water. Members of the latter group only are con- 
sidered here. 

The two dynamic factors just mentioned are the most important ooes 
that influence the biota of the lake. There are no currents of suiBcient 
rapidity to affect the animal life. The direction of the wind, whether 
ofF-shore or on-shore, may respectively lower or raise the level a few 
centimeters, especially when the wind blows lengthwise of the long 
narrow inlets, such as Conglomerate Bay (Fig. 11). Borne fixed or 
slow-moving species may accordingly be alternately submerged and ex- 
posed, while motile forms can at once adjust themselves to any change 
of level. Of far greater importance is the motion of the water caused 
by wave action. It is only on rare occasions that the lake is quiet. 
Qentle waves come in nearly all the time, and after storms become of 
great violence. Wave action is of itself sufBcient to inhibit the growth 
of shells along tbe exposed shores, where they might easily be torn 
loose and crushed against the rocks. Such forms are consequently re- 
stricted to the shores of the smaller bays or to the lee side of islands. 

Wave action is of importance further in determining the character 
of the bottom. Where the shore is exposed directly to the lake it is 
usually of massive rock, all the fragments having been washed down to 
deep water. In small shallow coves, where the waves break always in 
one direction there is usually a sloping beach of gravel extending across 
the end perpendicular to the direction of the waves. Every breaker 
sorts over this gravel so that it is nearly impossible for a fauna to 
develop. In lat^r coves or bays, where the violence of the wave action 
is reduced by distance, the gravel is finer or e\'en a beach of sand may 
rarely be formed. Along the steep or cliff-like sides of these coves the 
bottom is frequently covered with angular rock fragments too large 
to be moved by the water. These are frequently inhabited by shells. In 
general the development of a free littoral fauna demands quiet water 
where the animals will not be dashed on the rocks or stranded on the 
shore, and for attached species there is required either quiet watw or 
a firm bottom which will not be dislodged by the waves. A raor^ de- 
tailed discussion of this as affecting the distribution of shells will be 
given later. 

In the larger inland lakes, of which Siskowit Lake, the only one of 
the class studied, may be taken as an example, essentially the same con- 
ditions obtain as on Lake Superior itself. The difference in temperature 
and content of the water seems to be of minor importance. The waves 
in the larger lake can naturally reach a larger size, and their influence 
is felt far into the bays. Thus at the head of Rock Harbor, about six 
kilometers from the lake prosier, the distribution of shells and the al- 
most total absence of free forms indicate that even there wave action 
is of importance. In Siskowit Lake, although larger than Rock Harbor 
the force of the waves is so reduced by ever^' hendlnnd or island that 
on the quiet water in their shelter a rich fauna of such free forms as 



60 MICHIGAN SURVEY, 1908. 

water-Btriders and whirligig beetles is found on the enrface, while 
numerous muBsel shellB live on the silt or sand bottom. In Somner Lake 
and others of limited area the motion of the water has no measurable 
effect on the biota, and they wilt therefore be treated ander a separate 
heading. 

The distribution of Bhells along the shore, particularly species of 
lAmnaca and Physa. is of especial interest. Having relatively low 
motility they are correspondingly limited in their distribution and the 
factors governing it are more readilv determined. These will perhaps 
be made clearer by concrete illustrations. 

Tonkin Bay is a small inlet about half a kilometer long, opening 
to the east upon the lake, and with steep, approximately parallel sides. 
It is narrowed half way up by two beaches lying perpendicular to its 
length. By this the wave action on the upper part is reduced, but still 
may sometimes be sufficient to wash heavy driftwood upon the beach. 
Id the outer half the wave action is but slightly less than on the lake 
itself, and no shells are found. In the inner or upper half Limnaea 
atagnalis L. (Nos. 50, 54, 57), LUnrwiv nnarginata Say (Nos. 50, 57), 
and Physa sayii (Tap.) <Nob. 50, 57). live along both sides where the 
bottom is rock, but not across the ends. They live only on a rock 
substratum, which may be either hori7,ontal or vertical, and in water 
no to 45 cm, in depth. The larger species, Limnaea atagnalis, is more 
nbundant in the deeper water, and only the smaller species live at a 
depth less than 15 cm. They then prefer the vertical walls to the hori- 
zontal or flat bottom. 

Conglomerate Bay is a rocky inlet iFig. 11) similar to the one just 
■ described and about 1.6 km. long. Being wider at its mouth than 
Tonkin Bay the force of the wave action is felt farther up the hay. Kear 
the end the waves have little effect, as is evidenced by a sandy beach 
(Fig- 4), almost without driftwood. At the upper end of this bay alontc 
the north side Limnaea emarginata Say (Nos, 118, 12S) and Physa aayii 
Tap. (Nob. 118, 125) are found in water 15 — 45 cm. deep, in the deeper 
water on the tops of flat rocks, in the shallower water, also on the veiti- 
cal sides and in small crevices. They never occur on the sand or gravel 
deposited around the rocks, as is frequently the case near the sand 
beach at the upper end of the bay. The distance to which they extend . 
from shore is greatest opposite the concavities of the shore line and least 
opposite the small rocky headlands. Their distribution in both Tonkin 
Bay and Conglomerate Bay seems to be regulated mostl.v by the wave 
action, since they seek the most protected places, avoid the shallow 
water where the waves would strike them most, and do not live on 
loose or small rocks, gravel, or sand, which would easily be dislodged. 
The fact that the smaller shells are found at the least depth, while the 
larger Limnaea emarginata inhabits the deeper water, would indicate 
that the small size of the former renders tliom less easily dislodged by 
the waves. Again their gi-eater abundance on the north side suggests 
the possibility of a light relation. 

Siskowit Lake, with its rocky shores and large area, offers essentially 
the same condition as Lake Superior itself, and the shells have the same 
general distribution. Along the very gently sloping rocky shore near 
the outlet Limnaea stagnalis occurs in abundance, always at a depth 



ECOLOGY OF ISLE HOYALE. 61 

of 1040 cm. Alon^ the south Ride of n large island near the Houth shore, 
where they are sheltered from waves in every direction, the same spe- 
cies is abiindaot. They live on rocks in the full sun in \vater 10 — 45 cm, 
deep, with the optimum depth at 20—25 cm. They may occur on the 
tops or sides of rocks, but never on the sand between them. Associated 
with the Ltmnae-d, but much lees abundant, are Planorbia bicarhtatun 
royaUnais Walker (No, 210J, P. campanulatug Say (Nos. 210, 211), 
Lamp^Us htteolus (lAm.) (Sob. 210, 211), Anodonta tnarginata Say 
(So. 210), and Anodonta grandis fooiiana Lea (Nos. 210, 211). 

Opportunity was given to observe the behavior of Limruiea stagnalis 
(So, 217) in waves of some size near k small circular island half a kilo- 
meter out in the lake. The bottom was gently eloping, and either of 
solid rock or of large rounded fragments. There were no overhanging 
trees, so the shells were found in uniform abundance in the usual depth 
of water on all sides of the island. At the time the island was visited 
a strong wind was blowing, and the waves were probably nearly as high 
as they ever become on Siskowit Lake. One or two shells were seen 
which had been washed loose, and of course would be unable to re- 
attach themselves until the waves abated. It would be expected that in 
such cases the shells might be crushed or broken or the animal killed. 
That such may happen was evidenced by finding a few live shells which 
had been cracked and then healed, leaving an irregular surface. Their 
occurrence here and elsewhere only upon rocks of considerable size 
shows that they require a firm substratum, and where the rocks are 
free from any coating of slime they can certainly endure higher wares. 
Around the island under discussion the rocks were washed perfectly 
clean. 

The beach in front of the ramp at Siskowit Bay {Fig. 29) was inhabited 
by large numbers of (So. 200) Physa sayii Tapp,, Pkysa sp,, Limnaea 
stagnalis L., and Limnaea einarginata Say, so that more detailed obser- 
vations of them could be made, and a few experiments carried out to 
show their sensitiveness to the depth, or bathytropism, as it has been 
termed. The beach here is of rock with a gentle slope of about one in 
five, corresponding to the dip, except where blocks have worn off, leaving 
low vertical walls. The wave action here is very light, its force being 
cut off by a series of islands lying between the beach and the main body 
of Siskowit Bay. This was well shown by the conditions on ,\ugust 2, 
when there was scarcely a ripple inside the islands, although the bay 
outside was covered with whitecaps. The beach is covered with a thin 
coat of slime formed mostly of excrement from the snails. 

On such a beach snails may live close to the edge of the water, but 
the larger lAmnaeaa still occupy their usual depth of 1.5 to 4.5 deci- 
meters. About 10 A. M., on August 3, all the shells to a depth of about 
1 decimeter were gathered from a strip of the beach about 10 meters 
long. They were comprised in the following species: (So, 200) Limnaea 
stagnalis L., Limnaea cmarginata Say, Phyaa aayii Tapp., and Phyta sp. 
The smaller Physaa were especially abundant and about 200 of them 
were taken. Four hours later, at 2 P. M., 60 shells, all of the smaller spe- 
cies, had migrated upon the same strip. The only evidence concerning 
the way that they came is that one shell of Limnaea stagnaliit was seen 
to drift up over a low wall into the shallow Eone. This method could 



62. MICHIGAN SURVEY, 1908. 

hardly account for 60 of the Bmaller ones, however, appearing in so short 
a space of time. It may be taken b» indicating a geoeral and continued 
migration in all directione within their bathjtropic limits. 

It was noticeable that the large Limnaea emarginata and Limnaea 
»taffnali8, aside from the one specimen mentioned above, live at an aver- 
se depth of '3 dm. and never deeper than 4.5 or 5 dm. To test their bathy- 
tropism six of them were picked out of the deeper water by band and 
held in contact with the bottom in the shallow zone nntil they extended 
their feet and attached themselves. At this time the water was very 
quiet, moving just enough to cause a faint sound on the beach. But the 
size of the shell of the two Limnaem is so large that they offer consider 
able surface to the water and are consequently easily washed loose. Two 
of the six swung a little from side to side and were then washed off and 
carried by the nndertow into water 3 dm. deep, where they again at- 
tached themselves. A third, without being shaken by the waves, clung 
to the rock for some time, then suddenly let go its bold and drifted over 
a low ledge into the deeper water. Two others immediately started to 
crawl down the slope, and one in about fifteen minutes, the other in 
about half an hour, had crawled over the ledge into water 3 dm. deep, 
where 'they both remained stationary. The sixth remained attached, and 
in three honrs had crawled 2 dm. parallel to the shore, keeping at the 
same depth. The n^ morning, twelve hours later, it had disappeared, 
and of course could not be recognized in the deeper water. 

On Angust 4 two shells of Limnaea appeared in the sbaDow zone, bat 
it is not known whether they drifted or crawled up. They were there 
at least three hours. After they were last observed a fresh breeze 
sprang up from the east and the slight wave action caused by it prob- 
ably washed tbem down. 

The level of the lake varies somewhat with the direction and inten- 
sity of the wind, so that in front of the camp a strip of beach up to 
5 dm. in width may or may not be covered with water. The smaller 
shells, Phyaa nayii Tapp. and Physa sp., live in this zone in spite of the 
fact that they are sometimes out of water. So far as observed they are 
never exposed for any considerable length of time, so that they do not 
become dry. Then again the weathering of the rock has left bowl-shaped 
hollows a centimeter or so across and about the same depth, and the 
snails usually get into them. 

To summarize, the known facts bearing on the distribution of these 
four species are as follows: 

1. .Their lower limit is 4.5 to 5 dm. depth of water, governed possibly 
by the water-pressure or the food supply. 

2. The upper limit is for Limnaea stagnalis and Limnaea emarginata 
l.B dm. of water, for Phyaa sayii and Physa sp. the shore-line. The 
cleaner the rock and the less the wave action the shallower the water 
which they may inhabit. 

3. Their horizontal distribution is controlled by (a) full exposure 
to the sun; (b) a rock bottom; (c) a certain minimum of wave action. 

But two species of insects were collected which should properly be 
eoDsidered here, caddice flies and stone flies. The larva cases of the 
caddice flies were collected only in the outlet of a small stream 
emptying into Rock Harbor, in 1 — 1.5 ni. of water (No. 163 or 



ECOLOGY OF ISLE ROYALB. 03 

164), bat tbe iinagOB were common all along the diore of the lake, ee- 
pecially on the grarel beaches. One (No. 192) was taken on the boat 
about 2 km. off the south shore of the island. Stone flies were also fre- 
quently collected along the beaches, where they came op to breed. They 
were most numerous, however, on steep or even vertical cliffs with SKiuth- 
em exposure (Nos. 24, 80). Near the entrance to Conglomerate Bay 
(Fig. 2) they were seen collected in such a place by thousands. The 
water there was at least 4 m. deep. 

A few hair-worma, Qordvas aquaticm (L.) (No. 207), were collected 
in 2 to 3 dm. of water on the rock beach {Fig. 30) in front of the camp 
on Siskowit Bay. 

The various tnussels collected in Biskowit Lake and elsewhere, even 
though Bometimes associated with lAmnaea stagtMilU, belong rather to 
the associations of the smaller inland lakes. 

///. The Inland Lake. 

The smaller lakes are mainly surrounded by tamarack swamps, with 
the vegetation showing the characteristic zones, certain ones of which, aa 
the rushes, water-lilies and pond weeds, live in the lake itself. The bot- 
tom is covered with peaty mud or with slime, and the wave action is 
never severe enough to interfere with the growth of either faana or 
flora. Id many of the smaller lakes, in fact, the water lily zone is so 
wide and the open water so restricted that there is practically no wave 
action at all (Fig. 46) . Accordingly both fauna and flora are 
richly developed both in species and individuals. The fauna may 
be roogbly classified into several groups according to their habitat in 
order to facilitate description. The interrelations of the different spec- 
cies are complex in the extreme, and of course could not be properly 
worked out in such a short time as tbe lakes "were under observation. 

a. The Faiimi of tlic Bottom. In Sumner Lake (III, 5) [Figs. 18-22 
and in sheltered places in Siskowit Lake several species of shells live on 
the bottom in sand or mud and at a depth of from 3 dm. to 1 or 2 m. 
Planorhis trivolvis Say (No. 135) lives in the shallower water, prefer- 
ably in mud. It is nowhcK abundant, but was collected in both lakes. 
One specimen only was found in Sumner Lake in a little pool with mod 
bottom. Shells were commoner on the shoreward side of an island in 
Siskowit Lake, on a bottom composed of sand and mud. They were 
well buried under the sand and the majority of the shells were dead. 

Mussel shells, especially Anodonta marginata Say and Anodonta 
grandis Lea, were common in all the smaller lakes and at the upper 
end of Rock Harbor. They were most abundant in the deeper water 
with a sand bottom, particularly where there was comparatively 
little v^etation. In certain sheltered bays at the upper end of Sisko- 
wit Lake they were especially numerous. Muskrats carry them to the 
shore to eat, and leave the empty shells in heaps, which were conspiou- 
ooe sights along most of the lake shores. In Sumner Lake live shells 
were very scarce, but the piles of dead ones on the bank testified to their 
-former abundance. 

At the upper end of Bock Harbor some small shells, Planorbia Btonn'- 
natus Sav (89), Planorbia exacutua say (89), PUmoriia parvus Say. 
<89, 103,*164), Talvata tricarinata Say (89, 163), Valvata sincera »^4c 



64 MICHIGAN SURVEY, 1908. 

landeri Dall (8ft. 163, 164), AmnicoJa hmtrica Pile, (89, 163, 164), and 
Piaidium sp. (163, 164), and AmphipodH were dredged from a depth of 
1.5 to 2 metera near the mouth of a Bmall stream {Fig. 22) where the 
bottom was thickly covered with email twigs and other coarse vegetable 
debris. From the same place the caddice fly larvae were obtained, as 
mentioned previously. The same fauna was collected in the stream it- 
self, but only near the mouth, where the water was deep, the current 
slow, and the conditions io general much like those of a lake. May flies 
probably breed in similar places. No larvae were seen, but a few imagos 
were collected (No. 178). 

The fauna of the bottom shows a connection through the presence of 
Pisidium sp. in the last case with that of the small streams in the 
tamarack swamps and with that of the brooks, like the outlet of Siskowit 
Lake. The accumulation of vegetable debris and the more restricted 
amount of water are both approaches toward the conditions in the former 
places. In Siskowit Lake, where Planorbis campanuJatus, PUinorJiis 
bicarinatus royalenais and Anodonta grandw footiana were associated 
with lAmnaea- stagnalis, another transition was shown between the faunas 
of the inland lakes and the larger lakes as tvpifled by Lake Superior 
itself. 

b. The Free Fauna of the Water. No species were observed eicept 
fishes and leeches. The latter were abundant in Sumner Lake, especially 
among the water lilies and in the shallow water along the shore. 

c. The Fauna of the Surface. Hardly belonging properly to this- 
group weiv the small shells, Limnaca catascopium Say (220), Physa sp. 
(220, 221), VaJvata aincera nylanderi Dall (220), and Amnicola limoaa 
Say (220), found abundantly on the under side of water lily leaves. I^ir 
distribution is directly controlled by that of the water lilies, that is, 
near the shore, and in the larger lakes only in the sheltered bays. Pro- 
bably a third of the leaves had one or sometimes two shells attached 
to them. Water striders, Gerris remigis Say (No. 96), were abundant, 
usually near shore in the water lily zone, but occasionally out in the 
open water. Whirligig beetles, Oyrinua wu'nwtHs Fabr. (No. 219) were 
also common, but not abundant on the smaller lakes. In the sheltered 
bays of Siskowit Lake they collected in immense awarma, keeping mostly 
near the shore among the water lilies and under overhanging brush. 
Doiiacia proxinia (Nos. 171, 184) and Donacia cincticonii-a (Nos. 171, 
175) were abundant on Sumner Lake, resting on the water lily leaves. 
When alarmed they would fly a short distance cltwe to the water, making^ 
a little trail Miind them, and alight on another leaf. 

d. The Free Aerial Fauna. Dragonflies of several species are abun- 
dant along all of the lakes. They usually keep close inshore or over 
the water lilies, aud fly regularly in patrols around the lake, searching 
all the time for insects but keeping up a uniform rate of speed. Aeachna 
sp. was probably the most abundant, and associated with it were 
Enallagma hageiU Walah and Leucorkinia proxima Anth. The butterfly 
Argynnis atlantia Edw, also occurs (No. 169). 

The inland lakes may be regarded as small detached portions of the 
main lake, cut off from it by the lowering of the level of the latter. 
Since they are composed of stagnant water with little or no wave action,, 
where oi^nic material may accumulate in quantity, they support a 



ECOLOGY OP ISLE ROYALE. 66 

different fauna and their genetic development is along a different line, 
culminating however in the climax type or balsam-spruce forest. The 
only intervening stage is the tamarack swamp. 

IV. The Tamarack and Arhor Vitae BwampH. 

Nearly every inland lake in the Isle Royale region ia wholly or partly 
surrounded by tamarack Bwaiiips, {Figs. 1^, 19, 22, il, 47, i8). It is 
not necessary to discuss the general stnictnre of the vegetation, since 
that is described elsewhere in this report, but il may be indi- 
cated here that the ground cover is a spongy mass of sphagnum covered 
with a dense growth of erieaceous shrubs, such as Cassandra and Ledum, 
and that the trees are almost entirely tamarack and black spruce. The 
forest cover is open enough to allow ample illumination. Tamarack 
swampe may be found of all ages, from those developing at the edge of 
a lake to those which have completely covered the lake and are now 
dying as an association. Their surface is generally level, the older parts 
being successively somewhat higher as they are built up by the accumu- 
lations of peat. 

When the level is nearly that of the lake the beds of sphagnum are 
interspersed by little streams or pools of water, some of them being 
merely extensions of the lake itself, or some of them serving as inlet 
or outlet. The smaller ones have no bottom except the sphagnum itself, 
while the larger have a loose incoherent bottom of slime. In the larger 
of these streams are found small bivalve shells, Pisidium sp., embedded 
in the slime at the bottom (No. 230; V-5), and other material; and the 
beetles Baliplus ruficoUis DeG., Bydropontu trigtis Payk, and Agabue 
congener Payk. (No. 230, V-5). In the smaller ones, which are fre- 
quently only a decimeter or two wide and half as deep, there is no 
difference in the vegetation except for a little TJtricuIaria in the bottom. 
Animal life is there very scarce (No. 237, V-5), J>ut. included Pisidium 
sp. 

As the swamps become older the water is limited to small shallow 
pools, seldom more than one decimeter deep or three or fonr decimeters 
wide. Their bottoms are covered with dead leaves and sphagnum, and 
they are usually densely shaded by the forest growth above. In them are 
found small bivalves, Pisidium affine Pterki (77A, TflA), P. subrotundum 
Sterki (116, lf*l. \»2, 2371, P. subrotundum Prime (IIC, 237), and water 
beetles, HaUplus ruficoUis l)eg. (No. HC, 1-4) and 8cutopterus homii 
Cr. (No. 181, 144). The latter is restricted, so far as observed, to this 
single habitat in the pools in tamarack and arbor vitae swamps. Dragon- 
flies are the principal aerial insects, but are not abundant. A fly (No. 
240, V-5) was taken on the flowers of ^olidago neglecta. 

In still drier swamps, where there is no longer any standing water, 
{Fig. 14). ants are a characteristic feature of the fauna. They 
build huge, dome-shaped nests, 4 to 7 dm. high, composed within of 
sphagnnm and other vegetable debris, and smoothly covered on the 
ontside with leaves of Cassandra, doubtless to prevent drying. Formica 
adamtii Wheeler (No. 115, I-fi) seems to be the only species concerned, 
and a nest from which the collection was made was photographed. No. 
114, taken at the same time from a similar nest, has been identified 



66 MICHIGAN SURVEY. 1908. 

as FomUca dryag Wheeler, sa^neBting a poBslble cfmfnsiQD of the nnm- 
bera. No other inaects were observed except the omnipreeent black-flie« 
and mosquitoes. 

At the bead of the numerous fjord-like inlets along the shore there 
is usually a swamp tract extending for some distance inland in the 
same direction aa the inlet itself. The level is but little above the lake 
itself, but there is no permanent standing water or lakes as in the 
tamarack swamps. The standing water is limited to small scattered 
pools, seldom more than a meter acrose, and the forest cover is pre- 
vailingly of arbor vitae. The shade is exceedingly denee, and the 
ground is covered with tangles of underbrush and fallen logs. The 
fauna is accordingly reduced to a minimum, and the few forms col- 
lected were all dredged from the leaf-covered bottoms of the small 
pools, and included bivalve shells, Pyramidula striatella (Anth.), and 
PisidUtm subrotundum Bterk. (No. ISS), and water beetles, Scutopterua 
homii Cr. (No. 182). The latter were very scarce. 

Paunistically the arbor vitae swamp is very cloaely related to the 
later stages of the tamarack swamp, as a comparison of the species will 
show. At the ends and around the sides the swamp grades imper- 
ceptibly into the balsam-spruce forest. 

In connection with the swamps must be mentioned the fauna of the 
small rapidly flowing streams leading ont of the inland lakes. The 
bottom is usually rock or gravel, and the swift current prevents the 
accumulation of organic debris. In Benson Brook on the north side 
of Rock Harbor in stiil, deeply shaded places were dredged up (No. 149) 
Pallifera dorsaUs (Binn.), Pyramidula altemata (Say), PyramidtiUi 
striatella (Anth.), Zonitoides exiguus (Stimp.) and Physa sp. In the 
outlet from Siskowit Lake, in small pools 5-15 cm. deep witb a bottom 
of slime covered with loose pebbles, were collected several shells (No. 
238), Phyaa sp., Pisiduim media>iitm Sterki, P. subrotundum Sterki, 
and MuscuUum 8cc«rw (Prime). The cnrrent where these were col- 
lected was very slow. In the more swiftly flowing water nothing could 
be found. 

Owing to the peculiar geological structure of the Island the swamps 
have a generally oblong form with approximately parallel sides. Along 
the sides the swamps grade imperceptibly into the balsam-aprnce forest 
(Fiff. 43), and on the ends as well, though there the transition 
is more gradual and the facies are usually separated by an intermediate 
zone marked by dense thickets of alder. 

Y. The Oravel and Sand Beaches. 

The gravel beaches are found in but certain places along the shore 
(Fiy. 1), where the slope of the banks and the action of the waves permit 
the formation of the gravel deposits. Optimal conditions are found 
at the heads of the numerous inlets or coves; such as Conglomerate Bav 
(Fig. 4), and Tonkin Bay, already described, and many other similar 
places. They also occur, however, along the shore of the lake itself, 
where the wave action is at its minimum. Their distribution appears 
to be controlled principally by the slope of the bottom, since the gravel 
could not be piled up on slopes of too steep pitch, and they are almofltip 



ECOLOay OF isle HOYALB. 67 

Invariably in locationa so bounded by rot^ks or Bhore that the waves 
strike them always in one direction. An instance of this was seen 
near the light-house. A smaU inlet about 5 m. in length and width 
opened towards an island. Waves struck it in two directions, both 
diagonally, bnt rebounding from the rocks eontinned into the inlet in 
one direction. At its back was a small but typical beach, the only one 
in the immediate vicinity and likewise the only spot where the waves 
always came in the same direction. As a consequence of this directive 
action the beaches always lie at right angles to the direction 'of wave 
action. 

The gravel of which they are composed varies in size from fragments 
as large as one's flat to mere sand, but the biota of the sand beaches 
is so different that it requires separate discussion. There is no vege- 
tation,- bnt the beaches are frequently strewn with dry drift wood in 
which several kinds of fruits, dead insects and shells may be found. 
The gravel is dry on top, bnt is always moist at a depth of one or two 
decimeters or even less. The broader beaches have full exposure to 
tiie sun, hut the narrower are shaded, and all are bounded at the rear 
by a narrow but dense zone of alder. 

The fauna of these beaches is limited in species, probably o«-ing to 
the lack of food, although the number of individuals is relatively lai^. 

Caddice flies are rather common rnnning about over the finer gravel 
jast above the reach of the waves, or sometimes taking short flights 
(No. 10). Btoueflies are associated with them; they crawl abont 
Actively over the wet gravel near the water's edge and do not attempt 
to fly. They are frequently struck by waves which merely wash them 
a Mttle farther up the bank. A few species of ants are also common, 
running over and through the gravel (No. 38). They prey on dead 
-caddice flies or even on live ones when they succeed in capturing them. 
The most characteristic group, however, consists of several species of 
spiders, which are found in great abundance on the coarser gravel in 
the sun (Nos. 16, 25, .38, 39, GO), Lijcom pratensia Bmer., Pardosa 
lapidicina Emer., Pardoea groenlandioa Thor., Eho latithorax Keys. 
They run with great rapidity and at the least alarm crawl under the 
rocks, where it is almost impossible to find them. After the flrst alarm 
they usually show themselves in 10 to IB seconds, but being frightened 
again, they crawl for some distance under the gravel and are lost 
permanently. Many of them carry egg cases, and if forced to drop them 
they spin a web which they follow back in a short time. These spiders 
are very numerous, probably 10 or 12 to every square metre over all 
the gravel beaches. 

Other insects observed were, a small beetle (38) crawling over the 
■sandiest part of the beach; two spe<:ieB of small beetles (39) crawling 
through the coarse sand and fine gravel at the water's edge; a click 
lieetle, Corymbitea medianua Germ. (41) crawling over sand in a shaded 
place near a rock cliff; a Scarahaeid, Serica. vespertitm Gill, (43) ; a 
beetle, Macropogon rufipea Horn (60). Some fish worms 1 40) were 
also found buried 3 dm. deep in moist coarse sand under the gravel 
lieach in front of the light-house. They were above the level of the 
jiround water. Butterflies and wasps, which were so abundant on the 



68 MICHIGAN SURVEY, 1908. 

sand beaelies, were collected but once. The butterfly, Pyrameis cardui 
Linn. (39) flew out of the woods, rested a moment on the gravel, and 
then visited a dogwood flower. The angle wasp (il), Ammophtla sp., 
was seen flying low over a small area of sand near the water's edge on a 
gravelly beacb. 

^me fossil beaches were observed, rising several meters above the 
lake. The gravel was then thinly covered with lichens, and in some 
cases even -supported a scanty growth of flowering plants. "A beetle, 
(37) Leptura ckrysocoma Kby., was collected on a rose in such a place. 

The contents of the drift washed up on the beaches is of some inter- 
est as indicating a possible way in which new forms might reach the 
island. Here were found Limnaea atagnalis (19) ; a dead butterfly, 
Atiogia plexippus Linn.. (19) ; some dead ladybuga, Anatis ISpunctaiu. 
Oliv. (21) ; shells (21) ; butterflies (21) ; one snail shell, Polygyra alho- 
labri» (Say), badly broken but still containing part of the body (39). 
The v^etable drift (18, 21) included cones or fruits of jack pine, balsam, 
arbor vitae, and alder. 

Sand beaches are formed in the same way and under the same condi- 
tions as the gravel beaches already mentioned, but only where the wave 
action is much reduced by distance from the lake. The principal ecologi- 
cal difl'erence between the two lies in the presence of the sand, affording 
a fairly uniform surface, and a finer substratum in which various species 
may live protected from predaceoue ants and spiders. 

The principal beach studied was at the head of Conglomerate Bay, 
(Fig. ^1, and may be described in some detail. The beach was more than 
100 meters long, and divided at the middle by a small stream mnning 
through it into the bay. One portion was only 2-6 m. wide, and over- 
hung by alders. There the sand was always moist, and the fanna very 
scanty. The other portion was 10-20 m. wide, fully exposed to the sun, 
and sloping very gently back to the usual zone of alders. There was 
some drift wood scattered about over it. 

A warm sunny open place like this attracts many casual visitors from 
the neighboring woods. Three species of butterflies were especially char- 
acteristic. Papilio tumus (No. 29) was the most abundant. They flew 
back and forth along the beach at a general height of 2-3 meters, occa- 
sionally flying ont over the water and dipping into it now and then. 
They very seldom alighted on the sand. The red butterflies (No. 29) 
hovered low over the sand but when they alighted chose gi-ass or low" 
shrutm along the margin. No. 29 includes Pyrameia h/utUeri Fabr., 
Fi/rameis cardui Linn, and Basilarchia a/rthemis Dm. 

The black butterflies were not common (No. 29). They flew rapidly 
and irregularly over the sand and the edge of the water at a height of 
1-3 m. and very rarely alighted. Two other casual visitors were ob- 
served bat not caught; a redwinged grasshopper which flew over the 
sand at a height of 2 m., and dragonflies which hovered over the small 
stream. Both came from, and returned to, the woods. 

Peculiar to the beach were small blue butterflies, Phtfciodes tharos 
Drn. (No. 29), and two or three species of sand-wasps (No. 31), includ- 
ing Diodoatus n, sp,, Ammophila sp., and Santhoaariis latimamia Say, 
which flew rapidly over the surface at a height of about 1 dm. but very 
rarely alighted. When dead they were preyed upon by ants. One 

'8lc 



ECOLOGY OP ISLE ROYALB. 



or two species of flies (So. 31) (Cynomyia cadaverina Desv.) were aluo 
commoD. 

Crawling over the sand were ants (No. 30), spiders with eggs cases, 
PardoBa groetilandica TLor. (No. 3ft), and beetles, Bembidium cariniila 
Chaud. (No. 30). The latter were very numerous, and iuctuded two 
species. They ran rapidly and irregularly over the sand, and especially 
the flue gravel just back of the wet margin. When alarmed they try to 
hide under small pebbles, or sometimes fly a short distance. 

A dead shell of Limnaea stagnalis (No. 32) was found on the beach, 
and a dead Polygijra alholahrift in the small stream (32). 

VI. The EocJ: Beach. 

Where the slope of the shore is steep or the action of the waves severe, 
gravel or sand cannot accumulate, and the bare rock is left exposed. 
The ecological conditions affecting animal life here are so different from 
those of the gravel beaches that they require especial mention. 

Rising directly from the wiLter they are naturally exposed to the full 
force of the waves, (Fig. 3), which dash upon them to a considerable 
height, washing away all loose particles and effectually preventing even 
the most meager formation of soil. Beyond the reach of the waves, rains 
and drainage water act with greater or less effect in the same way. The 
vegetation is therefore limited to various species of crustaceous or 
foliaceous lichens, which are true lithophytes. Even they are absent 
from the lower portions where the wave action is more continued, and 
especially where the ice may scrape them off. Higher up the procumbent 
juniper and Cladonia appear and the whole eventually metres into the 
Cladonia clearing to be described next. Some idea of the zonal succes- 
sion of the different plants may be gained from the following table, 
showing the heights of the different zones on a rock beach near the 
Bock Ifarbor light-house, Figs. 6 and 7. 



Zone. 


HelBhi— teet. 


TDtal Height. 




7 fi. 7 m. 

4 It. a la. 

4 ft. in. 
B ft, 7 In. 
4 ft. 1 in- 

























The first two zones, to tlie height of twelve feet above the lake, are 
included here in the rock beach. Naturally these levels may vary with 
different localities, being lower in more sheltered places. 

Over the lower portion of the beach the fauna is practically without 
shelter or protection, and in the zone of foliaceous lichens shelter is 
afforded only to very minute species. There are sometimes small fis- 
sures in the rock, but only two species were observed to enter them. 
During all or part of the day the beaches are exposed to the direct rays 
of the sun, and the rock consequently reaches a temperature far above 
that ever reached bv the air. 



MICHIGAN SURVEY, 1808. 

The temperatures observed on July 11 may be given as an example. 



Time. 


AJr»t4n.lnsun. 


Rock mirlace. 


Rock. 




51 ' Fhr- 
68" ■ 
66° " 
6«- • 


eS<Bua) 
81 Caluwie) 








, ^ 




2 in 


M(sun) 









Tbe absence of plant growth also tends to limit the number and char- 
acter of species to predatory forms, and the number of individoals is 
small. 

A small rock beach jntted into the lake near the light-house, antf 
was at most but one meter higb. Although sheltered from the waves 
by an island, it was still completely flooded by even moderate waves^ 
Most of the surface was accordingly without vegetation, but besides 
tbe cnistaceous lichens there was one species of moss, a few plants 
of harebell, and several tufts of gross. Five species of insects were found 
on this beach, four of which were merely casual visitors. Borne spiders 
(No. 46), Pardoaa groenlandica Thor., wandered upon the rock from the 
neighboring gravel beach, but finding.no rocks to hide under they soon 
left. Ants {No. 46), Formica dryaa Wheeler, were rather common, but 
it was easy to see that they came from, and returned to, the gravel 
beach. The only food they obtained appeared to be the remains of dead 
caddice flies. A species of fly, Eydrophorus philombrius Wheeler (No. 
46), was very common on those parts of the rock which were constantly 
wet by the waves- They were seldom seen over the dry portions, but re- 
mained resting on the wet rocks. This fly was of common occ.urrence 
in the uplands and will be mentioned also under other headings. A 
few stoneflies (No. 46) were found on the wet rocks where the waves 
struck. The only species confined to the beach was one species of beetle, 
Bembidmm grapei, which ran over the snr&ce, hiding from time to time 
in tufts of moss. 

On a smaller rock beach exposed to the full force of the waves were 
collected a spider (No. 47) and an ant, Formica dryas Wheeler (No. 
47) ; a butterfly (Xo. 471, Ba-ailarchia arthemis Dru., was also taken while 
hovering over the beach. 

On a larger beach near by, the elevations of which were given in a 
preceding paragraph, the fauna was better developed. A jumping sptd^ 
was fairly abundant, and was a fine example of protective coloration, 
l)eing almost invisible against the gray rock backgi-ound. Another spider 
(No. 48) and red mites (No. 48) hid under the foliaceous lichens. A small 
beetle (No. 48) was abundant, running rapidly over the rock, never 
attempting to fly, but hiding in the crevices. A brightly colored red 
and black beetle was common. It ran rather slowly but flew easily. No 
ants were seen. Besides the forms just mentioned, which may be con- 
sidered normal members of the rock beach association, there was collected 
a caddice fly (No. 481 and a running spider (So. 48), undoubtedly a 
straggler from the Oladooia zone above. 



ECOLOGT OP ISLB ROTALEL 71 

At other times were collected on rock beaches ante, (No. 15) Cam- 
ponotua hercvleanua L., carrying away dead eaddice flies, and as acci- 
dental %'i8itors a CitniCT americana I-each (No. 106), a butterfly 
(Xo. 107), Basilarchia artkemis Dm., and a running spider (No. 103), 
Lycoaa pratengia Eiaer. 

In connection with the rock beaches may be mentioned the beach 
pools {Fig. 5), which are depressiODs in the rock filled with water by high 
waves. They are naturally most abundant on flat or gently sloping 
beaches, and their permanency varies with their size and depth, affecting 
evaporation, and with their height above the lake, affecting the frequency 
with which they are filled. In those which are permanent are found 
shells, lAmnaea emarginata Say (No. 58), and Planorhis parvus Say 
(No. 59), and a few insects, Rhantua binotatu$ Harr. and Corixa sp. 
(73, 74, 75). The water beetles and water boatman are strongly atereo- 
tropic, staying on the bottom or in, crevices, and leaving it only to dart 
qaickly to the surface for air. 

VII. The Cladonia Clearing and Jack Pine Ridges. 

The elevated position of the rock ridges and their physiographic rela- 
tion to the uplands are the two chief factors determining the succession 
of biota upon them. In reaponse to the rapidity of drainage, and the 
slowness of soil formation the first plant life to invade the rock beaches 
is a lichen association composed to a large extent of Cladonia rangifertTia, 
which carpets the rock to a thickness of 1 to 3 dm. With it are associated 
various xerophilouB shrubs and herbs, but no trees. Consequently the 
insolation is strong, and after rains that water not removed by mirface 
drainage is soon evaporated. The soil consists only of those thin de- 
posits formed by the disintegration of the underlying rock and the 
decay of the vegetation, and is held in place by the tufts of lichens. 
Such natural clearings in the forest are frequent near the lake (Figs. 
6, 7, 9), either on gentle slopes but little above the lake and consequently 
of late origin, or upon the elevated rock ridges {Fige. 8, 25, 26), where 
they are of much greater age. Their shape and size varies naturally 
with the topography. 

In these Cladonia clearings has been developed a very characteristic 
faunal association, rich in species and in individuals, and especially dis- 
tinct in the number and variety of insects. The fauna may be con- ■ 
veniently divided for discussion into three groups, aerial, terrestrial, and 
subterranean. Since the latter is the most nearly fixed in habit, it may 
be described first. 

1. Subterranean Fauna. In the shallow depressions and crevices of 
the Tovk{Figa.7,2o,26),&re thin soil deposits supporting a dense growth 
of various plants, especially the Cladonia lichens, the bearberry, and . 
dwarf juniper. Ants are frequent, running over the surface and ex- 
cavating below it, but they make their nests only in thje deeper crevices 
or under the densest growth of plants where the depth of soil is suffi- 
cient to allow them to make their excavations and to conserve the mois- 
ture supply. In the crevices they are usually 1 dm. or more below the 
sorface. Camjmnotus Jiercuieanua L. (22), Myrmica rubra L. (61), and 
Leptothorax canadensis Prov. (63) are the sjjecies generally represented. 
The nests are more frequent near the margin of the rock clearings. 



72 MICHIGAN SURVEY, 1908. 

where the eoil is better shaded. A nest of Formica sanguinea Latr. (No. 
72) vaB placed under a decaying limb, and the soil beneath it was 
largely composed of mintite fragments of rotten wood. This ant has 
two sorts of pupa cases. Another colony, Leptothorax canadeneis Prov. 
(No. 77), was also collected in Cladonia clearings. 

The lai^est species of ant (No. Q2) ,' Camponotus heroideanus L., is 
found always singly, and no nests were ever observed. 

Spiders also occur in the looser soil deposits, but most of them prob- 
ably belong to the snrfaee, such as (Ko. 71) Lycosa kochii Keys, which 
had an egg case attached, although buried under two cm. of soil. 
The largest spider, (No. 67) Coelotes sp, of which only one specimen 
was observed, is apparently entirely subterranean. It spins a pocket 
just about large enough for its own body, and when uncovered does 
not attempt to run, but buries itself in the soil or in crevices. A third 
species was a mite (No. 64), Rhyncholophut aimplesf Bks. 

Other species are found in fewer numbers, such as the flshworm (No. 
70), in soil under bearberry at a depth of 5 cm.; a shell, Zonitoitlea 
arboreus Say (No. 65); myriapods (No. 64), and a few other 
insects, including beetles, beetle lar^'ae, and one Jassid (No. 61). 

2. Terrestrial Fauna. Aside from the ants, which I have included 
in the first group, shells, spiders and grasshoppers are the most import- 
ant members of this fauna. Of the former but one species is included, 
Polygyra albolahris Bay. It was not seen alive, but their dead shells 
are abundant on nearly every Cladonia clearing as well as the drier 
forest covered ridges (Nos. 20, 33, 88, 93, 138, 145, 174, 197). The 
live ones are also found in damper places or even in swamps (No. 113). 

Spiders were numerous especially in the clumps of Cladonia, where 
they crawled ovct and under the mats, fretjuently carrying e(K cases. 
Three species were ob8er\-ed, Orwphosa brumalis Th., Pardoia atemalis 
Th., and Lpcoaa kochii Keys, (all No. 22). 

During the first part of July grasshoppers were infrequent, except the 
wingless stages, but during the last part of the month and in August they 
were extremely abundant. They are not confined to clearings with a 
copious growth of Cladonia or other vegetation, but are equally abundant 
on the most barren rock-ridges. Immature 8i>ecimens of Chloealtig coti- 
speraa Harr. (No. 22) were hopping over the lichens on July 6. 

Mature forms of Melanophm huroni Blatchl. and Circotettix verrucu- 
latmKby. (No 44, 35, 108, 131, 132), were very abundant. They fly 
well, making a clicking noise the while, and very rarely leave the sunny 
open ridge. Chloealtis cotMpersa Harr. (Nos. 143, 144) was collected 
in similar places from Prunus pcnnsylvanica, Diervilia, and Coptis tri- 
foUa. and the grasshopper Melanoplun alaakanus Scudd, (Nos. 146, 147) 
was taken on Onaphalium, Diervilia and grass. 

3, Aerial fauna. The light and warmth of the Cladonia clearing 
attracted many flying species, including the cicada, Tihiccn- rimosa 
Say, rar. (44. i08.'lll) ; bees, Mommctha. alUfrons Kby. (68), Xanthos- 
arun latimanvn Kay (68, 108), A', melanoph^a 8m. (108); the dragon- 
fiics, Acschna (No. 69), Ophiogotnphua colubrinua and Tetragoneuriri 
epinigcra Say (132); the butterflies, Papilio turnua Linn. (97), Ba^il- 
arrhia arthcmia IH'u. (97), Arj/j/njtM vtijrinn Tranier (97), B,m\ Argynnift 
atlantig Edw. (32), and hosts of blackflies, SimuJium vcniistiim Say. 



ECOLOGY OF ISLE HOYALK 73 

The butterflieH, Baailarchia arthcmis Dm. and Argynnig atlantis Edw., 
are so characteristic of thefle clearingB that we knew them by the com- 
mon Dame of "clearing" butterflies. The blackflies are abundant, and 
are prejed upon by dragonflies, probably the chief reason for the occur- 
rence of them so far from the swamps. 

Of particular interest was the small fly. Hifdrophonia philomhriiis 
Wheeler, mentioned before in connection with the rock beaches. They 
were nnmerons over all the clearings, bat they settled in especial abund- 
iince on the moist newly exposed soil which I UDCovored. It is probable 
that they do this only for the moisture or coolness, but io one case a 
number of them swarmed over the pupa case of an ant. (No. 66). 

Of especial interest was the fauna of the large complex of Oladonia 
clearings just behind the camp at Siskowit Bay {V, 3), Figs. 24, 25, 26. 
There was a uniform gentle slope from the margin of the bay back some 
distance inland, on which large areas were occnpied by the usual growth 
of Cladonia, juniper and bearberry. The whole was surrounded and 
intersected by balsam and spruce forest. 

Shells were quite rare, although a few of the usual species, Polygifra. 
albolahriK (Say) (233), were collected. 

The subterranean species of ants so common about Rock Harbor were 
not observed. TTiey were replaced by another species, Fomiica fusca 
K (223, 224, 226, 227), which built large circular flat-topped nests 
iFig. 28), a to 8 dm. in diameter, composed of earth and vegetable debris 
and covered with debris of balsam and spruce needles. Two sizes, a 
larger (223) and a smaller (224), were sometimes associated in the 
same nest. Many nests had been almost completely destroyed by the 
pileated woodpeckers. Spiders, Pardosa Btemulia Th. (No. 225). were 
frequently seen crawling over the ant's nests. Other spiders crawl over 
and through the Cladonia, dragging egg cases behind them, and crawl- 
ing into boles and crevices. 

Grasshoppers were abundant, as usual. Borne short winged nymph» 
of Melanoplus fasciatus Burnst-Walk.. (No. 208) were taken in thickets 
of Juniperua nana. They usually hide down in the juniper and will not 
jump out if frightened, but crawl down close to the ground, so that they 
are practically invisible. When once seen they can be picked up with 
the Augers. Sometimes they leave the clumps of juniper and jump or 
fl^y out over the Cladonia and rocks. These flights seldom exceed 1-2 
m. in length, but on one occasion one flew 6 m. high and disappeared 
among the balsam ti-ees. The adults of the same species (193, 201, 
208, 214), with full length of wings, fly long distances at a height of 
3-7 m. or more, making the usual clicking noise. They alight only on 
the bare rock or on short Cladonia, avoiding the other vegetation. One 
fiddling grasshopper, Camnula pcUticida Scudd. (No. 228), was also 
taken from mats of the juniper. 

Bumblebees, particularly Bombiis terricola Kby. (208), visited the 
flowers of Diervilla and ilelampyrnm. 

Other bees, including Taithredopais ncbrlloidrs MrOHl, Coelioxya 
moesta Cr., Xanthosariis nn'lanophca Sm.. and A', latimanua Say, visited 
the same plants. 

A small carabid beetle. Carabua aerraiua Kby. (No. 208), crawls over 
and through the t'ladonia, foraging. Leptura chryaocoma Kby. (208) 
was taktin in the same locality. Oi^jlc 

10 ~ " 6 



74 MICHIGAN SURVEY. 1908. 

The yellow clearing butterfly, Bas'ilarchia arthemia Dpu. (208), is 
■very common, flying in reRiiInr paths up and down the clearing at a 
height of about one meter, sometimes alighting on the ground and Bome- 
timeH on the flowers of Opvlaster. 

IJrocerus favicornia Fabr. and V. flavipennig Kby. (208, 209, 228) 
were specially common. They fly low, usually 2-3 feet above the 
■ground with a moderate but uniform velocity. They are searching for 
balsam trees in which they deposit their eggs, and were sometimes taken 
crawling over the trunks. 

-A small brown wasp flies low over the ground like an asilid. 

Asilid flies, Aailus annulattis Will. (208), fly low, 1-2 ft. above the 
-ground, alight on tufts of grass or Cladonia and crawl down into it. 
It could not be determined what they were hunting. 

Tliree species were taken on the flowers of the harebell, Campanula 
rotunifi folia. They were Coelioxi/s nivrsta Cr., Xanthosanie mclan^b/x'a 
Sm. and X latimanns Say. Insects were more numerous on the flowers 
«r Opii taster, from which were collected Tenthredopain neJyclloides Mf- 
Oill, ProBopis sp., Argynnig atlantig Edw., ErisMis dimitliatus Wied., 
J'hormia terracnovae I>e8v., P. rrgtiia Meis, and ffyetodcsinia serva Meia. 

A wasp, Eutii/pus americanus Cress. (235), was found backing over 
'the ground dragging a spider, Lycosa kochH Keys. At brief intervals 
it dropped the spider and ran rapidly back and forth looking for the 
liole to which it was taking its capture. It seemed to have a general 
idea of its location, but had to crawl always exactly to it. Having found 
it, a similar searcli was begun for the spider, and then the journey was 
resumed in a direct line toward the hole. 

The typical Cladonia clearings just described were almost invariably 
on the lower ridges or gentler slopes. They were surrounded, and 
eventually entirely covered, by the balsam-spruce forest. On certain 
<if the higher or steeper ridges, there was another intermediate stage in 
which the clearings were covered with jack pine. This was due ap- 
pajently to their position; the formation of soil was slower and the 
•drainage better, so that, even with a considerable depth of soil they were 
«till too dry for balsam or spruce, and were accordingly occupied by the 
,xerophile jack pine. In general ecological conditions they were but 
little ditfereut from the treeless associations. The ground vegetation 
was, as usual, Cladonia or bearberry, and the forest cover was scarcely 
leavy enough to make much shade. But the mere presence of trees 
indicates that there was a greater deposit of the soil. Under the bear- 
berry and Cladonia, the soil was quite thin, but there were more loose 
rocks, and larger and deejter Assures, which were fllled with soil. The 
effect on the fauna was to increase the number of subterranean species 
and diminish the number' of aerial forms. 

In the soil deposits up to 5 cm. deep there is practically no animal 
life, although ants crawl over the surface. Nests of Lasius niger h. are 
mmmon in crevices and under loose stones at a depth of 1 dm, or more 
'|No8. 79. 82). A nest of Lasiiut niger L. (No. 83) was excavated under 
tand at the side of a large stone. The stone formed the roof of shallow 
"excavations where the pupae were stored, and the vertical wall of earth 
at the side was honeycombed with rounded passages 1-2 cm, high, 2-4 cm. 
liroad, .and separated by thin partitions. Under larger stones their 

D, _, i..C0CH^Ic 



ECOLOGY OP ISLE ROYALE. 7S 

nests may be built at less depth, as one of Forinica fuaca L. (No. 100> 
at a depth of 4 cm. These looae i-ocks tend to conserve the moisture 
just as do the crevices. 

Beetle lairae are rarely found, owing to the abundance of ants which 
feed upon them. Tliey occur under rocks or in the deepest soil deposit* 
where the moisture is conserved. (Kos. 80, 82, 102.) No. 102 contains, 
two species of larvae, one a Cistelid, the other Draitcrivg sp. The latter 
when collected had been captured by an ant, FoiTnica fusca L. (No- 
102). A dead beetle, Dipolataxis liberta (103), was collected under aflat 
rock. 

Spiders are abundant, especially Draaaua neglectus Keys (No. 101)> 
Cicurina arcuata Kej-s (No. 102), and Lycoga pratcnsia Emer. (103). 
The former builds a small pocket-like web 2 by 3cm. in cavities under 
rocks, at a depth of about 1 dm. Spider egg cases were frequently found 
under stones or in rotten wood (No. 102). 

Myriapods were rarely seen. They seem to have regular runaways 
excavated through the wood or soil (No. 103). A dead caterpillar was- 
also found under a rock (No. 102). 

Besides the numerous dead shells of Polygyra alholaVris Say (Nos. 23,. 
27, 81, 187) which are common on the ground, especially near dead 
1<^, others were taken below ground. They occur at a depth of 1-2 dm. 
under angular rocks, ov at a- less depth under larger flat rocks. In 
either case their presence seenii* to be controlled hy the moisture (Nos- 
81, 102). Other shells were also rather common under rocks, especially- 
flat ones at a depth of 1 dm. or less (81). This single collection in- 
Huded Pjframidvla cronkheitei anthont/i Pils., Zonitotdes arboreus Say, 
yitrea binnojana (Nise), Strobilops vtrgo (Pits,). Under angular rocka 
down to a depth of 1.5 dm. Pjfrtimittula croiikheitei anthon^/i (Pils.) and 
Zonitoidcn arboreus (Say) were foimd. There are very rarely more 
than one under each stone. Most of them were dead, and the shells 
were fiequently broken, but a few were alive. At but one place were 
they associated with a Polygyra. and in this case the Polygyra wa« 
sealed with a merabnine across the orifice and was probably still alive. 
No shells were ever found under i-ocka with ant's nests. 

One jiynping spider, Lycota pratemis Kmer, (103), was caught on a 
dead jack pine tree. dm. from the ground. 

The fly {HydrophO'rug philombrius AVheeler) already observed on 
beaches and clearings was again common. Ordinarily they fly about 
near the surface in the sunniest places, alighting on the ground or od 
low plants. As soon as any moist soil is exposed they congregate on 
it in numbers, crawling over the surface, into ant burrows, and evea 
apparently attempting to eat the ant pupae. One species of ant was, 
seen catching them. 

Among other insects were bumblebees, Bombiis sp. (23), visiting the- 
flowers of Dipri-^illa- dU-'rvilla ; grasshoppers Circotrttij" rcrriiculatug Kby.. 
(27); cicadas, Tibiccn rhnosa Sny, var. (28, 84), fi-equent in the pine 
ti-ees. 

nil. The Balsam-Spruce Foirat. 

The ultimate tendency of all plant associations on Isle Royale is: 
toward the balsa m-spnice foi-est. The succession is Hmuetimes direct, 
Hometimes indirect; sometimes rapid, as upon the smaller Cladonia 



76 MICHIGAN 8URVBT, 1908. 

clearings; Bometimefl slow, as upon the jnck-piae ridges. Just as all 
temporary plant associations are occDpie<l by definite faumis of a com- 
position largely dependent on the plant eovering, so the climax aRsocia- 
tioQ of plants is also accompanied by a definite fauna, which mnst like- 
wise be regarded as the olimax auimal association. 

The succession of the dense forest growth brings into play a number 
of new ecological factors, which are not only of the highest inxDortancci 
in controlling the animal life, but are also retroactive upon the plant 
covering itself. In all the associations heretofore described physio- 
graphic changes have been procee<ling with comparative rapiditj-. 
They may be due to wave action, drainage, elevation, rock disintegra- 
tion or soil formation as direct agents, or to changes in the soil com- 
position, soil moisture, light, or heat through the indirect agency of 
the vegetation. Torresponding to the wide diversity in physical con- 
ditions there has l>een developed a fauna of many ai)ecie8 adapted to 
many different modes of living. Through the agency of the forest 
cover the light is reduced to a constant minimum, the temperature js 
made more uniform, the soil becomes of uniform character throughout, 
and the moisture is; kept nearly constant. Indirectly the ditfusie light 
is normally too weak to allow the growth of a ground cover of herba- 
ceous plants BO that the variety of food supply is reduced. In short, 
the change in from lipferc^eneity of ecological conditions lo homo- 
geneity, and the number of species varies diii?ctly with the heterogeneity 
of the habitat. This is true not only for Isle Royalc, but for any 
biotic association. Here, however, the homogeneity is esjiecially 
marked, because two species alone, the balsam ftr and the white spmce, 
are dominant throughout. 

The soil in the halHam-spnice forest is a damp closely packed leaf 
mold, sometimes deep, somcliines shallow over the rocks, and com- 
posed of decaying balsiim and spruce needles, mixed with decaying 
sticks and interwoven with fungus mycelium. When the forest is not 
so dense asijen and lurch trees may be growing, and their leaves also 
mix in the mould. In such phues tliei-e may 1h' a very thin 
ground cover of Anttr miicrojtltiiUus. Liunaca amcricntia and Pyrola 
rhUtrantho: otherwise the soil is without cover. Above this rises 
the dense growth of trees, the younger ones and the lower branches 
stunted or dead from lack of sufficient light. 

The insect fauna is composed almost entirely of auhterranean species, 
all few in number, and mostly colorless. A few species of spiders are 
seen, and a minute (^ollembolau, Tomocnua mfinr Bourl. (Xo. 140). 
Two sjK'cies of niyriaidida { No. 14(1) ai-e ralher abundant in the mould, 
one other larger sj)e<'ics was (teen once (So. 140), aud an Eiichytraid 
earthworm (Xo. 140), A few sjwcies of small sheila are rarely found 
at dejilhs of alMHit 3 cm. or sometimes on the surface. They are Pi/m- 
iiiielitld HtritiUlUi (. \.iith.), ■ Zom'tuidca aiboixuH (Say), Vitna hiiiiici/ana 
iXise), and Anri/duii sp. fllO), 

There are no anls except a large hiack species which forages singly 
over the surface, ('ampoiiotnit hrrculcaiiiis Ij. (No. 140), A single 
black f'arabid, Vatthjm !/rrg(triiiiiH Say (No. 140. 2^G), also runs over 
the surface and hides under old balsam cones. 

A few spocies of flying insects occur, especially mosquitoes, and wheu 



BCOLOOT OF ISLE ROYALB. 77 

the fresh mould is turned over a few of the tnoistDre-lovitift flies. 
Hydrophorus philomhriva Wheeler, appear and rest ou the moist ex- 
posed surface. 

The trees themselves shelter a more varied population. Most of the 
Buprestids and Cerambycids caaght iu the tent proltabl.v came from 
the forest. The dead trees of balsam or spnioe are attarked b,v wood- 
boring larvae, which construct a network of chambers just between 
the wood and bark. Borne of these turn into the wood uud extend 
to the center, following a longitudinal or tangential path for most of 
the way. These holes ma,v be filled with dust part of the way, but 
the frreateet portion in empty. They arc about 3 by 5 cm. iu diameter, 
of an elliptical shape, but at the ends sometimeit widen out into cham- 
bers a couple of centimeters broad. Two species of lanae occupy these 
burrows (Xo. 205). and in one was found a small spider, A$nanrobhta 
benneiti Blk. (Xo. 205). 

Tinder the loose bark of trees which have decayed further spiders, 
Atnaurobiiis hcnnctti Blk. (Xo. 205), frequently build their webs. A 
beetle. Calalhus arhvna I-e O, (No. li'2), forages here for food, and 
in one case a shell (142) was tnlipu. A nest of Formk-n miHt]vineii 
Tjatr. (No. 78) was found in the rotten wood of a fallen tree, hut the 
ants probably foraged' over a rock clearing near by rather than in the 
forest. Tn prostrate decaying logs the ftinna is not different from 
that of the leaf mold, and the Ranie BpecicB wei-e collected. 

A number of tlie ninshrooma of the genus Ph'iimtiiti were collected 
on dead trees and they were inhabited hy large numbers of lieetles* 
(220) Tritoimi thorarira Sa\', T. nuifiv T-w.. lioh'tohiitu riiK'ticotlis Say, 
and (Irophaoia sp. 

IX. Artifivial VU'urimju. 

The clearing about the <'anipK Iwth at Rock Harbor and at Siskowit 
Hay attracteti many species of in«ectn| p.irticula Hy strong fliers, sucli 
as Hymemoptera. Lepidoptera and Diptei-a, At Itock Harbor Cow- 
parsnip, Hcifu-lfum lanatttm, introduced in the island some way, was 
in bloom and it atlnicted a number of s)>ei'iea of bees and flics. 

In a similar clearing on Ibe north Hide of Km-k Ifai-bor a number of 
shells were collected from the under side of dead logs (150). iSome 
of these were observed at no other pbu'c, Tlicy included Polygyra 
albolabri$ (Kay). Ai-anthhiiila haiita (Say). HifuUina iappaniana (C. 
B. Adams), XoniioUh-s arbonti (Say), J'l/raiiiidiila nimkheiti anthonifi 
I'ils., C'whlicoiHi lithriru (.Miill), and Vallonia costata (SIflller), A 
plant of OiHilufitcr blooming in the same clearing attracted a multitude 
of insects (14S), including the flies i'lnlychinia peltatiis .\Ieigeu, 
Sfii/rphvs urniiaUg WilJiston, HiihaiTiifihoriu cyliHdrica Say, ErixtaJis 
diinidiatus Weed and TcirmoKt'niitt tirtfiialiK Ix)ew; the bees Htilictna 
vcrmtii Lowell. Xamhomms latiiiiaiiun Say, and Jtmiibiis terricola Kby; 
the beetle Lcptura chn/socoina Kbv. and liic lepidopfcron Cuindit stitiu- 
Tiia Bd. 

X. Stniiiiifirj/. 

Prom the lake, repi-ewiiting the most primitive rbalHtiit( WiXUWlli^ 
three line* of development culminating iu the climax a8»o(;iation : tirst, 



78 MICHIGAN SURVEY, 1908. 

through the tamarack swamp and peat bog; second, through the gravel 
beach and arbor vitae swamp; third, through the rock beach and 
Oladonia cleariogB. Physiographic forces have BOine direct part iu 
causing the successive chaugea in ecological factors, but most of them 
iire due to the retroaction of the vegetation upon the habitat. The first 
stages of the series are marked by a severity of conditions which limit 
the fauna to a few well adapted species. The intermediate stages have 
generally a wide variety of conditions, leading to the development of 
a varied fauna. The most noteworthy in this respect is the fauna of 
the Cladonia clearings. The ultimate or climax stage is homogeneous 
because of the domiuance of a few species, and the fauna is again lim- 
ited to a few well adapted siwcies. 




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ECOLOGY OF ISLE ROYALE. 



THE ECOLOGICAL IHSTRIBUTIOX OF THE BIRDS OF ISLE 
ROYALE, LAIvE SUPERIOR. 

On-O M'CHEAUY, AUaifLLTUKAL EXl'EltlMi:XT STATION", GENEVA, S. Y. 
I. I S TROD I! CT I ON', 

[n this report I shall discuns the habits of the different birds and 
their relation to their environment as found apon I«le Royale. The 
different localities visited will be deBeril>ed, the birds listed an found 
in pach locality, and the details of tht-ir habitn and distribution de- 
scribed. 

On account of the limited time, I van unable to examine a ler^ 
part of the inland, but representative localities were visited, so that a 
general idea of the bird life of the island can be grained from tbis re- 
port. For example, a nninher of tamarack swamps were visited and 
(^rtain birdn were found in each of these; it therefore seems reasonable 
to infer that these birds are found in the many other tamarack swamps 
which were not visited. 

Observations were made in five different localities by members of the 
Jliiseum party, hnt only those visited by the writer will be described. 
In connection with this paper the "Annotated List of Birds" ^onld 
be consulted. These localities will be taken up in the following order: 

1. Lighthouse Peninsnla. 

2. Trail to JlcCargoe Cove. 

3. West End of Rock Harbor and Trail to Summer Lake. 

4. Siskowit Bay Region. 

II. LlOHT-noUSB MISIXSL'LA. 

This station included the land between Conglomerate Bay and Rock 
Harbor. The conditions in this small strip of country varied very much, 
and on this account it will be divide<l into a number of stations as fol- 
lows: 

1. Lake Superior and Beach (Station I. 1). 

2 Spruce and Balsam Forest {Station I, 2 and 3). 

3. Tamarack and Arbor Vitae Swamps (Station I, 4). 

4. Jack Pine Kidge (Station I, 5). 

5. Sphagnum and Spruce Bog (Station I, 6). 

6. Valley at Head of Conglomerate Bay (Station I, 1). 

J. Lake Superior and Beach {Station I, 1). 

This station inclndcd the whole of Tonkin and Conglomerate Bays 
and that portion of Lake Superior and Rock Harbor which could be 
seen from the light-bouse. The water was deep, cold and contained 
very little vegetation. The shore bordering the lake was composed of 
jagged, desolate, wave-washed rocks (Figs 2, r>), and only in the 



82 MICHIGAN SURVEY. 1908. 

most pn((ft(ed portions of the ba.VH and harbors were trees found 
growing near the edge of the water (Fig 4) . This accounts for 
the fact that no sboi-e birds or vegetable feeding water fowl were ob- 
served here. 

The birds seen in this station were as follows: Herring Gull. Loon, 
American iferganser, Hooded Mergansei-, Spotted Sandpiper, Song Spar- 
row, Myrtle Warbler, Olive-backed Thrush, Tniw and Osprey. 

The Herring Gull was the oni.v bird seeu on the water in large iniiii- 
Imtb. At almost any time of the day there wei-e fifteen or twenty in 
sight, and sometimes they t-auie in large flocks to eat the refuse thrown 
along the shore of Rock Harbor by the ftsheriiien. Seventy-seven wei-*; 
onv^ counted, and occasionally the number was gi-eater. 

A\'hen not feeding on the water they passed the time soaring in the 
air or resting on the bai'e rocks. They seemed to pi-efer soaring during 
windy weather. With the head toward the wind they would move slowly 
upward and forward foi- some time, then tuni Ruddenly and soar 
away with the wind at a rapid rate, then swing around in a graceful 
curve and again mount upward. 

. The American Merganser, T^oon and Hooded Merganser were occa- 
sionally seen on the water. On .(uly 27 and 2S a female Hooded Mer- 
ganser and six young wei-e observed. These ducklings were yet small 
and could be overtaken with a row boat, but when pui-sued they escaped 
by diving. 

Thus tt will be seen that, excepting the Gulls, water birds were scarce 
and the shore bii-ds nearly lacking. Only one shore bird, the Spotted 
Sandpijier, was seen and that was observed two or three times; this 
wap pHibabiy a migrant. The other birds seen on the shore, were the 
Crow, Myrtle Warbler, Song Sparrow, and Olive-backed Thrush. They 
occasionally <'ame from the bushes and forests to feed there. Of these - 
birds the Song Sparrow was seen the most often, and almost every 
morning could be heard singing on the small rocky islets partially covered 
with bushes, which lay just east of the light-bouse. 

2. t^itntcr. a»f! Balstnn Forest [titatioti /, 3-3). 

In this forest of spruce, balsam and birches, there were many low 
rock ridges whose tops wei-e almost destitute of soil and trees, thus 
forming a long, narrow, natural clearing of not more than two hundred 
yards in length and from thirty to sixty yards in width (Fig. 8). 
Near the light-house there wei-e five of these ridges from thirty to two 
hundred yards ap;trt: while farther to the west there were more of 
them, but they weie farther apart. 

On account of these openings in the forest, there were many birds 
here that frequented partial clearings, yet no birds that inhabit large 
tracts of cleai'ed land, except the Chipping Sparrow, which occurred 
in the small clearing at the light-house. 

The birds found under these conditions were as follows: Red-breast- 
ed Nuthatch, Chip])ing Sparrow, Nashville Warbler, Black-throated 
Blue Warbler, BlacJt -throated Green W'arbler, Chickadee, Flicker. Gold- 
en-crowned Kinglet, Bay-breasted Warbler, Crow, MjTtle Warbler, 
Sparntw Hawk, Magnolia Warbler, Wilson's Thrnsh. 01i\-e-backed 



ECOLOGY OF ISLE ROYALE. 83 

Thmsli. Pine Siskin, Purple Finrli, White- throated Spari-ow, Trpe Swal- 
low, Barn Kiiv-allow, Sharp-shiniM'd Hawk and White-winged ('i'»)H«l)ilI. 

The Purple Finch, Pine Siskin, Sparrow Hawk. Sharp-»hinned Hawk, 
Bay-breaHted Warbler. Black-throated Hlne Warbler. Tree Swallow and 
Barn Swallow were only oecasionaily seen. 

Some of the Warblers were fouinion, and it was interesting to note 
the difference in the localitieK which they frequented. The Myrtle 
Warbler was most fi-eqnently seen nenr the shore. The Magnolia Warb- 
ler frequented the small spruce and balsam treea but was not seen on 
the slioi-e. The Black -throated Oreen Warbler was always observed 
in that part of the .forest where there wei-e many hirch trees, and the 
Xashviile kept near the partial clearings, 

Xests of the Myrtle Warbler. Chickadee. Go Idea -crowned Kinglet, 
Olive-backed Thrush and Chipping Span-ow were found in this locality. 
The nest of the Myrtle Warbler was foiuid July 7, on a small jack 
pine standing near the edge of a rocky cliff, which rose perpendicularly 
from the water to a height of about twenty feet. It was composed of 
small twigs, dried grass and pine needles, and contained four young 
about a week oid. \MiiIe we wei* near, the old bird approaclied the 
nest very cautiously. It would lly from ti-ee to ti-ee until within about 
tlfty feet of the nest and then drop down near the ground and fly low 
until below the nest; when leaving it flew along the edge of the cliff. 

Oq the same day a (lOlden-crowned Kinglet was seen to take a bit 
of moss and fly into a clump nf stunted spruce tives on a rock ridge. 
The tops of the spruce were so thick and busliy that it was impossible 
to see the nest from the ground, although the tree was not more than 
twenty-flve feet high. On climbing the tree a half tinished nest was 
found built mostly of gi-een moss. By July 21 the nest was finished 
and contained eight small eggs. This beautiful mossy cup was alioat 
four in4'he» in diameter and of the same depth, but the cavity contain- 
ing the eggs was still smaller, as the wall of the nest was about two 
and a half inches thick and lined with hare fur. 

In fi-ont of the lighthouse at the edge of the bea<-li, stood a small 
fqirace about twenty feet high, on a hotizontial limb of which was the 
nest of a Chipping Sparrow, composed entirely of grass. When the 
neat was found on July 5 it contained four young that had evidently 
just hatched. 

Ad Olive-backed Thrush's nest was found July S, in a low limb of 
a spruce that stood near the shore. It was found five feet from the 
ground, composed of grass and moss, and contained three very young 
birds. The old bird would not approach while I was near the nest 
and was so shy that the true owner of the nest was difficult to determine. 
Probably more iiests would have been found had we arrived upon the 
island earlier, as many young were able to fly when we came, and several 
immature Magnolia Warblers were found at that time in the bushes 
near the light-faonse. 

Birds were more abnndant in this locality than in any other of the 
same size. Why this was true, I did not determine. 



rfbyGOOgIC 



84 MICHIGAN SURVEY, 1908. 

3. The Tamarack and Arbor Vitae Swamps (Station I, 4)- 

This almoBt impenetrable swamp of cedar and tamarack, situated 
at tbe head of Tonkin Bay, extended back abont a qnarter of a mile 
toward the southwest. To cross this swamp was difficult on account 
of the fallen trees and numerons low branches, but a rock ridge extended 
from the bay through the middle of the swamp, almost to its western 
end, .and furnished a convenient route into it. This ridge in- 
Huenced tbe bird life of the vicinity because of its different ecological 
conditions. It was bare in places, but most of it was partially covered 
with birch, spruce and balsam. 

The birds seen in this swamp habitat were as follows: Nadiville 
Warbler, Red-breasted Nuthatch, Chickadee, Black-throated Green Warb- 
ler, Haven. Brown Creeper, Yellow-bellied Flycatcher, Hairy Wood- 
pecker, Winter Wren, Black-throated Bine Warbler, Flicker and Canada 
Jay. 

On July 11, nests of the Black-throated Gre«i and Nasliville Warblers 
were found on the north slope of the ridge within twenty-five yards 
of each other. The nests of the Black-throated Qreen was in a cedar tree 
about twenty feet from the ground. It was composed of grass, moss 
and twigs and contained young. The nest of the Xashville Warbler 
was in a! cavity in a thick bed of moss wbich covered the face of a 
small cliff five or six feet high. Here, in a soft nest composed of lichens 
and lined with grass, were found five young in the down. 

These two birds acted very differently when one was near their nest. 
The Black -throated Oreen would come within less than ten feet of the 
observer and scold while moving restlessly about among the branches. 
The ^Nashville Warbler was not as bold, for it remained up in the tree 
tops. It would hop on a branch, turn around a few times, turn anxious- 
ly toward the nest and then repeat the performance ; but it never uttered 
a sound. Here was one of tbe difficulties in judging what localities 
birds preferred. These two birds nested on the slope of a rock ridge and 
fed in a cedar and tamarack swamp. To which did they belong? How- 
ever, judging from other observations, I would say that if the natnral 
clearing had not been here the N,ashville Warbler would not have been 
found, while the Black-throated Green might have been. 

The Black-throated Green, Black-throated Blue and Nashville Warb- 
lers, Chickadee, and Red-breasted Nuthatch were nearly always found 
in this swamp, and these were in the more open parts where the trees 
were not so close together. I visited the thickest part of tbe swamp 
many times without seeing a single bird. 

4. Jack Pine Itidgc (Station I, 5). 

Tills habitat was on the north side of Conglomerate Bay and composed 
a portion of the south side and the top of a hill abont 100 feet high. 
The aide of the hill was dry and rocky, and was partially covered' with 
scattered aspens and clumps of jack pines (Fig. 13). Where there 
were no trees the ground was partially covered with mosses, lichens, 
bearberries, golden i^ods. etc. The top of the hill was bare rock with 
jack pines and a few plants growing in the crevices. Occasionally there 
was a small gully with other tiees growing in it. 



ECOLOGY OF ISLE ROYALE. 85 

On accoant of the desolate character of this locality few birds were 
found here. A Cedar Waxwiog's nest containing five eggB woe found 
Jaly 10. Juncoe and White-throated Sparroivs were occasionally heard 
singing among the jack pines. 

5. Sphagnum and Spruce Boy, (Station I, 6). 

This small bog, sltnated on top of the hill north of Conglomerate 
Bay, was covered witb sphagnum moss and hnrites with sev^al black 
spmce trees scattered over it. There were also several tamaracks an*! 
spruce at the edge of the bog. (Fiff- M), The birds seen here were: 
O olden -crowned Kinglet, White- throated Sparrow, Cedar Waxwing, ami 
Black-throated Gi-een Warbler. The Oven Bird and Wilson's Thrusli 
were heard in the forest near by. 

6. Valley at Head of 'Conglomerate Bag [Vicinity of Station f, 1). 

This location included the alders and the partial clearing at the 
mouth of the brook that emptied into the head of CoDglomerate Ba^r. 
Hie partial clearing, evidently diie to fire, as blackened logs were still 
lying aronnd on the ground, was covered with weeds, raspberry bushes, 
dogwoods and clumps of small birches. 

The birds seen here were : White-throated Sparrow, Canadian Warb- 
ler, Kedatart, Flicker, Winter Wren, Chickadee, Nashville Warbler, Mag- 
nolia Warbler, Olive-mded Flycatcher, Olive-liacked Thrush, Sparrow 
Hawk and Cedar Wazwing. The Bedstavt and Magnolia Warbler swined 
to be restricted to certain parts of this locality. The Redstart was always 
seen among the alders, while the Magnolia Warbler kept among a patch 
of evergreens at the foot of the hill on the north side of the habitat. 

III. TBIAI. TO MCr.\ltGOE COVE. 

This station included the country along the trail which ran from 
Bock Harbor to MrCargoe Cove. This trail started on the north aide 
of the harbor at the mouth of Benson Brook which it followed nearly 
to Lake Benson, then it crossed the hills to Sargent Lake and from there 
it went to McCai-goe Cove. As I did not make any observations north 
of the Greenstone Ridge, I will only describe that portion of the country 
between Rock Harbor and the top of the Ridge. In this portion there 
were several different oonditionR which will be descnbed in the follow- 
ing order! 

1. Ransom riearing (Station II, 1). 

2. Benson Brook (Station II, 1). 

3. Spruce and Tamarack Swamps (Station II, 2 and 5). 

4. Rock Ridge Clearings (Station II, 3). 

/. Ransom Clearing {Station II, J). 

This small clearing on the lowland at the mouth of Benson Brook 
was covered with grass and large clumps of alders, birches and aspens. 
These btishes scattered through the clearing formed an excellent habitat 
for birds, and, although the clearing was small, thirteen species were 
ohserved liei-e. They were as follows: Black-billed Cuckoo, Canada Jay, 



ivCoogIc 



86 MICHIGAN SURVEY, 1908. 

Soug Spnrrow. Aider Flycatt-lier, White-throated Sparrow, Redstart, Red- 
I'jed Vireo, ('edar Waxwiug, Wiltion'B Thrush, Olive-backed Thrush, 
Sparrow Hawk, Purple Finch and Pine Siskin. 

Kvery time thiH stntioD was visited there were one or two Alder 
Flycatchers aniou^ the aldev bushes, sometitaes on top of the highest 
busli and soinefiuies near the ground. They seemed to be always on 
the lookout for insecfH, and every few minutes tliey would fly several 
feet into the air and a snap of the bill told that some insect had been 
caught. They could often be located by their "pep" of alarm, and in 
the morning 1 fretpiently lieard them sing a short song. 

The Redstart and Nashville Warbler wei-e often seen among the alders 
also. Both were always on tiie move. The Redstai't kept flitting from 
branch to branch, only pausing an instant at each one to look for 
insects, while the Xashville Warbler would light on a limb and start 
to hop towurd the tup, looking an instant at each leaf us it passed. 

3. Bvmon Brook iStation II. 1). 

The londitions along this little brook are difficult to describe in a 
general way because tliey wei-e so diverse; every few rods there was a 
change. The little stream meandei-ed throngh dense forests of cetlar. 
spruci' and bir<h; through thickets of aldei-s, dogwoods and small 
Hiaplew: I'ushed thi'ough nm-row i-avines between bai'e topped ridge«, 
over i-ocks, thi-oHgli forests of birch and Jisi>en until it Anally reached 
the hiirbor at Ransom clearing. 

The birds found along this bi-ook were the M'hite- throated Sparrow, 
Redstart, Winter Wren, Red('^■ed Vii-eo, Cedar Waxwing, Oven Bird, 
Sparrow Hawk, Wilson's Thrush. Olive-backed Thrush, Blue Jay, Canada 
Jay. Crow, l'urj)le Finch, Sharp-tailed Grouse, GrinnelTs Water Thrush, 
Flicker, Magnolia Warbler, Hairy Woodpecker, Njishville Warbler, Red- 
breasted Nuthatch. Golden -crowned Kinglet and Chickadee. The Spar- 
row Hawk, Blue Jay, Flicker. Sharp-tailed Grouse, Cedar Waxwing and 
Purple Finch were seen more often in the clearings where there were 
berries, grasshoitpers and other insects. The Winter Wren and \\'ater 
Tlirush were always seen near the brook. The former frequented places 
where the iindei'gi-owth was thick. It was often observed flying aloD}^ 
the brook aud stopping every few yards to look under the leaves and 
logs for insects, and one was shot with a spider {Amaurohius hennetti 
BIk.) and two mosquitos in its mouth. Sometimes this shy bird would 
venture away from its damp retreat, perch upon the top of a tree 
and pour forth a melody that rivalled any song heard in these woods. 

The Oven Bii-d and Hed-eyed Vireo were nearly always found among 
the birches and asjtens. Tlie former very frequently was flushed from 
among the honeysuckle bushes on the ground, but the Vireo was always 
in the trees. The Magnolia Warbler, Kedbreasted Xuthatch, and Golden- 
crowned Kinglet were always seen in that part of the forest where there 
were several sjiiuce or cedar ti'ees. 

A large number of different species of birds was observed in this 
habitat, but that was because it was so large. In reality the country 
was rather deKolate, for with the excepti<m of some damp places along 
the brook, the original forest has all been bui-nt off and was only partial- 
ly replace«l by a Mccond growth of birch and as|>en. 



rfbyGOOgIC 



ECOLOGY OF ISLE ROYALE. g7 

3. Tamarack an<l Spruer Swampi* (Station TI. 2 and 5). 

About n quarter of a niilo north of Itennon Brook tlieif wa« a swamp 
similar to I, 5, exiei)t that it was larger and had more Bpruce and 
tamarack trees scattered through it. Tlie jrround was covered with 
Bphaf^um. Labrador tesi, pitcher plants, ett- hut appai-ently nothing 
that would attract birds ext-ept the trees. 

The birds seen hei-e were the Red-breasted Nuthatch. Marsh Hawk. 
Juneo, Canada Jay, Black- throated ftreen Warbler. Blaek -throated Blue 
Warbler, Thickadee, Golden-crowned Kinglet, White-winged Crossbill. 
Yellow-bellied Fl.vcatther, and tt'bite-throated S|)arrow. The Juneo 
probabl.v atra.ved here from a lai^ rocky clearinfc near b,v. as onl.v 
one was seen in the swamp, but it was lieard in the clearing ever,v 
time I visited it. 

About a quarter of a mile further on toward Greenstone Kid(i;e, the 
trail crossed another swamp similar to this one, though it was somewhat 
longer. Kince tlie conditions were the same in these two places, many 
of the same birds would he exi)ected to o«'cur in eaeii, and this was the 
case as will be seen l>,v comparing the list given above with the following: 
(Hive-sided Fl.vcatcher. Red-breasted Nuthatch, Nashville Warbler, 
Tanada Jay, Chickadee, White-winged Crasshill and Golden -crowned 
Kinglet. 

Near Forbes Lake there were two other swani]>s and in these the fol- 
lowing birds were seen: White-throated Hparrow, Canada Jay, Cedar 
Waxwing, White-winged Crossbill, Ked-breasted Nuthatch. Golden-crown- 
ed Kinglet, Chickadee. Nashville Warbler and Flicker. All these were 
found in both swamps with the exception of the Nashville Warbler and 
Flicker. 

There is a niarketl similarity in the lists of birds seen in each of 
these five swamps, and live of the Bi)efie8 were fonnd in all of them. 

-i. Rock Ridge Clearings iStation If. .i): 

This habitat consists of all the rock ridges which were crossed by 
the trail after it left Benson Brook. These ridges wei-e nearly all bare 
on the top, owing to the absence of soil. They had lieen burnt over 
sevei-al years ago and the stumps that are left show that they were 
oi-ipinally almost if not entirely covei-ed with forests. The tree's that 
were found in places where there was a little soil weve almost entirely 
aspen and birch. The birds found in this habitat were the Cedar Wax- 
wing, Juneo, Bay-breasted Warbler. Mourning Warbler, Robin. White- 
throated Sparrow, Olivebacked Thrush, Sparrow Hawk and Bed-eyed 
Vireo. 

Very few birds were seen in the clejirings, probabl,i- iKK-ause the heat of 
the sun drove theni to the shade, as mnst of the birds were olwerved at 
the edge of the clearings, in places where the gmund was jiartiall.v 
covered with ti-ees. 

IV. WESTEKN KNI> OF ROCK II.MIIIOE ANn TRAIL TO SI.'.MXBK I.AKK. 

This station comprised the western end of Hock Harlwr and a por- 
tion of the adjoining land. It was divided into Ave habitats. ^ 

hyCOOgIC 



88 MICHIQAN SURVEY, 1908. 

1. Harbor (Vicinity of Station III, 2). 

2. Small Islands (Rtation III, 1). 

3. Bulrush Zoue and T>elta (Station IN. S). 

4. Trail to Sunioer I>ake (Station III, i). 

a. Bircli Forest. 

b. Birch and Coniferous Forest. 

5. Runmer I^ke (Station III, 5). 

1. The Harbor iTiomitij of Station TIT. 2). 

In thts habitat the foUowinfi: list of fish-eating birds were fonnd : 
Ijooq, American ^Merganser, Herring Gull, Kingfisher and Bald Eaglo. 

An adult American Merganser and a number of young were observed 
about the middle of July, and about a week later another adult female 
and twenty-three young were seen. Although the young birds were 
quite small they were good ewinimers, and it was impossible to get near 
them in a row boat, except by cornering them in a small bay or in the 
end of the harbor. 

The l^oon was often seen and heard here, and once seven weiT .seen 
together. Occasionally one of the flock would swim around and around 
in a circle as fast as it could, splashing the water so that it could 
be heard for at least half a mile. Tt was impossible to get near these 
birds, not even close enough to shoot them with a shot gun, for as soon 
as they thought it was dangerous they would dive, to appear after :i 
few minutes very much farther away. It is very difficult tor the Loon to 
rise from the water, as it must fly a long distance liaising its wings 
and pushing the. water with its feet before It can get into the air. 

The Eagle was seen on a tree at the edge of the water. 

2. Small IsUinilx {Station ITT, 1). 

Near the west end of the harbor there wei-e two small islands partially 
covered with stunted cedar, spruce and birch trees, where many birds 
nested. The probable reason for this was that no squirrels inhabited 
the islands. On one island tliree or four rods long were found the nests 
of four Cedar AVaxwings, two Myrtle Warblers, a White-throated Spar- 
row and a Song Sparrow, and on the other island which was somewhat 
smaller, were a number of Cedar Waxwing's nests, three containing eggs 
or young, and the remainder being empty, most of them last year's nests. 
The Waxwing's nests were from three to fifteen feet from the ground 
and were composed entirely of lichens (Vsnea). These birds do not 
get excited as do many birds when their nests are disturbed. When 
I looked into these nests I did not hear a scolding note, althougli some 
of the owners were sitting on a tree not far away. 

Four Myrtle Warbler's nests, two old and two new, «'ere found. These 
nests were placed on spruce and cedar trees, from six to ten feet from 
the ground, and were composed of small twigs and grasses with a lining 
of feathers. One nest contained small young, July 21, and the other 
contained nearly fully fledged young. The White-throated Sparrow's 
nest was made of small sticks and grasses with a 'lining composed 
entirely of grass. It was on some hushes about a foot and a half above 
the ground, and contained one egg. ^ 

lyCoogle 



ECOLOGY OF ISLE ROYALE. 89 

3. Bulrush Zone and Delta {Station. Ill, S). 

This small grasa and sedge covered mareh was too ntmall to attract 
loaoy niarali Mrds, aad a pair of Bwamp Bparrows with, two jonng, a 
pair of Kingflshera aud Song Sparrows, a Red-winged Blackbird and the 
I,esiaer Yellow Jjega were the only birds observed here. The last two 
were only observed once, and no doubt they were only stragglers here. 

This small marah was aorrounded by a forest of s[H-nce, birch and 
balsam, and here the Golden-crowned Kinglet, Magnolia Wai1>ler, Chick- 
adee and Red-breasted Suthatrh were found. 

^. Trail to Sumner Lake (Station III, 4). 

Starting from the barbor this trail first went np a hill throng^ a 
birch forest, then across a narrow cedar swamp into a birch, spmce 
and balsam forest and down the hill to Siimoer Lake. As the birds found 
in the bircb forest were not the same as those found in the birch, spmce 
and balsam forest, the habitats will be distingaished. The cedar swamp 
was too small to be of any importance, and the birds in it were nearly 
the same as in the birch, spruce and balsam forest of which it will be 
considered a part 

a. Birch Forest. 

Judging from what had been observed before these bireh woods were 
visited, I expected to find the Oven Bird and Red-eyed Vireo, and upon 
investigation, many of both kinds were found. A family of Black-throated 
Oreen Warblers were also seen. Several Cedar Waxwings and White- 
throated Sparrows were observed along the edge of Rock Harbor near 
the trail, but they occurred almost everywhere along the edge of the 
Harbor irrespective of the kind of ti-ees. In rowing along the shore 
these birds were seen very much more often than any other. 

6. Birch and Coniferous Forest. 

This habitat was frequented by the Chickadee, .Golden-crowned King- 
let, and Red-breasted Nuthatch, the three most common birds in all the 
coniferous forests that were visited. The Winter Wren was heard 
in the cedar swamp. 

5. Sumner Lake (Station III, 5). 

This faabitat included Sumner Lake and the grassy marsh whicb sur- 
rounded it. Everywhere in the marsh the ground was soft, and the 
thick mat of grass sank under the weight of the body until the water 
poured into the shoe tops. The line dividing the grass and sedges from 
the forest was very distinct, but there were several stunted tamaracks 
and alders growing out in the marsh (Figa. 18-23). 

Many White-throated Sparrows were heard in the forest near the 
marsh, and at the foot of one of the alder bushes near the edge a nest 
was found hidden in a bunch of grass growing around the bush. Here 
in a well built nest of grass were two nearly Sedged young (July 18). 
Od the same day another nest of this bird was found on the other side 
of the lake, in a position similar to the one described above, but instead 
12 



W ■ MICHIGAN SURVEY. 1908. 

of young it contained four bluiah white eggs densely and irregularly 
viiri^ated with brown. Out in tlie marsh a Bittern was fliished from 
the grasB, and near by a deserted nest containing a bad e^ and the 
bones of two young was found. Tliis neet was only a depression in the 
tangled mat of giiiss in which it was situated. 

Two IjOons were seen on the I^ke ninny times, and these two birds 
were much tamer than Iioons usually are. for they swam very close to 
the bank whei-e I was standing. As soon as they saw nie one of them 
gave a weird and rapid "ha ! Iia I ha !'' and on being imitated it would 
i*eply evei-^' time. A Hooded Merftnnser, another fish-eating bird, was 
als(» observed here. 

V. 81BK<IWIT HAV RERIOK. 

When I arrived here in August the breeding seastm was practically 
over, Many young birds could Hy almost as well as the adults, and 
families were roving about the forests. Kandpipers were probably mi- 
grating then, and althnngh many were seen here it cannot be said that 
they bred. In two weeks other birds began to come from the north 
in large flocks, so that most observations were on habits of birds during 
migrntiou. 

Another evidence that the breeding season was over was .the decrease 
in the amount of singing. Tliis was flrat noticed on July 20, and in 
the next few days some species were heard for the last time. The follow- 
ing is a list of birds with the last date upon which they wete heard 
singing: Nashville Warbler. July 24; Myrtle Warbler and OIive-backe<l 
Thrush, .tuly 25; Wilson's Thi-ush, July 26; Magnolia Warbler, Klack 
and White Warbler and Redstart, Aug. 4; Winter Wren, Aug. 8. 

Although birds are more apt to be found in all kinds of conditions 
during migretion, yet many of them showed a preference for certain 
localities, so the localities in which the birds were seen will be given. 
Tliis station has been subdivided into the following habitats: 

1. Siskowit Bav and Shore (Htation V, 1). 

2. Trail to Siskowit Ijake (Station V, 4). 

3. Siskowit Lake (Station V. 6 and vicinity). 

4. Burning West of Outlet to Siskowit Lake (Station V, vicinitv 
of !l.). 

5. Ixjng and Menagerie Islands (Station Y, 10). 

i. SisJMicit Bay and Shore (Station V. 1). 

The conditions at this place were about the same as those at Rock 
Harbor, and almost the same species of birds were seen. Those seen 
here were: Herring Chill, Loon, Sciiiii) Duck, Solitary Sandpiper. 
Spotted Sandpii)er, Kingfisher, American Merganser and Osprey. 

J cannot say with any certainty how many of tliese birds bred in 
this vicinity, but the <iull and Merganser did. as u female Merganser 
with a flock of very small young was seen several times, and the Her- 
ring Gulls bred on the Islands south of the bay. The Ijooh. KingQsber 
and Spotted Sandpiper were observed nearly every day. The Solitary- 
Sandpiper was seen only once, (ui August lij. 

On August S four young Gulls were obtained from a flsheruian, imd. 



KCOLOQY OF ISLE ROYALE. gi 

we had an ojiportHniti' to stmly the habits of these birds. One was 
uearly full-gfown, while the other three were just gettiQg their winj; 
feathers. Ail were quite tume and tlie oldest would eat from the hand 
iind allow itself to be picked up: We were surprised to find how ch'au 
tlie«e younp (lulls were, for tlie nenta were as flifhy as those of the 
domestic Pigeon. They all seem very fond of bathing, and the largeNt 
one took a Imth several times a day. It 'nould swim ont into the bay. 
splash water over itself with its iiead and wings, dip itn head under 
water, then xhake itself; after rejwaling these perfonnanres sevenil 
times it would <-oine to the shoi-e, flap tlie wings and jump as if try- 
ing to fly. They were very particular abont keeping their bills cieiiu, 
for after eating they would walk to the water, immerse the bill and 
shake the head. 

2. Troi! to SinI:oirit Lair (Station T. .J). 

This habitat ini-iuded all the forest along the trail In'tween Siskowit 
Ray and Siskowit I^ke. If it had l)een in the breeding season it 
might have l)een divided into two or three different habitats, but the 
migrating birdu did not seem to show any preference for a particular 
forest. 

The birds seen at this station were ns follows: (Toldenci-owned 
Kinglet, rhickadee, Raven, Tigeon Hawk. Winter Wi'eu, Red-breasted 
Nuthatch, Bay-breasted tt'arbler. Red-eyed ^'ireo. Hairy Woodpecker. 
Magnolia Warbler, Black throated (ii-een Warbler, Brown Creeper. 
While-throated K|»aiTow, Tennessee AVarbler, Flicker, Canada Jay, 
.Tunco, Blue Jay. Pileated Woodpecker, Nashville Warbler, Sparrow 
Hawk, Chippiug Hparrow, (Siinnell's Water Thi-ush, I'urple Finch. Pine 
Grosbeak, Hharp-shinned Hawk. Myi-tle Warbler, Black-throated Blue 
'\\'arbler, Olive-backed Thrush, l>owuy Woodpecker, Yel low- liel lied Fly- 
catcher, and Cape May Warbler. 

The nests of only two hirds were fouud here, the ('hi<'kadee and Gold- 
en-crowned Kinglet. The Chickadee's nest was in a dead birch tree 
about ten feet from the groinid, and contained four young which were 
able to leave the nest August 11. The Kinglet's nest was in a spruce 
Tree about thirty feet from the ground. Both old birds wei-e obsened 
carrying insects into the tree, but the top was so thick that the nest 
could not be seen from the ground. On .Vngust 11 the young birds 
were still in the nest. 

The Nashville Warbler. Olive-backed Thrush, Junco. White-throated 
Sparrow and ('hipping Hjjarrow frequented partial <'learjngs or clear- 
ings in the breeding season but were found in the forests in the second 
week in August. On August 11 a flock of bii-ds were seen feeding in 
the top of a tall tamarai'k. They wei-e mistaken for warblers but on 
shooting one to identify it. it was found to be a Chipping Sparrow. 

.?. Kiskoirit Lair (Stntiim Y. C). 

This Lake was six miles long and about two miles wide at the widest 
part. The shores were mostly rocky, aud trees grew down nearly 
to the waters edge. The birds found hci'e were: Herring (Jull, Ospi-ey, 
Eagle, Spotted Sandpiper. American Jlerganser, Loon, and Kingfisher. 



lyCoogle 



92 MICHIGAN SURVEY, ISOS. 

The Song Sparrow and Grinnel's Water Thrush were also seen along 
the shore. 

Tile American Merganser, Ijoon. nnd Eagle nested in the neighborhood. 
Three diffei-ent families of SiergauHerB n-ere seen on the lake. One consist- 
ed of a female and three young, but I did not get close eooagh to the other 
tw« flocks to count them. When first ob8er\'ed, these two flocks were to- 
gether, but they sepaiuted when we rowed toward them. Two younsj 
Loons in the down were seen August JO. An Eagle's nest composeil 
of sticks was found -about 125 yards north of the lake, on top of a 
dead pine which was at least sixty feet high. The nest was four feet 
in diameter, and contained one young bird nearly ready to Sy. 

-!. Burning West of Outlet to Shhoidt jAike (Station V, 9). 

Here the original forest had all been burnt away and was only 
partially replaced by a seccoid growth of birch, mountain ash, aspen, 
wild cherry, June berry, and northern maple. Between the trees the 
ground was covered wi^ grass, currants, fire weed and other plants. 

The stream that formed the outlet of Siskowit Lake formed the east- 
em boundai'v of the burning. The birds found in this partial clear- 
ing were as follows: Purple Finch, Cedar Waxwing, Hawk Owl, White- 
thi-oated Sparrow, Chickadee, Redstart, M\Ttle Warbler, Flicker, Ked- 
eyed Vireo, Black and White Warbler, Xashville Warbler, Sharp-tailed 
Grouse, Water Thrush, Olive-sided Flycatcher, Chipping Sparrow and 
Song Sparrow. 

The Hawk Owl bred some place near here, as a< young bird with only 
down ou its head was taken August 4. This owl was seen flying around 
the clearing in the middle of the day and in the bright sunlight. The 
young bird was quite tame, or rather it was ignoraoft of the ways of 
man. It flew from one dead stub to another uttering a peculiar screech 
ofi it flew. The old bird was seen about a quiarter of a mile away on the 
top of a dead tree, but was wary and flew away. 

Along the stream there were several dead trees still standing, and 
on these trees eight to ten or more Myrtle Warblers were seen 
many times. These warblers sat on the limbs and watched for flies 
like flycatchers, and every few minutes the snap of a bill sounded 
the death note of some unfortunate insect. They did not sit in one 
place as long as a flycatchet does, but on the other hand they were not 
constantly in< motion like most warblers. 

Very little can be said about the other birds that were seen here. 
The Purple Finch and Cedar Waxwing fed on the berries here, and 
a Orouse was taken with berries and grasshoppers in its crop. The 
Water Thrush was seen near the lake and stream. 

J. Long and Meimgerie Islands (Station Y, 10). 

These two long narrow rocky islands were on the soutli side of Sis- 
kowit Bay about three miles from the mainland. Long Island was 
covered with trees except for a wide belt along the shore which was 
washed clean by the waters. Menagerie Island, on which the light- 
house was situated, had very few trees on it, as the top was barely 
out of the reac^ of the waves in severe storms. 



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ECOLOGY OF ISLE ROYALE. 93 

Menagerie Island was visited twice, on August 6 and 16. The birds 
seen liere were; Song eparrow, Barn Swallow, Tree Swallow, Herring 
Cull, Spotted Sandpiper, and Humming Bird. 

The Bam Swallow huilt in the boat-houBe and under the cliffs alongc 
the Bhore. On August 16 the neetH under the cliffa contained young 
nearly ready to fly. These cup shaped homes were composed of moss 
and mud, lined with feathers, and placed on small projections of the 
rock. 

The light-house keeper, Mr. J. A. Malone, told us that the Tree Swal- 
low built in the tower; but at this time the young were probably gone 
as none were observed entering the light-honse, although many were 
flying around. 

Long Island was visited on August 6. but no obsen-atious were made 
on any birds except the Gulls, These birds nested here by the thou- 
sands. The nests were among the rocks, some being just beyond the 
reach of the waves of ordinary storms, and others back among the 
bushes. They were from one to two inches thick, and composed of 
grasses, sticks or moss, depending on which of these materials was 
found near. Most of the nests were .on the south side of the island, 
and only a few were found on the north shore. At the approach of 
the boat the young Gulls that could not fly swam out into the water 
or hid in the bushes, while the old birds flew around overhead utter- 
ing their weird notes of alarm. 

XI. Summary. 

Hiis brief review of the birds foimd in each of the habitats studied on 
Isle Royale will give an idea of the birds that should be expected to occur 
in similar habitats of the island which were not visited. Of course only 
the common birds will be mentioned, because preference cannot be de 
termined by a few observations. The habitats of this rugged and hilly 
island presented a variety of conditions. There were bays, lakes and 
harbors, with rocky shores, wave-beaten and desolate. There were 
Kwamps that were covered with sphagnum moss and low bushes with 
here and there a black spruce or tamarack tree, other swamps that were 
covered with a dense forest of cedar and tamarack. There were clear- 
ings and partial clearings, forests of birch, containing scattered bal- 
sams and spruce, and still other forests of spruce and balsam con- 
taining a few birch trees. The characteristic birds of each of these 
habitats will be discussed in the order just given. 

1. Water Birds. The water birds found on the harbors and small 
lakes were the Herring Gull, Loon, American Merganser, and Hoodeil 
Merganser. Of these birds the Herring GuH was the most abundant 
species and could always lie seen on I^ike Snperior amd quite often 
on the smaller lakes on the island. The .American Merganser pro. 
bably ranked second in abundance. The T^on wns (juitc numerous, 
and at first it seemed as if they were more abundant that the Merganser, 
but in time it became evident that the Merganser was the more nnraer- 
ouB, though much less conspicuous, as they did not make any noise, while 
the Loon is very noisy and can often be heard a mile away. The Mer- 
ganser frequented the hays, harbors and larger inland lakes. The 
Ijoon was seen very often on the larger bodies of water, but seemed to 



&4 MICHrOAN SURVEY. 1908. 

prefer tlif siiitiner laketi more Itian llie other water bii-ds. as every little 
lake cDritiiiiieil a pair of t>oons. Yowng Merpansei-s aud Gulls were 
oftjeu seen, bnt, strange as it may appear, young Loons were only seen 
once, Ait^roW 10. 

The Otjprey, Eagle and Kingfisher were also seen several times, but 
only the latter was seen around any of the smaller lalces, and it was 
not often seen. These lakes alK>iinded in small fish and would have 
Iteen a good feeding gi-ound for Kingfishers, but there were no sand 
banks aronnd Ihe small lakes where it rould have nested, and this-may 
have iK^n the reason for its absence. There were two sand banks 
along the shores of Rock Harbor, and these were used as nexting sites, 

2. t^hoiT BinlH. The Solitary aud Spotted Sandpipers were seen 
along the shore, but these were probably migrants as only oue or two 
Spotted Sandpij>ers were seen before August 1. 

Although they were not shorn birds the Tedar Waxwing, Winter 
Wren aud White-thi-oated Sparrows wei-e often seen and heard while 
i-owing along the shore. The Cedar Waxwing would sit on the tops 
of the dead trees and every few minutes would (ly out over the water 
after insects. 

Herring Oulls nested on the shores of the sniallei* islands in large 
numliers but very few nested on the main- island. There is a reason 
why tliey choose (he smaller islands instead of the mainland, and it 
. is pi'oliflbly Woause there are no minks, lynx or other oarnivors on 
these small islands. The Gull seems to place its nest on the shore at 
1-aiuloni. wilhnnt any view to protwtion oi- secret-y. and if there were 
mink or lynx about the yonng would soon all be killed by these animals. 

The Barn Swallow nested underneath the cliffs along the shore at 
Memigerie Island and at Scoviil I'oint. The Song Sparrow and Myrtle 
Warbler weiv often seen feeding on fiie shore, and both were found 
breeding near it. The Song Kparrow fi-equented the small rocky islands 
in front of the Jight-honse, one of the islands in the west end of 
Hock Harbor, and also Kansoni Plearing on the north side of the Har- 
bor. Kven in tliis clearing it was never seen far from the water. The 
M.vrtle Warbler was found breeiliug on the uortii shoi-e of Tonkin Bay, 
and on an island in the west end of Ro<k Harbor, 

S. JSirdu Fifqiifittiiig Niiftuipx. The characteristic birds of the 
tamarack-sprnoe swamps wei-e the Pedar Waxwing, Chickadee, Red- 
hreasted Xntbatch, Golden -crowned Kinglet. White-winged Croashill, 
Canada Jay. Xasbville WaHder and White-throated Spari-ow. Pro- 
baiily none of these birds were foimd here simply beeanse it was a 
swamp, for all frefjuented other localities. The White throated Spar- 
row, Cedar Waxwing, and Nashville Warbler are characteristic of par- 
tial clearings, and this was realty a partial clearing l)ecause the trees 
were SI) far apart. The Wltite-winged Crossbill, Red-bresisted Xuthatch 
and (ioldeu-crowned Kinglet are characteristitr of coniferous forests, 
and as the ti-ees in the swamp wei-e nearly all coniferous trees, this 
would tlierefoi*e'be their natural habitat. The White-winged Crossbill 
fewls on the seeds of the tamarack ti-ees, and during the first few 
weeks of July it was only seen where there were tamarack trees; dur- 
ing the latter part of July, when the seeds of the spiiu-e became more 
mature, ihey were seen many times in the .spruce ami balsam forests. 



ECOLOGY OP ISLE ROYALE. 95 

The seeds of the tamarack mature quicker than the Hpriice, hence the 
CrossblllH would prefer the tamnracka during the earlier part of the 
siiiamer. The Black-throated Green Wnrbler wan characteristic of 
forests where there were a number of large bircli ti-ees, and this bird 
was oqI.v »een in those Bwamps which had several of these trees around 
the edf^e. Indeed the only true tiwanip bird seen here was the Marsh 
Hawk, and tliat waM only seen once. 

In the thickeat part of the cedar swaiup only a few birds were seen, 
and these wei-e the Winter Wren. Ohickadee. Red-breasted Nuthatch, and 
Brown Treeper. The Canada Jay, Nashville Warbler., Black-throate<l 
Blue Warbler and Black -throated Green Warbler were seen where tlio 
trees were tall and farther apart. 

4. Rirtlx of Clearings oiirf Partial ('IrtirinffH. The characteristic 
birds of the clearings were the Ohipping Sparrow. .Tunco, White-throat- 
ed Sparrow. Flicker, Cedar Waxwing, I'lirple Finch and Sharp-tailed 
Gi-onse. 

The Cedar Waxwing and Purple Finch were often seen feeding on 
berries in the clearings, and a Flicker was observed scratching in an 
ant's nest and eating the ants. Many ants nests were fnnnd scratched 
to i)ieces. probably by these birds. 

The cliaracteristic bii-ds of the partial clearings were the White- 
throated Sparrow, Cedar Waxwing. Chickadee, Olive-backed Thrnsh, 
Wilson's Thrush and Nashville Warbler. 

5. Birds Frequenting the Forests. In the forests of birch or aspen 
the Ked-eyed Vireo and Oven Bird were quite abundant, and in many 
Bniall tracts of birch and aspens these were the only birds seen. Other 
birds seen many times in the«e f oi-ests, wei-e Wilson's Thrush. Chickadee. 
Black -throated Green Warbler and Canada Jay. The character iertic 
birds of the spruce and balsam forests were the Chickadee, Red-breasted 
Niitlintch. Golden-crowned Kinglet. Magnolia Warbler. Canada Jay 
aod Wilson's Thrush. The Magnolia Warbler seemed to prefer places 
where the ti-ees were not very high, for on the small rocky knolls which 
were covered with stunted spruce and balsam, this bird was more 
numerous than elsewhere. 



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ECOLOGY OF ISLE HOYALE. 



THE FALL MIGRATION OF BIRDS AT WASHINGTON HARBOR, 
ISLE ROYALE, IN 1905. 

Br MAX MINOR PEET. 
I. INTBODCCTIOX. 

Oar observations of the fall migrations of birds at Washington Har- 
bor extended over the period from Angnst 18 to September 22. A 
basty examination was made of the bird life here before migration had 
really set in (August 5 to 8) , and the obsen-ations gathei-ed at this time, 
together vith the records obtained the previous rear, gave us nn insight 
into the conditions existing there. This was important, as migration 
had commenced while we were still at Siskowit Bay. 

Isle Roj-ale is situated about fifteen miles from the north shore of 
I^ke Superior, and ties nearly northeast by southwest. Sitiiatn) as 
it is several miles from the north shore and with an unbroken stretch 
of water 100 miles across lying south of it, the island iiuikes an ex- 
cellent point for the migrants to stop before crossing the lake. The 
birds seemed to center at Washington Harbor as if focussed there from 
the north shore, and in all probability the birds observed there repre- 
sented the avian life of many square miles on the mainland. Records 
were kept of the species seen each day and are given in tabulated form 
at the end of this paper. 

II. THE ESVIRONMEXT. 

1. The Clearing. Under the head of clearings, we include the thi-ee 
artificial clearings and the nari-ow roads conuectiug them. The first of 
these was situated on the sliore of Washington Harbor, near its head 
and close to the mouth of Washington River. The trees bad been en- 
tirely cleared away over an area of several acres, making a rectangular 
clearing which had been seeded to timothy and short grasses. The waves 
had cut away the soil along the shore leaving a nearly vertical bank 
two or three feet high, in some places overlianging the water. The 
land gradually rises from the water's edge, more rapidly at tiie south- 
ern end where a low bluff is formed. On this bluff the <'lub-house 
stands, and below, nearer the lake, is a little group of four small 
houses, the largest of wtiich ^ve used as a. camp. Other buildings were 
also located in this clearing. Part of the clearing was overgrown witli 
brush and small trees. These had been burned ami tlie debris left 
where it fell. Many small bushes, weeds, and vines sprang up among 
the fallen logs and branches, forming on ideal retreat for the smaller 
birds such as warblers and sparrows. The rank growth of the vegeta- 
tion made it almiost impossible to penetrate any distance into it. Here 
the Lincoln Sparrows were most abundant during their migration. 
Asthe soil was very shallow, the timothy grew short and scattering, 
and probably furnished little protection for the birds, as it was cut 



98 MICHIGAN SURVEY. 1908. 

about tlie middle of August. Near tlie road leading to the second 
clearing to the north vaA a siiiall upot cleared for a frarden. This 
bai-e ground was tJie favorite feeding place of the Horned Larks. On 
the i«hor1, steep slope which skirted the road to the second vtenring. 
thirty OP forty stnmps had l>een left. These were the favorite perches 
for the Sparrow and Hbarp-shiimed Hawks, and the tops of many 
of thf'ni were rovered with tlie harder portions of grasshoppers, these 
insects foraiiug one of the principal foods of these birds. Thre^ nar- 
row roads or trails left this clearing, one to the second clearing, one 
to the head of Siskowit Bay, and the third to I^ake Desor, The first 
of these was kept oi>eii and had originally been much wider than at 
pi-esent, being uarmwed by a fringe of alders, birches, and small bashes 
together with young balsams and spruces. 

Tlie second clearing, consisting of 3 acres, was divided into two parts, 
a grassy tract nnd a garden in which potatoes, carrots, etc., were 
grown. Fro'ni this a i-oad ( Fig. HH) led to the third clearing, 
<'alled Wendigo. which was about the same size, and contained 
two old log houses and two or three decaying sheds. Few 
of the stumps had been removed and hawks used them as per- 
ches. The ground was ovei'gi'own with short, nearly dead grass. 
White-footed mice were abundant in these clearings after ni^tfalU 
and mnuy Northern Hares were seen along the roads just at dusk. A 
iiuri-ow road wound past the clearing and off along the base of the 
bluffs for a mile or more to several abandoned cuts made by the old 
mining company. It was along these roads, which ran approximately 
north-east and south-west, that the bulk of the migi-ants passed. Even 
during the heavy migration comparatively few birds were obsened in 
the deiine forests. aJtbougb many passed along the river. It has been 
generally noticed that many birds, the smaller migrants in particular, 
as the spajTOWR and warblers, prefer the borders of clearings, and a 
long narrow road through heavy timber and bordered by bushes and 
small trees, appeared to be an ideal place for them. All the clearings 
were surrounded by the dense, coniferous forest except the first which 
fi-onted on the lake. 

2. Th/" Forent. The forest may be considered to consist of all that 
portion which has not been entirely cleared of the native trees. The 
major part consists of balsam and spruce with a heavj- undergrowth 
of ground hemlock, and in places along the river there are dense 
thickets of alder. The soil in the depi-essions is damp, with small poolfl 
of water standing on the decayed leaws. Waaliington River f^ceKS 
through the lower portion of the forest. It is a stream sixty or seventy 
feet wide near its mouth, but it rapidly diminishes in size, so that 
near Wendigo is not more than five or six feet across. Howe^-er it 
l>e4-onies much iiioi-e rapid and the Imnks are cox'wed with refuse ami 
fallen logs and hramhes. 

Few resident birds were found in the dense, dark forest, and still 
fpwei' migmnts wei-e found there. During very severe weather tho 
Chipping Sparrow sometimes retreated to the protection of the balsams, 
but it never wandered far from the open. The l^'hite-throated Spar- 
row was quite common, breeding in the foi-est along the river, and even 
during niigration it was fonnd most abundant in the nnderbrush. The 



- BcoLoay OP isle royalb. m 

Sharp-BhiDiied aud Sparrow Hawkfi rarely remained here, except dur- 
ing the night, or in very Btormy weather. The wArbters wei-e Bcarcely 
ever found in the heavy timber, but along the more open ptirt of tlio 
river and in the aldn* thii-kets the>' were abundant. By far tlie most 
eommon warbler along the river was GrinneH's Water Thrush. This 
bird was confined aliuoet entirely to the forent, and especially to that 
portion bordering the stream where fallen logs and rubbish furnisheil 
their favorite haunt. They Reenied to lie . migrating in pail's, but no 
immature birds were seen with them. Dui-ing the Mtorniy period laMt- 
ing from September 2 to 5, the Water Thruali came out into the road 
and clearings. The Wiltton and Olive-backed Thrushes bred in tlie 
forest, but during migration they preferred the open and were only 
occasionally found in the heavy timber. The path skirting the river 
waa also a favorite route for them. The maple brush which 
tiordered the forest in many places was the favorite habitat 
of the Hermit Thmsh. This and the diminutive Winter Wren were 
sometimes met with among the very densest conifers. . Among the 
other birds occurring here were the Browii i'.i-eeper, Qolden-crowned 
Kinglet, and Ked-breasted Nuthatch. Chickadees were neai"ly always 
present. This habitat was chosen by nearly all of these birds during 
migration, probably because it furnished the right kind of food and 
excellent protection. Many <rther species were occasionally met with, 
bot they were only wanderers and no particular significance can he 
attributed to their occun-ence here. 

8. Food. The clearing afforded abundant food for nearly all spet^ies. 
The grassy meadows and dry hillsides were infested with great swarms 
of grass hop[)ei-H which rose up before one as a buzzing cloud. Nearly 
all the birds taken, among which miglit be mentioned the Rharp-shinncil 
and Sparrow Hawks. Thick-billed Redwing, Rusty Grackle. Flicker and 
Nightliawk, fed to a gi-eater or less extent on these pests, as was shown 
by an examination of their stomachs. Many other species of insei'ta 
were abundant, blackflies, deer flies, and "no-see-ums"' being at times 
almost unbearable. The I)eer Mouse was very plentiful, and also the 
Northern Hare, as many as twenty or thirty of the latter being seen 
at one time feeding in the road between the first and second clearings. 
These animals together with the large niunber of Ked Squirrels found 
along the edge of the road furnished abundant food for the owls and 
migrating hawks. Seeds were plentiful and constituted the principal 
food of the Savannah and other sparrows. Wild red raspberry buslies 
were common and these berries together with several other kinds werf^ 
greedily eaten by many of the birds, especially the Cedar W'axwings. 
A few wild flowers grew in the clearing and these were oci-asionally 
visited by the Ruby-throated Humming Rird. Insect life characteris- 
tic of coniferous forests was probably abundant because the Brown 
Creeper, Chickadee, and Oolden-crowne<l Kinglet fed here almost ex- 
clnsively; otherwise this habitat did not appear to furnish much food 
for the migrants. 

III. THE WEATHER CONDITIONS AND JIIORANTS. 

1. Wrather Cim^Jitions. Throughout the iteiiod of thirty-five days 
daring which observations were made on migratiou at Washington Har- 



100 MICHIGAN SURVEY, 1908. 

bor, a daily record was kept of the direction of the wind, temperature, 
and the general weather conditions. The barometric readings, taken 
at Port Arthur, thirtj-flve miles nfearly due north from the Harbor, are 
from the daily weather maps. A comparison of my observations on 
the weather with thosfe from Port Arthur shows that the conditions 
at the two places were much the same, so I feel safe in assuming the 
barometric pressure at the island to be approximately that recorded jnst 
tn the north. An examination of the daily weather maps for this period 
shows that the same isotherms and isobars include boiii localities. Un- 
fortunately Port Arthur is the most northern station on the daily 
weather map that could have any appreciable effect on the bird life 
(if Isle Boyale. A reference to the areas of high and low pressure in- 
dicates that conditions similar to those on the island probably prevailed 
over a large ai-ea to the north of it. 

The records for the entire thirty-five days are included in the follow- • 
ing table. The readings were made between 7 :30 and 8 A. M. Other 
readings vere made during the day and where these are of importance 
I will give them under the particular discussion upon which they bear. 
All temperature readings were in Fahrenheit. The dates of the large 
bird waves are starred. 



Dste. 


Baromcm. 


Tempegi- 

tuie, F. 


Wind. 


Sky. 


18 


2»!9 

iS:S 

30.3 

20. e 

1:1 

30 


55 

1 
1 

44 

:19 

.■■.2 

i 
1 


8. 

!"■ 

None.' 

i 

Ii: 

II 
'ft: 

iZ 

ll 

w: 












































































38;! 

30.1 
30.2 

30. 1 

ii 

.10.2 

i:: 
II 




























































20 


K:' 



2. The Bird Miffiantn. a. Warblers. Many warblers nest on the 
island, and so the first indicutiou of migration in this family was the 
tendency to flock pi-epanitory to the trip south. At first these flocks 



ECOLOGY OP ISLE ROYALB. 10! 

(nnsistf^ only of the parent birds and young, but as tlieae wandered 
about tbey were joined by other families and, impelled by the gregarious 
instinct which is so strong after the breeding Beaaon, kept together and 
formed one large flock. Beginning to feel the migratory- impulse they 
were restless and waudered about over coasiderable territory, probably 
being joined from time to time by other families and often by other 
forms, for a flock of migrating warblers is rarely composed of a single 
species, aa are the flocks of so many birds. Smalt bands of Myrtle 
Warblers were seen feeding in the balsam trees on August 18, and on 
August 19 the first flocks of Tennessee Warblers appeared. However, 
these were scattered and composed of only a few birds, mostly adults. 
On August 30 I saw the first Redstarts, and from then on different 
species were constantly making their appearance. On the 26 the bulk 
of tlie Black Poll Warblers began to arrive, only a few admits being 
seen among the hundreds which came to the clearing. It is a siguificant 
fact that, in all cases where the yoimg were not in company with the 
adults, the latter and not the former, as some have reported, preceded. 
In tlie case of the Tennessee Warblers three days elapsed before there 
was any noticeable number of young, while toward the last of 
the migration the young greatly outnumbered the adults. Throughout 
the entire migration, however, the immature Blackpolls outnumbered 
the old birds, in fact the latter were very rarely seen. Only two Black 
and White Warblers were observed, and only four small flocks of 
r- Black-throated Green Warblers. 

The principal feeding grounds were among the alders, birches and 
balsams which lined the more open parts of the road. In the narrow 
strips where the high conifers bordered the path, the bird life was 
scanty, and when these portions were enconntwed by the migrating 
warblers they were qaickly passed, often in a single flight. The ilyrtle 
Warblers were the only ones observed to linger among these large trees. 

The food of the warblers consisted largely of insects, most of it being 
gleaned from the leaves and twigs of the bushes, but some was taken on 
the wing. The o|>en area here affoi-ded a greater supply of insettn than 
the forest, aud this may possibly have played a part in the choice of 
this {inrticular habitat. 

On cold mornings, when the thermometer registered about 45° F. or 
below, the warblers would remain hidden in the dense underbrush, 
not appearing until about nine o'clock, when the sun would be quite 
warm and the usual morning fog be dispelled. This fog bung over the 
haobor nearly every morning aud frequently was so dense that Beaver 
Island, in tl|e harlKir, was almost invisible. It was often blown back over 
the clearings, and until it raised, the majority of the birds remained 
quiet. The height of the migrating movement seemed to be from the 
middle of the afternoon until nightfall; how far into the night it ex- 
tended I was unable to ascertain, but the cries of innumerable birds 
could be beard until nearly morning. These cries, usually of a single 
faint syllable, were possibly uttered to help keep the birds together. 
When the migrating flock had to cross the clearing it was a noticeable 
fact that they rarely flew directly across, thus exposing themselves to 
the attack of the numerous hawks, but instead kept near to the ground, 
making short flights from bush to bush, and where these were scattered 



102 MICHIGAN SURVEY, 1908. 

thev alighted directly on the ground. This waB especially noticeable 
in the case of the Palm Warblers, which often lingered to feed in the 
grase. 

The Sparrow and Sharp-shinned Hawka were the principal enemies 
of these birds, de\oi]rinR manr earh day. Pnring the large bird wave 
of September 12, the Pigeon Hawk also played a conspicuous, part in 
their desrtruction. The influence of the hawks will be taken up under 
the discussion of that family. 

During the first days of migration the warblers moved aloog very 
leisurely, the same flocks apparently remaining about the clearings 
all day, but toward the latter part of the season the birds hurried for- 
ward, taking their food as tliey moved along. There were several 
warbler waves or periods of great abundance, the first occurring on 
Auffnst 23. This one was made np almost entirely of Tennessee War- 
blers, adwlt and young being about equal in number. For the re- 
mainder of the migration, however, the young outnumbered the adults. 
The second, made up largely of immature Blackpolls, arrived on August 
26. The third wave, consisting principally of Palm Warblers, occurred 
on August 30. The last wave, and by far the largest, occurred 
September 12. At this time the clearings and roads were full of 
warblers, nearly every species observed at Isle Royale being represented 
to a greater or leas extent. These waves will be dealt with separately. 

During the heavy rain and wind storms of September 1, 2 and 3, 
the warbler migration was at a standstill, the birds keeping under 
cover as much as possible. The Blackpolls and Palm Warblers were 
the only species which seemed to be unaffected by the weather. These 
beautiful warblers were obsci-ved feeding in the open clearing during the 
heaviest rains, but even they did not undertake to migrate against the 
strong wind, so far as 1 could determine. 

b. Spairotcs. The sparrow migration b^an much later in 19fl5 than in 
1904, some of the most striking examples being Savannah, Aug. 17, 
White-crowned, Aug. 28 and Lincoln, Sept. 1, 1904. On August 18, 
(1005) the first day obser\'ations were made, Chipping and White-throat- 
ed Sparrows were seen, The Chipping had gathered into flocks and 
roamed about the clearings, feeding near the borders, while the White- 
throated were still in single families hunting about among the dead 
leave» in the damp underbrush. Many of the White-throated Sparrows 
were still too young to migrate, some having a little of the first down 
on them. On August 22, an immature Vesper Sparrow was taken, 
the only one found on the island. A few Song Sparrows were pre^nt 
from August 21 to 24. These were the only ones seen and were probably- 
migrating at that time. The next few days the number of both Chip- 
ping and White-throated Sparrows was materially increased, large num- 
bers of young of both species making their appearance. Very few of 
these had moulted the first plumage. ^ot until August 31 were any 
other species seen, then large Hocks of Savannah Sparrows, both young 
and adults, came to the clearings. All were in perfect fall plumage. 
It seems peculiar that none of tliese birds were seen before this date, 
because between August 5 and 8, I saw several, and obtained one im- 
mature barely able to leave the nest- The food was obtained along 
the roads, in the meadows, and about the houses, where several lost 
their lives by entering deserted rooms and not being able to find their 



ECOLOGY OP ISLE ROYALE. 103 

way out. The Sharp-Bhinned Hawks proved to be their worst enemy. 
The first ftocks of Savannah Sparrows to arrive remained for several 
diays, their numberB constiintly increasing. On September 5 many 
of the Chipping, White-throated, and Havannali Sparrows left the island, 
and for the next two days only a comparatively few were seen, then 
others came in from the north and the flocks were rapidly increased. 

It was noticeable that most of the birds which migrated from the 
island on September 5 were adults, the young remaining until a later 
time. The Savannahs showed the least fear of man during migration 
of any of the sparrows. On September 12, with the great bird wave, 
<>uine the Lincoln Sparrows. Throughont this and the next three days 
hundreds of these birds were seen. As a mie they kept secreted in 
the burned brush and weeds of the first clearing, but individuals were 
met with all along the road, where they were seen bunting among the 
fallen logs and underbrush for insects. 

Chipping Sparrows remained throughout the entire period during 
which observations were made, but probably none of the individuals 
first seen remained throughout that time. This seems the more likely 
afl on aevernl dates the bulk of the sparrows of alt species left, while 
more came in later from the north. 

c. Hawks. During a few days spent at Washington Harbor early 
in August (Aug. 5 to 8) only a few Sparrow and Sharp-shinned Hawks 
wei-e seen, but by August 18, many individuals of both species had ar- 
rived, liiese remained here to feed on the swarms of grasshoppers in- 
fecting the meadows, and on the small birds, as warblers and sparrows, 
which were easily caught in the exposed clearing. The first few days 
the Sparrow Hawks outnumbered the Sharp-shinned about 10 to 1, bat 
QK the season advanced their numbers became more equal and toward 
the last the Sharp-sbinned outnumbered the Sparrow Huwks, both 
because of a steady increase of the former and because many of the 
latter left the island for the south. When the observations were first 
made the adult Sparrow Hawks were as numerous as the immature, 
bat toward the last of September the adults had nearly all left and 
many more immature had taken their places. Some idea of their 
number may be gained from the statenient that more than thirty were 
counted at one time, sailing over the first clearing. Until the middle 
of September the immature Sharp-shinned greatly outnumbered the 
adults. These immature were full size, but did not have the spotted 
plumage of the adult. The females of both species greatly predominated. 
Toward the end of September many male Shnrp-shiDned, both im- 
mature and adult, appeared. These two species of hawks fed on grass- 
hoppers to a considerable extent, but many crops of both species were 
found filled with the remains of Tennessee, I'alm and Blackpoll War- 
blers, Savannah Span-ows and other species not determined. As a 
rule the older hawks were the ones which destroyed the birds, and this 
may account for their migration from the island at the same time that 
"the large warbler and spaiTow wave passed, while the immature hawks 
remained. Pigeon Hawks were recorded from time to time, but not 
until September V2, when the lower end of the island was suddenly 
flooded with bird life, did they apjiear in any numbers. On this date 
several flocks of (J or 8 were seen in different parts of the clearing. 



104 illCHIGAN SURVEY, 1908. 

Tliey were preying principall.1' on the eparrowe, and were creating 
gi-eat Iiavoc anioug llieni. Coming with the great wave they remained 
throughoat the day and passed on with it that night, on); one being 
seen the next morning. 

Tlie migration of the hawks is thus seen to have been intimately con- 
nected with the migration oC the smaller birds npon which they preyed, 
and seems to give at least one instance of bird migration being in- 
fluenced by tlie food supply. 

During cold, rainy niorninga the hawks rarely appeared in the open, 
usually not until about 9 o'clock. This, however, was the time the 
warblers appeared on such days, and this may also have determined 
their appearance. 

d. OiHs. Only two species, the Great Horned and Acadian Owls, 
were seen. These wei-e residents at this time and only concerned mig- 
ration in that they sometimes preyed upon the migrants. Their effect 
was. pi'obably slight. 

e. Thrushes. Six species wei*e observed migrating, the Bluebird, 
Robin, Wilson's. Olive-backed, Gray-cheeked and Hermit Thrushes. A 
pair of Blue Birds nested in a dead Birch at Wendigo, and this family 
left the island about August 22. On the 2ith a small flock probably 
consisting of two families appeared at the first clearing and remained 
about the tangled bmsh imtil .Vugiist 31. when they also left. No others 
were seen escept on Hepteuiber 11 and 12. Robins were seen twice 
during August, but on Hepteniber 6, the first real migratory mo\'e- 
ment was initiated, and from then on the number rapidly increased. 
Kmall flocks numbering a dozen or so wandered about the clearings 
and open woodland. Many disappeared on the night of September 12, 
but the number was soon replenished, and at the time the observations 
were closed the Robins were quite abundant. 

The most common of the Thrushes was the Wilson's, They bred on 
the island and showed no indications of the migrating spirit until the 
latter part of August, when they gradually increased in numbers and 
moved about to a greater extent. After September 6 they became 
rather scarce, and none were seen after the 14th. Their place was 
taken by the Olive-backed, and later the Gray-cheeked became abundant. 
Many immature Olive-backed were seen but this species had nearly 
disappeared when the great flocks of Gray-cheeked arrived (ta September 
12. They showed little fear and did not seem to be frightened at the 
report of a gun. The flocks of the Gray-cheeked were made up of 
immature and adult birds, all in perfect fall plumage. 

f. Other Birds. Following the breeding season the woodpeckers wan- 
der about the island milking what might be called a local migration. 
Perhaps some of these birds leave the island in the fall and probably 
others come in from the north. In one iustance, that of the Flicker, 
their numbers are enormously increased during the latter part of 
.\ugust and all through September. It is very improbable that any of 
these latter birds winter on tlie island. Flickers were seen every day, 
but the number gi-eatly increased toward the last of September, and 
from the report of residents the number continues to increase until late 
October when they api>ear to leave the island. Many were found dead 
without any apparent cmiMC, and it was reported that in the latter 



ECOLOGY OF ISLE ROYALE. 105 

part or October imudreds were fouud dead eaeli year. The Downy 
aod Hairy Wood]>eckers visited the Hearings occHHionaU.r, ns did the 
P ileal ed Woodpecker. 

Several {>airs of Kingflshei's frequented the river bankx, and one 
pair neftted in a sand bank, i-earing 7 younf^. These birds gradunll.v 
diBappeared, uutil on Kepteinber 15 the last OQes left. Families of 
Sedbrea&ted Xuthatohes oecasionally vinited the cleariuj^s, often ac- 
companied by Cliifkadeee, During Ahgust, Crows were commonly 
seen, but by the latter part of September they bad entirely disappeared, 
n'hether to the south or not it was impossible to determine. Se\eriii 
species of flycatchers and vireos were seen migrating, the Alder, Green- 
crested, and Least Flycatchers being seen several times, while only out- 
Yellow-bellied was found. Both young and adults of the I^ost Fly- 
catcher were seen, usually tc^ther. 

One pair of Chimney Swifts was obsen'ed circling over the river on 
An^st 19. Whether these were migrating I do not know, but tliey 
■were the only ones noted here in IJKta, One of the most conspicuous 
species during August was the Thick hilled Redwing Blackbird which 
came to the cleai'ing in flocks numbering from about 30 to E50. Flocks 
composed of young and adults arrived nearly every day. The propor- 
tion between the two seemed to be about equal, or if anything, in favor 
of the adults. Xone were in the black breeding dress, and only a few 
males had the red on the shoulders out of the pin feather stage. The 
majority left on August 26, a few wei-e seen on August 29, 31, and Sep- 
tember 2, and two young were found on the 8th, 9th and 10th. A single 
specimen was taken September 16 and another, partially moulted, on 
the 20th. 

There were many instances where only an individual or a singlo 
flock of a certain species was seen. Among these might be mentioned 
the Catbird observed on September 12. which was the only one the ex- 
pedition noted either in the Porcupine Mountains or Isle Royale during 
bcth years, the I'hitadelphia Vireo, Blue-headed Vireo, Chimney Swift, 
Vesper Sparrow, Humming Bird, Migrant Shrike, Black-throated Bhw 
Warbler, Black-thi-oated Green and Black and White Warblers, Yellow- 
bellied Flycatcher, Red-eyed ^'ireo, UDid Kingbird. Of these, only two 
of the Philadelphia Vireo and Chimney 8wift were seen, and but single 
individuals of the Migrant Shrike, Black and White Warbler and Ruby- 
throated Huonming Bird. 

IV. LARGE BIRR W.WES. 

During the period from August 18 to September 21, six large bird 
waves passed over the island. Sometimes the waves were composed 
principally of one species, and again several species occurred in vary- 
ing numbers. These bird waves were mostly from the north, although 
small ones, consisting of the birds which had accumulated on the island, 
took place at various times. 

A bird wave may be recognized, first, by a sudden increase of indi- 
viduals, second, an increaso of species, or, third, by a sudden decrease 
in the number of birds which were residents or had gradually accumu- 
lated OD the island. During the large wave of September 12, all of 
14 



108 MICHIGAN SURTE!T, 1908. 

these evidences were preBent, but nsually only one or two were recog- 
ni?^, the most proaonnred of which was the great increaae of birds 
n« they paseed nloDg tlie ronds from one clearing to another. 

In some cases the bird wave marked the date of first arrival, at other 
times it simply marked the arrival of the bnik. The bird waves were 
p-nerally sharply defined, so that their relation to the atmospherir 
changes could be noted to the beat advantage. The bulk of the migration 
took place during the nights of bird waves, although there was a con- 
stant going and coming of certain specie* throughout the fall. Being 
almost constantly in the field the writer had excellent opportunities to 
Ktudy the migration when it was most pronounced, that is, during the 
large waves. 

1. First Wai/C. The first wave observed occurred on August 23, and 
consisted principally of Tennessee Warblers, immature and adults being 
alwmt equal In number. At 7:00 A. M., the temperature was 58°, a 
rise of 6° in the last twenty-four hours. The weather was clear, and 
it was in fact one of the most beautiful autumn days of the season. 
A light northwest wind prevailed, the barometer standing at 29.9 inches 
Mow). This wave lasted for three days. The day previous the weatlier 
conditions had been about the same, except that the thermometer stood 
at .'»2''. An area of low pressure (29.75) was advancing towards the 
iislsmd and on this date was central over Assiniboia. On the 23d this 
an-i was central over Isle Royale and the area to the north and northeast 
of it. On August 24 a low area was centered over northeastern Missouri 
while the high pressure which followed it reached very nearly to Isle 
Koyale, thus lowering the temperature to 53° with a northwest wind and 
barometer reading of 30.2. On the 25th of August the high (30.3 inches) 
included the island and the area directly south of it. The weather was 
clear, no wind, and a fall in temperature to 50°. On this date the 
beginning of the large wave of Blnckpoll Warblers comm^iced, many 
large docks appearing before nightfall. On the fourth day of the wave 
(August 26) the barometer fell to 30.2 with an increase to 54° in tem- 
perature. There was scarcely a breeze, and the day was for the most 
part humid and cloudy. The bulk of the Blackpolls arrived during the 
previous night and throughout the day. The wave lasted for se\'eral 
days, decreasing gradually in volume, so that it was difiScult to tell ex- 
actly when it stopped, if in fact it did not grade off into the next one. 

2. ■S'cro«</ Wove. On August 30 great flocks of Palm W-irblers in- 
vaded the island. On the previous day the area of low pressure was 
central over the i-egion a little to the west of Isle Boyale, with a bar- 
ometric pressure of 29.9. a temperature of 61°, and wind northeast with 
rain. On the 30th the low area had passed on to the St. Lawrence Val- 
ley and the advancing high pressure was over Manitoba. The barometer 
stood at 29.8 with a northwest wind blowing 4 miles per hour and a 
temperature of .^16". This wave also lasted three days and might have 
continued longer but for the severe gales which set in on the night of 
September 1. 

The second day of this wave (August 31) the area of high pres- 
sure (30.1) was central over all of Northern Michigjin. a considerable 
area north of it, and south to southern Wisconsin. With the high pres- 
sure came a drojt in tenii)enitui'e to 4C°. The day was clear with a north 



ECOLOGY OF ISLE ROYALE. 107 

wind blowing about 4 nii)«8 per hour. The Palm Wapblers continued to 
come in large flocks and were by far the most abundant bird at the 
Harbor. The vanguard of the migrating host of Savannah Sparrows 
appeared early on the morning of the Slst, and by evening the clearing 
was (airl.v covered with them; more came during that night and all 
the next day large flocks were arriving at the flrst clearing. There was 
no appreciable increase during the four succeeding days among the bird 
migrants. The third day of the wave (September 1) the barometer stood 
30.1 inches, temperature 52°, with a northeast wind and a cloudy (*y. 
This day marked the close of tliis wave both of birds coming to and 
leaving the island. This wave might have l)een checked either by the 
gales which follon-ed or the change of the wind from north to northeast. 
A few birds attempted to cross during these succeeding days, as many 
were killed by striking the lighted windows, etc., or were found in an 
exhausted condition. These were probably part of that steady stream of 
migrants which continues to paes south during the fall, without any 
marked wave and in general disregard of the weather conditions. 

:{. Third Wavt: Thin wave was noticed flit<t on September 5 when the 
bulk of the Chipping, White- throated, and Savannah Sparrows left the 
island, and on September G the first real migration of the Hobin com- 
menced. I have considered these two days as parts of one wave, con- 
trolled by the same conditionH, for probably the same influences acting 
at the saiiie time caoeed the sparrows to leave the island and the Robins 
to leave their more northern home. The weather conditions were suth 
as seem to be niost favorable for fall migration. On September 5 the 
high pressure had advanced to an area lying from Winnipeg on the north 
to Memphis on the south, and extending east nearly to Duluth. The 
barometer stood at 30.1 I high) with a northwest wind blowing six 
miles per hour and a temperature of 39°. The day was {tartially cloudy, 
but no rain fell. On the 6tli an area of high pressure had formed over 
the region directly to the north and northeast of the island. A moderate 
northwest wind prevailed with the bai-ometer at 30.1 inches, and ther- 
mometer 42°. and the weather was clear. A few Robins had been noted 
befoi-e this, but these were only scattered individuals or pairs, but on 
this date a large number came to the island, both youug nud adults being 
seen, although the latter greatly predominated. 

4. Fourth Wave. On September 12 the largest wave of the season 
occurred. For number of species as well as individuals it could scarcely 
be compared to the other large waves, a total of 41 species being ob- 
served in actual migration on this day. On the previous day the low 
area was central over Lake Erie, and a similar area was formed over 
the Dakotas. The barometer stood at 29.8 inches, the temperature at 
52", with a north wind and a clear sky. It was cloudy, however, on the 
north shore. On the morning of the 12th the low area was central over 
Sew England, while the western one had moved south and had been 
followed by an area of high pressure central over the Dakotas and 
Western Minnesota. The island lay between the isobars of 30.1 and 30.2 
inches, with a temperature of 42^, and a northwest wind averaging 8 
miles per hour. The day was clear and seemed i>erfect in every way. 
The birds were so plentiful in the clearing at 0:30 A, M., as to attract 
my attention from the windows. Unlike the other bird waves, the 



108 MICHIGAN SURVEY. 1908. 

motioii was continuons, scarcely a break occurring in the steady stream 
of migrants as tbey passed along the road. There was very little linger- 
ing by the way, although when the birds arrived at the first clearing 
they often scattered about, feeding on the myriads of insects infesting 
the grass and shrubbery. Many of the birds after collecting into great 
flocks, sometimes numbering more than a hundred, rose directly from 
the clearing and taking a south westerly direction, left the island, pre- 
sumably for the south shore. As a rule the birdie flew dii-ectly down the 
Harbor and the fishermen and tourists at Washington IxJiind reporte<1 
that never before had they seen such numbers of birds except during 
the spring migration. These observers reported that the majority of 
the flocks passed at a considerable height above the island, many of 
them probably belonging to the same flocks that left the clearing four 
and one-half miles ap the bay. Some -species, particularly the warblers 
aud sparrows, flew from the ground in practically the direction they 
took when on their way, but others as the American Pipit, Bna^ Qrackle, 
Horned Lark and Thick-billed Redwing flew around in great circles, 
often hanging around the border of the clearing' for some time, as if not 
quite decided whether to go or not. No flocks of the last named bird 
were seen on this day, however. As I passed up the road toward Wendi- 
go that morning I met flock after flock of Palm Warblers, Grey-cheeked 
Thrushes, and Savannah Sparrows, The pn^ressive movement of the 
migratory birds was clearly shown as they passed in a southwesterly 
direction along the road from Wendigo to the clearing at the Club 
House. 

Darting everywhere were Sbarp^hinned and Sparrow Hawks, while 
every few miniites a Pigeon Hawk would dash by. All the birds seem- 
ed restless as if impelleil by some uncontrollable spirit to keep ever on 
the move. Warblers, thrushes, sparrows and flycatchers were coDStantty 
crossing and recrossing the path in front of me. 

During the night of September 12 nearly ail the birds left the island. 
Towards evening the temperature gradually dropped, until at 9 P. M, 
it was only ZS". The morning of the 13th was one of the coldest I 
experienced on the island, ice remaining on the water pail until nearly 
noon. During the night the area of high pressure had advanced until 
at 7:00 A. M. it was central over Isle Royale with a barometer reading 
of 30.4, temperature at 26°, and a brisk west wind. The sky was clear, 
as is usual under high barometi-ic pressure. 2tfany Lincoln Sparrows 
remained, as well as numerous flocks of Gray-cheeked Thrushes. But 
the great flood of migrants had passed on the previous night. Howe^-er, 
the wave set U|» by these verj' favorable conditions was not yet over. 
Large flocks of Homed Larks numbering from about 60 to 200 or more 
came to the clearing, feeding on the injects and seeds in the open 
meadow and on the cultivated ground. On this morning many dead 
birds were found, among which was aji adult male yellow-bellied Fly- 
catcher, the only one of this species seen. 

The following is a list of the 41 migrants which composed this re- 
markable wave of September 12: Chipping Sparrow, White-throated 
Sparrow. Sharp-shinned Hawk, Sparrow Hawk, Blue Bird, Flicker. 
Myrtle Warbler, Tennessee Warbler, Phoebe, Redstart, Least Fly- 
catcher. Hermit Thrush, Spotted Sandpiper, Pigeon Hawk, Robin, Olive- 
backed Thrush, Golden-crowned Kinglet, Cooper's Hawk, Wilson's 



BCOLOQY OP ISLE ROYALK. 109 

Thmsh, Solitary Sandpiper, Slagnolia Warbler, Palni' Warbler, Oven- 
bird, Bay-breasted Warbler, Blue Headed Vireo, BlackpoU Warbler, 
SaTannah Sparrow, Black -throated Ore«i Warbler, Grinnell's IVater 
IMirtish, Marsh Hawk, Catbird, Black-throated Blue Warbler. Naah- 
Tille Warbler, Philadelphia Vireo, Bed«yed Vireo, Linoolo's Sparrow, 
Connecticut Warbler, Ruby-crowned Kinglet, Gray-cherfted Thi'ush, 
Broad-winged Hawk and White-crowned Sparrow. 

The following were seen on September 13: Chipping Sparrow, White- 
throated Sparrow, Sharp-shinned Hawk, Sparrow Hawk, Flicker, Her- 
mit Thrnsh, Pigeon Hawk, Robin, Palm Warbler, BlackpoU ^I'arbler, 
Savannah Sparrow, Marsh Hawk, Lincoln's Sparrow, Gray-cheeked 
Thrush. White-crowned Sparrow, Homed Lark and Yellow-bellied 
Flycatcher. 

5. Fifth Wave. On September 16 the bulk of the Rusty Crackles 
arrived. A few had Iteen seen the day previous, but only scattered in- 
dividuals composing the vanguard of the large, noisy flocks to follow. 
The area of highest pressure was off the Kew England states, while 
the low pressure centered in Kansas. The barometer stood 30.0 inches, 
temperatnre 50°, and an easterly wind with a cloudy sky. While this 
was one of the smallest of the sharply defined waves, it presents a 
tnarked contrast to most of the others. Although the area of highest 
pi-essure was not near Isle Royale. as during most waves, the barometer 
litood at 30.0 inches, which was higher than for the area to the west 
and south ; the wind was from the east and the sky cloudy in con- 
trast to the northwest wind and clear sky of the other waves. 

6. Sixth Wave. On September 18 the high area was far to the 
east (Maine) and the low centered over Kansas. The barometer stood 
at 29.S inches, with a clear sky, northeast wind, and a temperature of 
53°, Like the last this was comparatively a small wave and only in- 
volved a single species, the American Pipit, which came in large flocks 
numbering from perhaps 100 to 200 birds. 

V. THE BBLATION OP WEATHBll TO MIGBATIOX. 

Gooke ('88. p. 16,) makes the following statement in regard to the 
relation of temperature and barometric pressure during migration, '"The 
area of the lowest pressure is nfever stationary but con.**tantIy moving, 
and in an easterly direction. It may be moving northeast, east, south- 
east, and rarely north or south; but never northwest, west, nor south- 
west. The usual direction in the Mississippi Valley is a little Bonth 
of east." Warm wa^-es. which are associated with areas of low jires- 
sure, therefore b^n in the northwest, and move toward the southeast, 
'"It is a law of the movement of winds that they go toward areas of 
lo^' pressure, and from an area of high pressure." "But an area of 
low pressure is followed by one of high pressure, producing an opposite 
effect, and the isotherms which bent north to welcome the coming of the 
low area turn rapidly southnard l)efore the icy breath which blows 
from an area of high pressure. Thus the cold and warm waves both 
come froml the same quarter, and both move in the same direction: 
that is "the direction in which the area of low pressure is advancing."' 
It will thus be seen that the temperatnre and the direction of the wind 
over any given area are both associated with the barometric pressnpdi- 



110 MICHIGAN SURVEY, 1908. 

and the inovemeiit of its high and low arefls, and since "low preesnre 
is generally accompanied by clouds and rain, while areas of high pres- 
sure are cloudless" it will be seen that this important etement is also 
associated vrith the barometric pressure, Tims we see that the four 
striking factors which influence migration, namely teauperature, direc- 
tion of wind, condition of the weather, and barometric pressure are 
correlated and work together, the same factors being always associated 
together and giving the same resultn. 

To detei-niine the true relation of these factors to migration we 
must discover the most favorable conditions for this moveiDeDt, and 
then we can coi'i-elate the atmospheric changes which are taking place 
with the corresponding migratory moi'emenf. Of course many birds 
are constantly passing to the south throughout the fall, iiTCspecti^-e 
of the weather conditions, but the changes which will set great num- 
bers moving onward simultaneously must be the ideal conditions for 
migratiou. If this be true the time to study this relation of the weather 
is during the great waves. 



TABLE OF BIRD WAVES 








DatF. 




Tempen- 


Wind- 


Sky. 




riMT Bin 


WAVE. 






Aug. 23 


29.9 


i 


IS 


B 


















S8;f 


Is 


Vr 










CloudT. 




THIHD Bia 






Sept. -I 


30.1 

FOVBTH BIB 


« 


!i:S: 


^ly cJoudr- 






*•"■!!:;::::::::,;:::.:::;;:::::::::;:;::: 


S.i 


42 
28 


N. W. 


Clear. 

Clear, 






















Sept. 18 


29-8 


S2 


fj. E 


Cte«T. 



1. Influence of M'ind. A reference to the table of bird waves shows 
that on six days of the thirteen during which large waves were ob- 
served, the wind was from the northwest. Two diays were without 
appreciable wind, on two, the wind was from the northeast, and upon 
other days it was from the north, east, and west, but npon none of 
them was it fmiii the south, soutlieast, or southwest. A. n<H^hweet 
wind prei-ailed the first two days of the first wave, the third and fourth 
days being without wind. Tlie second wave commenced with a north- 
west wind, which changed to north on the second, and to the north- 



ECOLOGY OF ISLE.ROTALE. Ill 

east OD the third day. The northwest winds prevailed both dajs of 
the third ware, while the fonrth started with a northwest wind, and 
changed to west on the succeeding da.r. The fifth wave was )>ei-uliar 
in having an east wind and the sixth a northejist one, both of which 
brought birds of different species, and from a different direction than 
those with nopthweet winds. These two waves were also mnch smallra' 
than the preceding ones. It will thus be seen that the great bnik of 
migration took place with a n(K*thwest wind. 

2, Iitflarufc of TcmprratHrc. Since fall migration pi-evails at a 
tinte when the temperature is gradually falling, the records for a wave 
near the first of the movement would be much lii;{ber than those at 
the last, so this factor can be considered only in a relative way, i. e., 
we must not compare the temperature at the fli'St and. last of the 
season, bnt simply consider the tempera tiu-e immediately preceding and 
following a wave. The average temperature for the thii*teen days was 
47°. This low average was partially due to two days of very low 
temperature. All the waves but one were on a falling temperature, 
and in this cnse the mercury had fallen from the day previous. As 
-a falling or low temperature is the cause of the high barometric pres- 
sure, which in turn with the passage of the high, causes the north- 
west winds which are so favorable to migration, it will be seen that 
a falling or low temperature ii« perhaps the first reqnisite for the bird 
wave. The low temperatnre also influences the food of the migrants, 
billing off the insects, or driving them to shelter, and in this con- 
nection may prove to be very important. 

:{. Inftncmi: of Baronutric I'rrsHuiT. One of the most striking con- 
ditions was the high barometric pressm-e under which these lai-ge waws* 
took place. On ten of these thirteen days the barometer stood at Sti 
inches, or above, the average of these being 3(M7. The lowest pressnits 
was 29. 8, the average for all bedng Sll.Oi). None of the waves took 
]>laee on a falling buroineter, but whei^e there was a change the pre**- 
sure was rising as: first wave 29.9. Hit.i, 30.3. 30.2; second Wave 29.8, 
:t0.1, 30.1; third, 30.1, .30.1; fourth 30.2, 30.4. As before stated the 
direction of the wind is due to the relation of the arenn of low and 
high pressure to the region under consideration, and it is in this con- 
net'tion that it bears upon the problem of migration )ilienotiicmi. 

Cooke in his discussion of the effects of atmospheric changes on 
spring migration shows that at this season the large movements took^ 
place on low or falling baroaneters, and stated that it probably would 
be found that in fall the opposite conditions existed and mignitinn 
would occur on the rising or high barometric pi-essiires. This was found 
true at Isle Bovale and probably is true for all fall mignition. 

4. Condition of the Hky. It will be noticed that on 9 of the 13 
days of bird waves the sky was clear, and on the remaining four it 
was simply cloudy, fflo waves occurring during rainy weather. In 
spring the wa^-es usually occur during cloudy nights; in the fall, as 
witnessed here, the opposite is the case-, and the bulk of the fall migra- 
tion can be said to take place on clear nights. 

H. Summarif and Cmiclusion. From the data submitted we see Ihat 
fall migration as witnessed at Isle Royale occurs, in the majority of 
cases, with a nni'thwest wind and a falling temperature with its rising 



112 MICHIOAN SDHVE7, 1908. 

barometer, and clear aky. (Cf. Smith, '07, p. 223.) It therefore seems 
evident that low temperature and high barometric jttcssnre, with the 
prevailing northwest winds and clear sky which accwnpany them far- 
niahes the most favorable conditions for the bulk of the fail migratioo. 
It will be noticed under the head of Migration Routes that: a few birds 
prefer northeast instead of a nortliwest wind. The conditions which 
would be favorable for the migration of these birds would occur after 
the passage of a high and while the approaching low was still some dis- 
tance off. 

It is desirable that similar observations be carried on at other favor- 
able localities in order to further test these conclusions, and determine 
whether thej are of general application to the fall migration. 

VI. roi;tE3 of migration. 

From the observations made during the falls of 1904 and 1905, it 
seems that Isle Royale ties directly in the path of a very strong mi- 
gratory movement. In the fall there was a great massing of bird 
life. For some unknown reason the path of densest movement was 
very narrow, at least appearances pointed to such a condition. Tbia 
apparent narrowness of the route through the island was strikingly 
shown on September 9 when a trip was made across it from Washing- 
ton Harbor to Siskowit Bay, About 15 miles were traversed, em- 
bracing every environment from clearings to high hardwood forests 
and damp cedar swamps. Nearly a day was spent hunting over the 
clearing and adjacent forest near the head. of the bay, but scarcely 
any migrants were observed. A few Black-throated and Tmnessee 
Warblers and a few sparrows were seen, while an occasional Sparrow 
or Sharp-shinned Hawk was met. This was not due to a lack of food, 
as gras^oppers and other insects were very plentiful. At Washington 
Harbor the reverse was the case; here on September 9 and 10 I saw 
many migrants, the majority of which wwe not seen at Siskowit at 
all. These observations at the harbor were made in the morning before 
leaving and in the late afternoon of the following day when I re- 
turned from the bay. While at Siskowit scarcely a bird was heard 
passing over, although at the harbor they could be heard throughoat 
the night. The path apparently extended lengthwise of Isle Boyale 
with Washington Harbor and the r^on lying between it and the 
north shore of the island as its diameter. 

In a recent paper, Tavemer ('06) makes the statement that perhaps 
a migration route lies between Isle Boyale and Eewe^iaw Point. From 
the observations made on the island, I am led to believe that such a 
route does exist and also one lying much to the west of this point, 
perhaps to the Apostle Islands and the mainland lying Southwest of 
them. These conclusions were drawn from a consideration of the fol- 
lowing facts. The route taken by the majority of the migrants, both 
those which passed slowly across the island and those observed fiying 
overhead, whether by night or day, lay nearly southwest. During the 
latter part of August and parts of September, the nights irere un- 
iiMially bright, so that migrating flocks could often be seen high in the 
air even when not crossing the face of the moon. The cries of mi- 
grating birds, heard mostly on cloudy nights, usually came from a 



BCfOLOQT OP ISLB ROTALE. 113 

northeasterly directioo and died away in a sonthwesterly one. Cer- 
taiu birds, as the Thick -billed B«d-wing Blackbird and 'Lincoln 
Sparrow, which were found commonly at the island, are very 
rare or do not occnr at all in the r^ion directly south 
or southeast of it The Thick-billed Red-wing has never been taken 
at any point in southern Michigan. These birds, being of western origin, 
have gradoally worked their way east where tiiey have found suitable 
breeding grounds, but it seems probable that in their fall migration they 
move westward and join the throng passing down the migration route 
traversed by their ancestors. It seems probable that the greater portion 
of the migrants which leave Isle Rojale, moving in a southwesterly 
direction, continue thus until they reach the Mississippi Valley, where 
they are joined by birds from other regions, and all move down this great 
liighway of bird migration. 

It was observed that nearly all the large bird waves were associated 
with northwest winds. As the birds probably take a southwest course 
this gives them the beam wind which seems to be most favorable for 
their flight. Of course it was impossible to tell from what directiOTi 
the birds came to Isle Boyale, but it seems reasonable that they should 
choose a beam wind when leaving the mainland, since they arrived 
shortly after at the island flying with such a wind. If this proves 
to be tme, the majority of the birds coming to Isle Boyale are from 
the north or northeast. 

The data for the supposed roilte to Keweenaw Point is slight com- 
pared with that for the southwestern one. Two species, the Busty 
Gracfcle and the American Pipit, were observed migrating in this direc- 
tion. During their flight the wind was from the niH-theast giving them 
the beam wind which a number of observers have noted to be the one 
preferred by hawks and gnlls during their migration. Under "Perils 
oif Migration" an instance is cited where a number of birds were caught 
by a storm while crossing to the east of the island and were driven 
to Washington Harbor. These birds were possibly crossing to Kewee- 
naw Point. Probably the majority of the bdrds which strike this point 
are from regions lying to the northeast of it, and arrive there on north- 
west winds as do those birds which n^grate across Isle Boyale. The 
theory that many birds skirt the Great Lakes, 'as brought out by 
Tavemer, explains the absence of several species from Southern Mich- 
igan which is not done by the discussion of the routes from Isle Boyale. 

VII. TUB PERILS OF MIORATION. 

Dixon in his "Migration of Birds," divides the perils of migration 
into three important classes : flrst, those arising from fatigue doe to 
the mechanical part of season-flight; second, those arising from the 
natural enemies of each species; and third, those arising from Munders 
and fatalities on the way. These three classes were observed in vary- 
ing degree during the fall migration at Washington Harbor. 

1. Fatigue. Between the north shore of Lake Superior and Isle 
Boyale, the distance is so short that unless unfavorable winds inter- 
cept them the older birds would have little trouble from fatigue due 
to the simple operation of flight; but the young, which often commence 
migration soon after being able to fly, would experience considerable 



114 HlCHIOAN SURVEY, 1908. 

strain on their frail bodies in even bo short a flight. This was vividly 
Bhown in the number ot exhausted young found after every bird wave. 
During September, immature warbler* and sparrows were often founil 
in the morning in an almost completely exhausted condition after their 
night's flight. This was especially evident among the Tennessee and 
Rlackpoll Warblers. On the morning of September 13, following the 
day which witnessed the largest bird wave, I picked up many dead 
birds. Nearly alt were warblers, the Tennessee seeming to have suffered 
most, although the Palm was a close second. A few inunature Savan- 
nah Sparrows, one adult Yellow-bellied Flycatcher, and ae\'eral young 
Flickers were also found dead. These dead migrants were seen in the 
clearing, along the sroads, aud on the banks of the stream. An ex- 
amination of these victims showed no outward indieoition of the cause 
of their death. Only a few were emaciated to any extent. It was a 
noticeable fact, however, that none of the birds found dead were in 
the prime, fatty condition of most of the other migrants taken. The 
conclusion therefore seems probable, that the birds must be in the 
best condition possible to miake a successful migration flight, and that 
the greatest mortality among the migrants lies in that class which for 
some unknown reason are not in prime condition. As there were no 
other reasons evident which could have brought on this high death rate, 
it is probable that death had been caused by severe exertion, coupled 
perhaps in a few instances with lack of food and unfavorable weather 
conditions in which to recuperate. This seemed the more plausible 
considering the fact that in only a comparatively few cases were the 
victims adult birds, while, as before stated, many of the immature 
warblers and sparrowH had only been ahle to fly for a ^ort time. 

After heavy storms, especially those from the southeast, many adult 
as weli as young birds -were fonnd in an exhausted condition, their 
plumage presenting a dilapidated appearance, the wing and tail feathers 
broken, and showing genera) evidence of a hard struggle with the wind. 
Some of these birds may have been caught by the storm while cross- 
ing from the north shore to the island, but as the birds appeared to 
be blown before the wind I think thait at least part of themi were over- 
taken while crossing the take considerably to the east of Isle Royale, 
perhaps toward Keweenaw Point. Overtaken by the storm and with 
no place to take refuge they were gradually blown in the direction 
of the island where they were found the succeeding morning in such 
an exhausted condition. Some of these birds would even allow them- 
selves to be picked up and handled without showing any fear. The 
birds which suffered moat were the Palm and Tennessee Warblers. 
Michael Hollinger, a i-esident on the island for several years, told me 
that often, especially ini spring, he had seen Washington Harbor "liter- 
ally covered" with floating birds which had succumb^ in their struggle 
against the storms and had drifted in from the open lake. The peciUiar 
shape of the harbor and the lake currents tends to mass floating bodies 
at this point. But the loss as shown by those collected at the harbor 
oonld be bat ai slight proportion of the vast numbers which must have 
jterished in the open lake. 

Se^-eral flshermen said that after heavy gales in late fall and early 
spring, the shore at Washington Harbor would be strewn with the life- 



• ECOLOGY OF I8LB ROYALK. IIB 

le«s bodieB of birds thrown up by the ■*'BTea. During the fall of 19H3, 
birda w-ere several times reported as lighting on the ehipe coming into 
the island, andt the fishermen eecured several which lit on their small 
boats after a storn^ when about two miles from land. They reported 
the birds as very tajne and allowing themselves to be handled freely. 
The birds secured were several small Bparrows, Tennessee and Palm 
Warblers, a Saw-whet Owl, and one adnlt Bobin. Theae birds were 
all encountered near the southwestern end of the island. They had 
probably been blown out of their course and were striving to reach 
the nearest point of land, as no birds would be coming from the south 
at this time of j-ear nor would any so completely exhausted have at- 
tempted to leave the island. 

2. Natural Enemiea. Without doubt the greatest natural enemies 
of the birds during migration were the Hharp-shinned and Sparrow 
Hawks. At times the Pigeon Hawk made great havoc among the 
smaller birds, and the Owls aim played a small part. Probably weasels 
and minks fed to aome extent rm the migrants, which they canght while 
the birds were resting. These animals, however, onJy destroyed com- 
paratively few, as remains of their victims were seldom fonnd. Like- 
wise the Owls probably destroyed only those which came direftly in 
their path, the abundance of the Varying Hare furnishing a food much 
easier to procure. This undoubtedly saved a large number of mdgrants. 
Of the other animals, the Lynx also fed largely on the Hares and so 
probably molested the birds very little, while the family of house cats 
kept at the clnhhonse were more than supplied by the n'umi9er of small 
birds which met death striking against the windows, etc. 

The early migrants were preyed upon very little by the hawks, prin- 
cipally because the Sharp-shinned Hawk had not arrived in any ap- 
preciable numbers, and secondly, the great swarms of grasshoppers fur- 
nished an abundance of appetizing food. As the season advanced and 
both species of hawks grew more uumerons. their effect on the bird 
life increased. None of the smaller birds were safe, away from the 
protecting boughs of the conifers and alders, and therefore were con- 
fined almost exclusively to the edge of the clearings. The Sparrow 
Hawks fed both on grasshoppers and on warblers and sparrows, while 
the Sharp-shinned fed almost entirely on the latter. During the bird 
waves the hawks became more numerous, this being especially true for 
the great wave of September 12, On this date great numbers of both 
Sharp-shinned and Sparrow Hawks made their appearance, as well as 
many of the Pigeon and a few CoopCT's and Broad-winged Hawks. 
The Pigeon Hawks in particular timed their migration to that of their 
victims, appearing and disappearing with each successive wave, very 
few remaining on the island. The majority of the Shapji-shianed also 
kept pace with the retreating birds and by the time the bulk of the 
warblers and sparrows had passed they too had gone on. Among the 
birds which suffered most heavily ntay be m«itioned the Tennessee, 
Blackpoll, and Palm Warblers, the Wilson's, Olive-backed, and Oray- 
cheeked Thrushes, and the Chipping and Savannah Sparrows. Great 
daring was shown by the Sharp-shinned Hawks. Sometimes so eager 
were th^ in pursuit of their i»a-ey that they would dart within a few 
inches of one's head. 



lie MICHIGAN SnRVBT, 190S. 

3. Blunderg and Fatalities. A comparatively new danger vbich 
besets migratory birds on tJie IslaDd is the fatal attraction of the lighted 
windows of resorts and the light-housefl. During the migration scores 
of warblers, chiefly Tennessee and Palm, killed tliemselvea by sfriliing 
against the lighted windows of the Club-house which stood 4n a clearing 
near the Harbor. Many also met death by tlie same means at Washing- 
tori Island, which is situated at the entrance to the harbor. Among 
tbe species killed at the latter place were the Tennessee, BlackpoU, 
Myrtle, Magnolia and Palm Warblers, Gray-cheeked, Olive-backed and 
Hermit Thrushes, and several species of sparrows. On September 2, 
during a. hard storm which lasted several days, five Olive-backed 
Thrushes were fonnd dead by Wood beneath the windows at the hotel, 
and on September 5, a Oray-chedied Thmsh was found dead at the 
same place. This latter was the first one of this species seen^ no other 
being observed until Beptembei- 12. Alt the birds were killed on the 
north side of the buildings. Sometimes after cloudy nights numbers 
of small birds would be found on the north porch of the Club hou^ in a 
dazed condition, probably from striking the building the night before. 
Both young and adults were found, the young being the only ones 
killed on clear nights. 

The light-house keeper at Menagerie Island in Siskowit Bay, Sir, J. 
H. Maloue, reported that hundreds of birds lost their lives every spi-ing 
and fall at his light alone. It was mainly on cloudy nights that the 
birds struck the lighted windows and the lantern, but some were killed 
on other nights. 

REFBBBNCBS. 

Oooke, W. W. 

1888. Report on Bird Migraticm in the Mississippi Valley in the 
years 1884 and 1885. 
Bull. No. 3, Div. Economic Ornithologj', U. S. Dept. Agr. 
Smith, Frank. 
1907. Advantages of Migration Records in Connection with Bird 
etudv in Schools. 
School Sci. and Math., 7, pp. 221-224. 
Taverner, P. A. 

1905. A Hyperlaken Migration Boute. 

Bull. Mich. Omith. Club, VI, pp. 1-7. 



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THE KrOLOGICAL Sl'CCESSIOX OF BIRDS." 

BY CHARLES C. ADAMS. 

"Of all truths relating to phenomena, the moat valuable to «a are tboM which 
relate to thetr order ot Buccession. On a knowledge ot theK la founded every 
reasonable anticipation of future facta, and whatever power we possess ot In- 
fluencing those facts to our advantage," — John Stuart Mill. 

"Indeed, some geologists aeem to take pride in lack of knowledge of principles 
and of their failure to explain the facts obaerved in the terms of the elementary 
scleaces. ' I have heard a man say: '1 observe the facts as I And them, unpreju- 
diced by any theory," I regard thia statemeni as not only condemning the work 
of the man, but the pmltlon as an impossible one,,.. The geologist must select 
the facts which he regards of aufflclent note to record and describe. But such 
selection Implies theories of their importance and significance. In a given case 
the problem Is therefore reduced to selecting the facta for record, with a broad 
and deep comprehension of the principles involved, a deQnlte understanding of 
the rules of the game, and appreciation of what Is probable' and what is not prob- 
able; or else making mere random observations. All agree that the latter alter- 
native Is worse than useless, and therefore the only training which can make 
a geologist safe, even In his observations. Is to equip bim with such a knowledge 
of the principles concerned as will make his observations of value." — Pbeside.vt 
C R. Vas Hise. 

I. IXTBODfCTIOX. 

-Almost everv obsener of aniiimiR liiis noted that certnin kinds of 
birds are uaually found nssocinted in certain conditions, as, for ex- 
ample pflrticnlar species of Kaiid[>i)>ers and plovers upon the sand.v 
beaih, or the Mendonlark and Dickcissel upon certain prairies; but 
tills is rarely considered a subject w-orthy of'serious scientific study. 
To discuss the significance and (-alue of such ecological study and sug- 
gest phases for investigation is tite object of this pai)er. By the ecologi- 
■cal distribution of birds is meant that correlation between envircm- 
mental conditions and the occurrence and association of certain species 
of hirds. In such study special attention must be de\'oted to the places 
of breeding; nevertheless the associations of birds at all seasons of the 
year are of importance. It is not the isolated occuri-ence of these 
8i>eciea, but their relative abundance, the association of certain species, 
and thpir' persistent wcurrence in such conditions which is signifi- 
cant. In the literature of omitliologv- there is a vast amount of isolated 
data bearing on tiiis subject, but very little of it has been organized 
and systematically studied. 

When once the fa^-ts and general ecological relations have been de- 
termined, BO that the representative bird associations or societies of 
given localities have Iwen correlated with their ]iroi>er environ- 
nients it will then lie possible to determine how one society l>ecomes 
"transformed into another, whether this is due primarily to other birds 
or to other environmental influences, A knowledge of the succession 
of bird societies and of the laws of change will not only lend to new 
ideiis as to the intluence of the envii-onnient. but will also have a 



• Reprinted, wilh llie sddHlon of cliapler VI, (rom the Auk, 25, ji 



M«r;Coogle 



122 MICHIGAN SURVEY, 1908. 

marked influence npoii the practical field studies of the bird student. 
It should lead to a more intelligent understanding of the relation of 
birds to the world about them, or even better, to the world of wliicU 
they form a part. 

Attention ebould further be directed to the fact that simply tlin 
occurrence of the bird in a definite habitat is not by any means the 
sole aim of such wort. The influence of the environment should l»e 
studied, in its bearing upon all phases of bird life. Not only slioiilcl 
the most favorable habitat or optimum^ be recognized, but also the in- 
fluence of the less favorable conditions; thus the nesting site, composi- 
tion of the nest material, food, abundant'e, feeding grounds, mignttinj; 
habits and all like relations are needed for an adequate and exhaustive 
study of the ecological distribution and succession of birds. 

It is tlierefore not surprising that such requirements will be difti- 
cu!t to meet because the facts themselves are difficult to secure. Then 
there are further difficulties which ai-e due to the limitiitions of tlie 
student bimiielf, and are psychological in their nature. As examples <if 
this class of difficulties two may be cited, because they are of fi^eqiient 
occurrence in all kinds of scientiiic work and not by any means confined 
to the study of birds. For, contrary to our youthful ideas, naturalists 
have the same limitations as humanity in general! We may divide 
naturalists into two classes, depending upon their primary type of mind. 
First, those who tend to see only the infinite detail of isolated facts and 
ob8er\'ations. Tliis type of mind is jMirtirularly impressed with tlwt 
multiplicity and variety in natui-e, and is one to which a general state- 
ment is almost a cause of irritation because there are usually exceptions 
to any general statement. The constructive imagination seems feebly 
developed in this type. To this class belongs many extremely valuable 
and useful students, because of the data which they, often with extreme 
conscientiousness, collect. They are collectors of facts rather than stud- 
ents of relations. To the second class belongs that type of mind whose 
primary interest is in genei-aliaations, principles, relations, and wbieh 
tends to -neglect isolated facts and observations. The constructive im- 
agination is liable to be developed in this type. Tliis includes many 
extremely valuable and useful students on account of their tendency lo 
condense, sift and formulate great masses of isolated facts. They are 
students of ideas and relations rather than collectors of "facts." 

Each class, esijecially the well-marked ty[)es and extremes, often find* 
it difficult or impossible to understand the point of view of the other 
class. This frequently leads to misunderstandings and often to mutual 
contempt. Cope and Marsh clearly illustrate these two types of mind» 
among our American naturalists. 

By this time some may wonder why this subject has been introduced. 
It lias been with a definite purpose, because fi-equently these opposed 
points of view cause delays in the development of many subjects. Thu» 
a forewarning to students of bird ecology may produce good results if 
the individual student makes a ronscious effort to counterbalance such 
deficiencies as go with his particular type of mind. In the past, details 
have tended to pi'oduce confusion through the neglect of general ideas. 
It is rarely that a word of warning on this subject is out of place, be- 
jMUDse the balanced "golden mean" investigator is nevei- too abundant. 



ECOLOGY OF ISLE ROYALB. 123 

The quotations at the head of this article have au immediate bearing 
upon tiie subject at this point. 

Not only is habitat preference, the asaociation of avian species, their 
succession, and the laws expressing these relations of much interest, but 
they are of much importance scientifically as well as in a practical way. 
It is therefore desirable that naturalists realize the necessity of under- 
standing the "roles of the game" if the true relations of birds «re to be 
studied to the best advantage. No adequate substitute has yet been 
devised to replace a grasp of general principles. 

Throughout this paper emphasis is placed on the definiteness of the 
dominant major environmental influences and compleies because the 
irregular features have apparently received undue emphasis and have 
retai-ded the recognition of certain important detinite relations. 

II. REPRESENTATIVE LITERATCHE OX HABITATS AND SUCCESSION. 

1, Habitat Prefcrenfc. The American literature on habitat prefer- 
ence and succession, as a subject of special investigation, is very limited. 
By succession is meant the change or i-eplacemMit at a given place of 
one or several species (an association) by others; as when a swamp is 
invaded by a dune and the representative swamp birds are replaced by 
those of the dune; or even a^in when the dune becomes fixed by vegeta- 
tion and is inhabited by still another association of bird life. -This is ^ 
a much neglected subject; however, isolated observations on habitats 
are abundant in the biographies of the various species. The fragment- 
ary- character of these biographies tends to make them composite and 
they lose what pecularitics they may have which are due to a response 
on the part of the bird to its particular conditions of life. These un- 
fortunate limitations clearly show that here is an extensive field worthy 
of careful investigation. The work already done will be a usefnl'guide 
in many cases, but the student who wishes to develop this subject must 
turn to the fields and forests rather than to the literature, both for 
his inspiration and his data. 

Perhaps a further word should be added concerning the limitations of 
the composite life-history method, as this will aid in making clear the 
kind of work needed in the future. This composite or generalized method 
of describing habitats and life histories and the response of birds to them, 
tends to lay undue emphaKiK upon the aceragc conditions of life and habits, 
and tends to neglect those detailed responses to the environment which 
reflect the laws of local influence. These results are similar to those pro- 
duced by systematic students who are "lumpers" and who do not recog- 
nize local races or varieties. Thus a nest may be built upon the ground 
at the base of a shrub or bunch of grass, or in the brush, but what 
conditions determine such sit^? In a dry meadow a Hong Sparrow may 
build directly upon the ground, but in a swamp, in order to have a dry 
nesting site, it builds in a willow shrub. In many cases the causes of 
these differences will be difficult to determine, but in others it is a 
relatively simple question for any one familiar with the species to solve. 
There are also geographic variations of habits as well as those of 
habitats just cited, and for this reason it is necessary not to confuse such 
variations with those confined to some restricted area. These locqlL. 
and ge<^airiiic relations are very intimately related, but they are stib^ 



124 MICHIGAN SURVBT, 190S. 

jects which can only be worked out in detail wlien local studies gire 
proper attention to local enyironmental responses. 

In the following account of the literature no attempt is made at 
completeness, but the papers cited are helieved to be representative. 
These pi^rs will help to give some idea of the kind of observations 
and records already made, and wilt be suggestive as to future work. 
Mention will first l>e made of tbe literature on habitat preference, and 
then of that on succeeeion. 

By far the best discussion we have found on habitat preference of 
the birds of a given region is that by Townsend ('05) on Essex County. 
MassachusettB. Ilie primary avian environments are described, the 
representative birds listed, and their preferred habitats are briefly dis- 
cussed. niuB, the ocean and its birds, the sand beach and its birds, 
the sand dnnes and their birds, the salt marshes and their birds, and 
the fresh marshes and their birds, give a general idea of the subjects 
treated. B^^arding the birds of the sand beaches, he remarks : "Among 
the Plover, the Black-bellied, Bemipalmated, and Piping Plovers are 
above all birds of the beach, although the first two are occasionally* 
found in the marshes, while the last-named rarely strays from the beach 
and the adjoining sand dnnes. The Golden Plover, although at timies 
found on the wet sands, is much more likely to hunt for food on the 
dry sands above the highest tides, or still farther inland, while the 
Eilldeer genra'ally avoidB the beach altogether, preferring the fields" (p. 
21). And regarding the birds of the sand dunes he remarks: "Savanna 
Sparrows nest in numbers at the foot of clumps of tall beach grass 
throughout the dunes, and on the edges of the tidai inlets from the 
marsh. The nests of tbe Bed-winged Blackbirds and the Bronzed Grack- 
lee are abundant in the bogs and groves of the birches. The Crow, 
in the absence of tall trees, builds perforce in the stunted pines and 
birches, at times only ten or twelve feet from the gronnd'' (p. 34). In 
the case of the Crow, note that he records the response to the dune 
wivironment. 

While TowasCTid recognizes changes in the environment, as in the 
dunes and beach (pp. 21, 30), yet he does not see their relation to 
the bird life in the definite -wnj in which he sees their habitat pref- 
erences, nor does he appear to clearly recognize the fundamental re- 
lation of association within the breeding habitat. To him the environ- 
ment ia static. However an excellent feature of his work is the record 
of seasonal changes in the bird life of the various habitats. In thin 
connection attention should be called to certain papers which will 
greatly aid in the study of the dynamics or changing environmentul 
factors which influence sea or lake shores bordered by dunefl and 
' BwampB; conditions represented on the MassachusettB coast. Gilbert 
('85) has discussed the general principles and top(^raphic features of 
lake shores and Gulliver ('1)9) the shore line of the sea. But in ad- 
dition to these physit^raphic forces, the vegetation also has a dontinat- 
ing influence upon bird life. For genera] principles relating to this 
subject Cowles ('01) should be consulted for his discussion of the vegeta- 
tion of inland Bhoree and dunes, and Ganoug {'(Ki and '06) for his 
treatment of the .Atlantic coastal conditions. These authors discuss 



, Google 



ECOLOGY OF ISLE ROYALB. 126 

the SQCceesion of the vegetation, a factor of the atmost importance in 
the study of avaiu Bui;ce«aion. 

While considering Townsend's reenlts, it nmy be well to outline 
briefly a general euccesmon of bird life along the shore as indicated 
by his records. It is evident from the map accompanying his volume 
that the currents and waves are constantly modifying the coast line 
and f^H-miing spits, bars and islands ; and that the barrier beach area is 
increasing, and thus tending to become continnous at the expense of 
soDie of the ocean habitat. As the continuity of the beach develops, 
the area of swamp land behind it tends to increase and thus to farther 
restrict the open water and increase the swamp haibitat. The beach 
sands, once free from the waves or ground water, are caught up by the 
winds to form dunes, and may migrate into the swamps and thus trans- 
form them. Thus with the extension of the beach the sea birds are re- 
placed in dotninance by the shore birds, and a succession is produced. 

In a similar manner the dunes encroach upon the swamp, and swamp 
birds are succeeded by those of the dunes. As the wandering dunes 
become anchored by vegetation and forests grow upon them, still other 
birds will invade thrai. Thus all stages may be expected, from ocean 
to beach birds, onward to those cliaracteristic of wandering and fixed 
forested dunes. These relations are outlined simply to indicate the 
problem and its causes, which need detailed investigation. 

Tn Michigan a few habitat studies have been made. One in the Por- 
cupine ^fountains, on the south shore of T^ake (Superior, and another on 
Isle Koyale. Both are by McCreary; the paper on the latter area is pub- 
lished in this volume. The summer birds of the Porcupine Mountains are 
listed (M«*-"peary 'OC) by selected' localities and the habitat preferences 
are discussed as follows: water birds, birds frequenting the shores and 
banks of streams, birds frequenting grassy meadows and alders, birds 
frequenting tamarack and cedar swamps, birds frequenting hemlocks 
and maples, and birds frequenting the cliff and mountain top. In its 
emphasis upon habitat preference this paper is the only one so far seen 
which at all approaches Townsend's discussion of this subject. Mc- 
Creary's work was done without a knowledge Of Townsend's. 

In southeastern Ihlichigan,, Brown ('00) made a locality study and 
outlines the habitats as follows: birds fonnd in orchards, birds of the 
ojieo woods, birds of the open fields, birds of the thicket, and birds of 
the marshes and river. Brown's paper is intermlediate in character 
between the preceding papers and those of an economic nature, to be 
mentioned later, because the area studied has been so much influenced 
by man. 

There are a few papers which, althongh primarily faunistic or geo- 
graphic, contain habitat data. Such, for example, is Ridgway's (74) 
discussion of the birds of the Wabash Valley and ('89) the birds of 
the Illinois prairie (pp. 13-16). An exceptionally good paper of thifi 
character on the Louisiana birds is by Beyer, Allison and Kopman ('06), 
although its aim and method of treatment differs from that of Town- 
send. The bird life is, however, closely correlated with the vegetation 
and the physical conditions of the State. 

The papers previonsly mentioned have been written from a r^ional 
standpoint. The study however of all the various conditions frequented. 



126 MICHIGAN SURVEY. 1908. 

by a given speciee or some natural group is also an important and 
Delected method of ecological study wbich poaaeflsefl certain important 
advantages. As an illustration of this method may be mentionetl 
Palmer's ('00) study of the Maryland Yellow-throat. He has shown 
that different varieties have different habitat preferences, Jacobs ('041 
has given us an interesting habitat study of a single species in Pennsyl- 
vania, the Golden-winged Warbler. 

Let UB now turn to another class of habitat studies, those which 
through man's iufluenre throw only a subordinate li^t upon "natural" 
habitats and succession, and are primarily of economic importance but 
contain valuable habitat data. 

An interesting and rather unique paper belonging to this class, based 
on observations in Southeastern Michigan, is by Watkins ('00). It is 
entitled 'Michigan Birds that Xest in Open Meadows.' A few of his 
statements explain his point of view: "To make ntore plain the limit 
and scope of this treatise, which, of necessity must be longer than I 
hoped, I will include in my list only such species as I have found nesting 
u{>on the ground in the open fields and meadows, excluding those found 
nesting upon the boundary fences or ground; also those nesting in the 
open marsh land which are undraioed and boggy to the extent of being 
unlit for hay or pasture" (p. 67). The paper contains numerous notes 
on the habitat preference and variations in these traits. 

By far the most comprehensive and thorough study of any limited 
farni area is that by Judd ('02) of a farm in Maryland. In this paper 
habitat preferences are clearly recognized, and discussed rather fully 
(pp. 12-20). The birds are associated thus: — birds that nest in the 
open fields, birds that depend on covers, birds of less limited distribution 
{consists largely of remarks on haunts), and birds of varied distribu- 
tion. His last two sections are rather miscellaneous in character and 
show that the principles of classification for habitats were not clearly 
defined in his own mind. 

The only other paper discussing habitat preference in detail is also 
the latest upon the subject, and is by Forbes ('07), This is a pre- 
liminary report on a bird census across the com belt of Centra) Illinois 
in the early autumn; a study of the feeding grounds and preferences as 
influenced by the dominant crops of the arear traversed, corn, pasture, 
and stubble. By means of this census, the hRbi^dt prefei-ences for differ- 
ent crops and the association of species in them is statistically deter- 
mined. The paper is particularly suggestive for its bearing on the sub- 
ject of dominance; however, the suggested method of study has even 
greater significance when applied during the breeding seaeon. Doubt- 
less opinions will vary as to the validity of the method aa applied by 
Forbes, even by thtwe who ^-ould approve of it for the detailed study 
of a limited area, or a breeding habitat. For large areas some co- 
operative method m«iV be necessary. 
■-2. Succession. Turning now to the literature on succession, it is 
'^und to be extremely limited in amount. So far as known to the 
writer, only two American authors seem to have realized the existence 
of succession. In his discussion of the liiotic succession in the Por- 
cupine Mountains of Michigan, Ruthvcu (*00) clearly included the birds. 
alth<mgh they did not i-eceive separate treatment, and might for this 



ECOLOGY OP ISLE HOTALB. 127 

reason be overlooked. His poaition is clearly stated (p. 43) as fol- 
lows: '"OwiDg to the dependence of forms of life on their environ- 
ment, biotic (.'hnngeti are neeeHxarily closely i-elated to en vii-on mental 
cluinges. These biotic changes may occur in two ways; the forms must 
("ither be able to respond to the new conditions, or be supplemente'l 
by other forms. That they tend to become adjusted cannot be ques- 
tioned, bat in many caaes at least, this adjustment Ia(pi behind the 
elian^Dg conditions, and the fomw are replaced by others from ad- 
jaceut habitats which are adjusted to the conditions toward which the 
particular habilAt is changing, thus bringing about a succession of 
Mocieties." In speaking of the biota of the hard-wood forest he further 
says: "This region has been reserved for the last, for the conditions 
are evidently those toward which the other habitats tend to be changed 
under the present conditions. , . .This society thus represents the climax 
so<iety of the region. It consists of the forms that are adapted to or 
associated ii'ith the conditions which prevail in this region in the last 
stages of the mirtual adjustment of all the environmental processes. 
As the processes become adjusted to one another, the habitat of the 
climax society is increased at the expense of the other habitats, and 
the associated biota tends to become of general geographic extent in 
the region." 

The only other paper discussing avaio succession is that by Frothing- 
ham ('06). and this is not a "natural" succession but one influencetl 
primarily by man. He clearly expresses a bird succession correlated with 
the reforestation of bui-ned lands. The area studied is the Michigan 
foi-est reserve on Higgins Lake. The n^ion was originally corered with 
"White and Norway pine, hut repeated flres first killed off the pines, later 
the oak and maple; and finally the dominant vegetation is sedge, sweet 
fern, huckleberry and prairie willow. With the Are protection afforded 
by the reserve, Frothingham anticipates a revei-sal of the above succes- 
sion of destruction, and further remarks: "With the types of v^jeta- 
tiou which mark tlie different stages of the plant suci-easion just de- 
scribed there seem to be correlated certain definite bird forms. These 
forma are for the most part such as frequ^t observations in northern 
Michigan have identified as generally characteristic of the respective en- 
vironments." This is follon-ed by lists of birds characteristic of differ- 
ent kinds of v^fetation. While these lists do not correlate perfectly 
with the implied succession, yet thfe general statement of the problem 
!<' clearly ezpt-eased. 

The burning of forests has long been known to change the character 
of the v^etation and fauna of areas, but this is often referred ^o as the 
change of a "life zone." Thus Merriam ('99. p. 47) states that a fire 
in the- Canadian zone on Mt. Bhanta is followed by the Transition 
zone and remarks: "But in the meantime a new growth of Shasta 
flr has started, and in ten or twenty years is likely to overtop and 
drown out the Transition zone species, enabling the Canadian zone to 
reclaim the burn.... But on the steeper slojtes, especially rock slopes, 
if the vegetable layer is burned off, the (lower) zone which creeps up 
to replace the (higheri one destroyed l>ecomes {>ennanent or nearly 
so. . . . Deforcatration of an area therefore tends to lower its zone posi- 
tion." Birds are not mentioned in this discussion nor the relation of 



126 MICHIGAN SURVEY, 1908. 

"zones" to tbe general problem of succ^sioo. Bach "zones" are thus 
only particular phases of BUccessioQ. 

It ifl thus Been from the above outline of literature that habitat pre- 
ferences have been outlined for a few widelv eeparated localities and 
for Bome a^cnltural conditions, but there has been no comprehensive 
diRCUBsicv of the jH^blema of habitats and succession, even in a pr^ 
Iimittar7 manner, either fromi a scientific or economic standpoint. This 
fact seems rather remarkable in viev of the great utility of a knowledge 
of the general principles underlying economic practice. There are, 
however, certain phases of biotic succession which have been discussed 
by a few authors. Theae subjects have either been discussed in a very 
general manner or are detailed discussions of special re^ons or groups 
of plants and animals. For this reason, perhaps, their bearing upon 
other groups than those speciflally mentioned are very likely to be over- 
looked by those who take little interest in any subject or discussion 
which does not specifically mention their specialty or locality. This 
phase is mientioned in order to show that while avaJn successions have 
iH-en considerably neglected, advances have been made elsewhere, by 
means of which some general principles appear to have been fairly 
well established. This is particularly true of plant succession, as 
shown by the writings of Cowles ('01), and in considerable detail by 
Clements ('05). The discussion by Clements will be particohiirly valu- 
able to tbe student of avian succession. 

III. THB IMA30B AVIAN ENVIRONMENTS. 

Ab has been seen in the preceding review of the literature on haunts, 
no comprehensive discnssion has been given of the environmental influ- 
ences or ecological distribution of (extra-tropical) North American 
birds. Various authors have discussed their ge<^Taphic distribution, 
and certain geographic variations have been referred to certain environ- 
mental influences, but a general ecologic treatment, as contrasted with 
a primaril,!- faunistic one, has not been made. This is remarkable when 
we recall the fact that the collections of North American tnrds are, 
considering the large area concerned, the beet in the world both as to 
quality and as to quantity (Ktejueger, '03). This means that there 
have been many trained collectors; but what has become of the notes 
and obsert'ations on the environments and conditions of life of these 
birds, which must necessarily have been known to successful collectors? 
Part of these observations have been published, and periiaps no one is 
to blame because more have not; but the point of significance is that 
w<e have, in fact, hardly made a banning in the careful detailed study 
of the bird environment and its development as a distinct field of study. 
In common with the remainder of the North American biota, several 
general principle.^ are known, but they do not appear to be current 
among ornithologists. 

The following discussion and su^^^tions on the larger environmental 
units attempt only an outline of certain phases of the problem, in order 
to call attention to certain principles which seem useful as a background 
for the intelligent study of bird habitats and succession. From such 
a standpoint as this, the dominant infiuences of given areas and environ- 
ments are of particular interest and of fundamental value. By focuSB- 



ECOLOGY OF ISLE ROYALE. 129 

ing attention upon the importance of recognizing these dominant en- 
I'iTonmental inflnences, we niny hope to escape some of tbe confuaitm 
which appalB those who are keenly intpresBed with- the chaos and com- 
plexity of the problem. These dominant factors are usually not single 
isolated forces, but resultants of several or many influences. Thus, 
as in the case of the v^etation. it is not one factor, but a complex, 
which influences different birds in different ways. Nevertheless there 
is what may be called a mass or dominant effect. 

A major habitat unit may be considered as a combination of condi- 
tions which are dominant in a t^rtain area. The ifrj/ dominance means 
that a relatU-ely limited number of forces or comphrxea are operative. 
With departure from such a center of influence the dominance changes, 
as other influences are encountered and other dominants are established.' 

When we consider that certain ecological groups of birds are world- 
wide in their environmental relations, it becomes evident that such 
characters are of fundamental importance. Thus water birds may 
occur in any part of the world where water is ' a dominant environ- 
mental factor. This is not a simple ecological group of birds, bat one 
of the greater units of association which may be subdivided into many 
minor classes; as those which frequent tbe sea, and others the inland 
bodies of water. The shore birds form another natural ecological 
group, and also the inland birds a third. There may thus be considered 
to be three primary ecological groups of birds which are closely correl- 
ated with definite and dominant environmental influences: Thus: — 

1. W^er blrdB. 

Tbose trequentlng the sea and the adjacent rocks on which tbey oeet, 
and Inland waters. 

2. Shore and Marsh birds. 

Those frequenting shores of all kinds, seas,. lakes, swamps and rivers. 

3. Inland birds. 

Those frequenting deserts, grass lands and forests. 

' Of course these ecologic:il classes are not sharply defined, and yet 
they are so distinct that they can be easily recognized. It should be. 
noted that the above groups are closely correlated with certain domin- 
ant physical features of the earth— the sea, the shore and the inland 
environ ments. 

Tbe relative abundance and dominance of these classes of birds will 
be determined largely by the dominance of such physical conditions 
as most distinctly favor a particular ecological group. Thus at sea 
the water birds are dominant; on shore, the shore birds; and inland, 
still other kinds. The linear character of the shore habitat and tbe 
adjacent breeding grounds gives it a rather unique character, as the 
two other habitats occupy lai-ge expanses. However, the swampy, some- 
what shore-like conditions of the far north most nearly approach, for 
the shore birds, the expansive character so usual for water bodies and 
inland areaa 

In the present discussion the emphasis placed upon the inland vegeta- 
tion does not mean that tbe dominance of other influences is not recog- 
nized, but simply that it makes a convenient and fairly reliable index 
to many other environmental ivfltiencea. as. for example, the climate 
and topography. A further important advantage of the plant index is 
17 • 



130 MICHIGAN SURVEY, 1908. 

that the st-ieace of plant ecologv and many of its general priocipIeH and 
methods are applicable to birds- A general knowledge of plant eeo\t>gy 
is therefore becoming one of the most valuable tools in the hands of the 
field ornithologist. Everj- field naturalist has obser\-ed the general cor- 
relation of certain birds with certain kinds of vegetation. This rela- 
tion is clearl.v eKi»re».sed by Ridgway ('8i), p. S) us foliowM: "There 
is probably no better index or key to the distribution of birds in any 
countrj' than that afforded by the character of the vegetation; should 
thifi vary essentially within a given area, a corresponding difference 
in the bird-life is a certainty." This phase of the subject clearly illus- 
trates the oft-i¥peated experience of naturalists that in order to tUor- 
ougjily understand one subject — perhaps the favorite one — it becomes 
necessary to study another, or even several. Thus iui order to know 
the bird life of a region it has become necessary, to study the ecolo- 
gical relations of its vegetation. 

The study of eoolt^cal plant gec^^phy is an extensive one, but many 
of the details, so important to the botanist, are of much less concern 
to the ornitholr^ist, who needs primarily to knoT the major plant as- 
sociations or formations and their snccessional relations. This im- 
plies ability to re<^'C^rnize dominance among plant si^ecics and the gen- 
eral method of transformation from the dominance of one to that of 
another. 

By a plant formation i^ meant that association of species (or plant 
society) which is correlated with those conditions which tend to pre- 
vail over a large (ceographic area in the last stages of mutual adjust- 
ment of nil en vi ran mental and biotic processes. Snch an association 
or formation tends to occupy sach an area to the exclusion of all others, 
■end is thus a climax society. 

But absolute dominance of a formation does not occur, because local 
conditions break the monotony where streams, water basins, bare rock, 
and similar influences may interrupt the desert, grasslaiud or forest, 
and produce minor habitats and associations of both plants and 
animals. 

It is not my purpose to discuss in detail the varioos plant formations 
of fextratropical) North America, but to outline those which are of 
evident omitliolopical utility. The following may be recognized pro- 
visionally : — 

1. The Arid Deaerte of Southwestern U. S. and the Mexican Plateau. 

2. The OraselHiida of the Great PlaloB. 

3. The Deciduous Hardwood Forest of Soutbeastem U. S. 
1. The Coniferous Forest of Eastern Canada. 

6. The Giant Conifer Foreat of the Pacitlc Coast and the Rocky Mountafna. 

6. The Barren Grounds or Cold Desert. 

7. The Alpine Deserts, 

A mere inspection of this list of avain and vegetational formations 
shows that the recognition of these large environments is relatively 
simple. It is also seen that they represent fairly definite physical 
or environmental complexes of such fundamental importance that there 
can be no doubt as to their general validit.v. As to the relative value, 
influence- iKiundaries. and the dynamic relations of these formations, 
much is already known, but not as an organized bod.v of facts and prin- 



ECOLOGY or ISLE ROYALE. 131 

ciples. It will also l>e noted that these regioDR do not closelv correspond 
with cuiTent faunnl ai'cas, although there is a very cloee correlation 
in some cases. An avian fnrniatioa may. in general terms, be coonid- 
ered the anal(^ue of a vegetational fonuation, altliough this do^ not 
imply tbat they necesnarily huve the same boundaries. 

As the literatare treating (►f the \-egetation of tlie»e aivas is extensive 
and scattered, a few papers will be rited as* an index to others: — 

1. Arid Deserta; Bray, '06: CoTlIle and MscDougal. '03. 

2. OrasHlandB or Plains; Clements, 'OS; Pound and Clements, '00. 

3. Southeastern Hardwoods: Cowles, '01; Harper. '06; Transeau. '05. 

4. Eastarn Canadian Conifers; Wblttord. '01; Transeau. '03, '05-'06: Oanong. 

'03, '06; Harvey, '03. 

5. Rocky Mountain and Pacific Conifers; Wbltford. '05; Gray and Hooker, 'SI; 

Piper, '06: Young, '07. 

6. Alpine; Merrlam, '90, '99; Covllle, '93; Fernald, '07. 

These environiueutal nnit areas a« found to-day, are the result of 
many succesaions which, in some vase» at least, reiu-li i-atber far back 
into the past. This is because some m'<-upy aneient land areas, such 
as much of the Houtheiistern Hardwood area. On the other hand, some 
occupy relatively new regions, that is, at least with regard to the dom- 
inant factors no*- in contnd. as in the glaciated part of Xorth America 
and on the <'o4istal I'lain. Ho far as the present is <'on«-emed wm'h rela- 
tions clearly show that these ai-eas are only the end results of extensive 
past changes or snccetJiHions which represent the termimi) branches and 
cross sections of development. It is to the study of snch ivgioiis and 
associations that we must turn for the fundamental organisation or 
nssociational relations of the various elements which compose not only 
the environments but also the associations of animals. 

In order to make as definite as possible the structural and rcoloffical 
characteristivs of these formations, certain general relations are here 
formulated. Throughout this paper it ^nuld be remembered that the 
individual birds and associations of given areas form tlie units of com- 
parison. Such a distinction is necewary because many species show con- 
siderable geographic vai-iation in habits and in the habitats frequented. 
The writer clearly recognizes the risks and difficulties of siu-h an at- 
tempt. They are delibenitely pnt in their present form to imitr criti- 
cism omt qualification from field workers. It is desii-nbJe to know the 
validity of these formations, their internal e<-ological relations and dy- 
namic tendencies, their relation to dominant envifonniental influencea. 
etc, A complete list is not attempted, and some of the statements may 
he only fragments of hirger generalizations; but it is just snch i-ehitions 
as these which will develop if the entire subject is considered critically 
and synthetically. Kome of the leading characteristics of these larger 
environmental nnits and their avian fontnitioiiH may be briefly outlined 
as follow^s: — 

1. The dominance of a limited number of ph^-sicnl conditions or 
complexes, as climate, topography, vegetation, aninuils, etc., in a given 
area prodwes the iH-imary envii-on mental units and formations. 

2, Secondary environmental dominance is shown b.v a secondary avian 
association. Thus in the Xortheastem biotic center there is a second- 
an' dominance due to wafei* l)asins in the forest area. ,- > 

i,C.ooglc 



132 MICHIGAN SURVEY, 1808. 

3. A formation or climax society is compoeed of a relatively (and 
usually absolutely) limited number of species wbich are dominant in a 
given environment of geographic extent. Such dominance, in general, 
implies extensive range, relative abundance, and ability to ind^nitely 
succeed or perpetuate iteelf under given conditlona. 

4. Where dominance obtains, avian variety is limited so that the 
greatest div«^ty occurs where local influences prevail, and at the mar- 
gins of the formation. 

5. Correlated environmental and biotic dominance produces That 
may be considered a iiotic base, stratum, or optimum, from which de- 
partures may be considered less favorable. This is a relative equilibrium, 
resulting from complete environmental and biotic adjustmetit, under 
gi\-en conditions. 

6. Tn each formation there is a normal inter-adjustment of the avian 
species and indiN-iduals, in addition to the adjustment with the domi- 
nant physical environment. The former is dominated by their straotnre. 
habits, and Ihe instincts or behavior; hence the colonial breeding or 
spacing, migration, etc. 

7. Each large environmental area or formation tends to have a full 
complement or set of species, of div«-Be but supplementary ecological 
character, such as water, shore or inland birds. One set is likely to be 
dominant. 

8. Relative stability in an association is correlated with the climax: 
dominance, and generally with extreme and slowly changing local in- 
fluences. Fluctuation is correlated with intermediate conditions. 

9. Diversified aasociatione and isolation are greatest with imperfect 
dominance, but dominance itself produces isolation of the climax as- 
sociation. This diversification produces associatitms surrounded by 
others and hence their isolation. 

lU. The taxonomic elements in different formations vary much, but 
there are close analogies in the kinds of taxonomic and ecological groups 
in different formations. — as the Mniotiltidte of the New are repre««itecl - 
bv the Svlviidae of the Old World. Cf. Osborn '02. LeConte, '50, 
1>. 239. Cf. No. 7. 

11. The roughly zonal arrangement of societies about the climax 
society (formation) or the environmental optimum, is primarily due 
either to local reversals, the lagging influence of local or neutral condi- 
tions, or to the influence of adjacent formations. This is a result of 
the retardation of the complete cycle of successions. 

12. The primary environmental conditions tend to encroach upon 
all others. The local conditions thus tend to become transformed in 
the direction of the dominant environni«it and to be appropriated by 
it. The corresponding avain associations are thus given a deflnite 
dynamic trend. 

13. The mobility of birds during the breeding season is very generally 
overestimated. The presence of the nest and young renders tbem for a 
time relatively sedentary. There are many causes influencing this, such 
as other individuals, proximity of food for young, homing, instinct, etc. 



3y Google 



ECOLOOT OF ISLE ROYALK. 133 

IV. MiNOE AVAIN EXVIKONMEXTS AXD THBIft ASSOCIATIONS. 

We have seen that the larger geograf^c environments or formations 
are characterized by definite conditi<ms and associations, and at the 
same time that even throughout these favorable r^ions the climax as- 
sociation is not distributed with absolute uniformity because of local 
variations in the physical features, such as vegetation, water basins, 
streams, mountains, etc. For the student of local bird life the real work 
begins when one attempts to examine into the causes and influences ex- 
erted by these conditions which break the monotony of the formation 
and make possible a diversified avifauna. But birdfi do not always re- 
spond as closely to slight local influences as does the vegetation, and for 
this reason one must learn by experience just what size of units must 
be used. Thus in the forest a few wind-falls will attract but little 
attention, bat a bum of a few acres will have a noticeable influence in 
harboring those species of birds which frequent openings; while swifts 
and swallows ignore many local influences which dominate other species. 

It should also be noted that whenever possible it is of distinct 
advantage to examine all habitats in their original state, uninfluenced 
by man. 

Instead of discussing the leading features of local conditions and 
their societies or associations in detail, only an outline of them will 
be given, and that in a form to facilitate use and revision, 

1. ftfinor environments are primarily dependent upon local conditions, 
and are thus in a sense correspondingly independent of the dominant 
forces of the region. This is, of course, a relative condition. 

2. Minor environments are, as a rule, relatively limited in area. 
In general their limited area favors their short duration, but age is 
primarily a result of the rate of change. 

3. Marked isolation, even when of extensive linear extent, — as a 
shore line, along a streain, or an elongate rocky ridge, — is also character- 
istic of minor environments. 

4. Minor environments tend to become encroached upon by the 
dominant regional influences and ultimately to be<'ome extinct. The 
succession of societies in local habitats is a declining one, while that 
of the geographic or climax habitat is an increasing and ascending <me. 

5. Local habitats produce most of the variety within the dominant 
area, and make possible a diversified avifauna. The structural dif- 
ferentiation within a formation (zones, etc.) is thus largely, in addition 
to variations in the formation itself, of local origin. 

6. Local associations or societies, in general, furnish the essential 
clues aa to their earlier successions which have attended the evolution 
or development of regional dominance. The variations in these are due 
both to the kind of life and to the influence of adjacent associations and 
centers of dominance. 

7. Marginal societies are particularly liable to variation in com- 
position, due to the combined influences of adjacent formations or centers 
of dominance as well aa to local conditions. 

8. Comparative studies of local habitats will form the most general 
and practical guide in the determination of the successions in the forma- 
tion. 

9. Local habitats and societies, in common with the larger environ- 



MICHIGAN SURVEY, 1908. 



mental complexes, are characterized by the doDiinance of few phyeical 
and biotic factors, and by a limited Dumber of epeciee. 



V, AVAIX SUCCESSIOX. 

1, General Remarks. Since the b reeding i^onnd a are fundamental 
importance in the ecol(^^ of birds. the"sTudy of tliem in such, situations 
furnishes the Rreatest source of innight into tiieir life relations. By 
an avain association, formation or soeiety is meant different combina- 
tions of species wltich reffuJarlij occur together in the mine breedinff 
habitat or area. These breeding {^rounds must be considered broadly, 
and include not only the nesting site but also the feeding grounds, 
even when they are jihysically very different, bemuse e<-ol<^ically these 
conditions foi-m a unit dui'ing the breeding season. 

It is well known that when a given set of physical conditions are 
dominant, as in a dense conifer forest, a swamp or an extensive orchard, 
relatively few individuals and kinds of breeding birds are charactOTstic 
of such conditions, except in the case of those nesting in colonies. The 
field relations of these colonial and isolated bi-eeders are quite different. 
It is also of imj>ortance io i-ecall that nbundam-e i« a relative term, with 
a very different meaning in the case of seed-eating and predaceous 
species. 

Bearing in mind these conditions, bird succession means a change 
fi-oni the dominance of certain species or associations to that of others. 
Thus in the beginning a slight change in abundance of a species may be 
noted, with a corresponding decrease in another; and this proportioo 
may continue to change until the intruder becomes dominant and the 
rival fojrm may disappear entirely. Tliis process of change, as a rule. 
is not limited to a single 8]>ei-ies, but usually involves several or all 
of the members of the association, as when a dune invades a swamp 
and the swamp birds are completely replaced by those fi-equenting the 
sand dunes. 

2. Succession on Isle Royalr. With these preliminary considera- 
tions in mind, we will turn to the ecological succession of bird life 
upon Isle Royale, I>ake Kui>erior. The field work ujion the island was 
CiU-ried on by a party from the University Museum of the University 
of Michigan, under tlie dire<-tion of the MTiter. Aside from succession, 
the general ecol<^ical relations of the birds were studied by Otto Mc- 
Creary and Max M. Peet, and elsewhere detailed descriptions of the 
i-egion and detailed notes will be published. The TVTiter has based his 
main retoi-ds of habitat preference upon their work. For this outline 
of suc^-ession only 'the primary features of the location need be given. 

In the present treatment an attempt will he made to follow the 
genetic suc(ression, at least in its broader outlines. Vanous qualifica- 
tions and reservations have been made, and otliers will follow, so it 
is hoped that no confusion will be produced by this method of treat- 
ment. 

Oeoijraphlcalhi, Isle Hoyale, Michigan, is an island in Lake Superior, 
near the North Shore, not far from I'ort Arthur, Ontario. The topogra- 
phy forms a pait of nu ancient peneplain of niodei'ute I'elief, glaciated 
and with an abundance of elong-ated low ridges and valleys with numer- 
ous water basins. Tlie >toil. which is locally absent, is generally humic 



ECOLOGY OP ISLE ROYALE. 135 

or mixed in ohaTnctep, boi-dering and in the depre«siona; but is mineral, 
stony and reaidual elsewhere. The combined shore and beaches are 
extenaive. largely stony and gravelly, and contain but little aaud^ mncb 
of the Hhoi-e line is rocky and precipitous; many outlying islands. 
Vegetation, herbaceous in shallow inland waters and a» a ground cover 
except where the shade is too dense, and upon rocks; shrubs on pro- 
tected beaches, in moi-e open places in the forest and in bums; the 
forest consists of Tiimarack, Black Hpruce and Arbor Vitje in bogs; and 
elsewhere in inesophytic conditions of Balsam Fir, Arbor Vitie, White 
and Yellow Birch, and rarely Hugar Maple, I'pon the drj- ridges, Jack 
Pine; and in bnnied areas, Aspen and Paper Birch. Climate, seasonal 
changes very pronounced; winters very long and cold, and summers 
short and cool; a relative humidity of about 80% in December and 
of about 7ttfo in July (cf. Jofaoson, '07) ; a mean temperature for 
January 7.97° F.; and' for July, I>2.:i4= F. (Port Arthur data). Early, 
deep snows. Pntlartom animals, as the Lynx, Marten, weasels, Red 
Squirrel and bats are directly in competition with the birds for food, 
or prey upon the birds. 

The alwive environmental fartorx are dominant featui-es nnd gi^-e us 
a general pictnre of the conditions, largely in terms of common ex- 
perience. In the life of the birds, however, a complete reassortment 
and change of intensity in these factors occurs when they are com- 
bined as habitats. The suiTOunding lake, the nnmerous baj-s, small 
lakes and ponds compose the aquatic habitat and make it a characteris- 
tic feature. The very iri-^ular and extensive shore line and limited 
beach ai-ea characterize the coastal border, while inland, excepting the 
main bodies of the few larger lakes, the encroachment of the bog vegeta- 
tion upon the shores is such as to prevent an extensive development of 
sandy open beaches. The above mentioned habitats are oi>eD unforested 
areas; the remainder of the island, with the exceptions of the bare 
rocky ridges, the clearings and burned over areas, are fJS^ed. Very 
extensive swamp forests abound in the elongate valleys and the borders 
of the water bodies, and are composed of Tamarack, Black Hpruce and 
Arbor Vitse. Tlie uiesophytic forest occurs on drained ai-eaa and is 
characterized by Balsam Fir, White Spruce and Pai)er Birch; the 
bui-ned areas by second growths of aspens and Paper Birch. Then tliere 
are also influences which are exerted upon the bird life in general, as 
for example, uitgration. In this case, undoubtedly both external condi- 
tions and the habits and the behavior must be correlated. Another 
general and dominant influence should be reiterated here, and that is 
that all open areas tend to become invaded with vegetation and finally 
forested, whether they are lakes, ponds, bogs, rock openings on the 
ridges, bums or clearings. The mes<»phytic Balsani-npnu-e foi-est tends 
to monopolize all habitats, and gives a definitenefs lo all succession 
upon the island. 

From a genetic standpoint the past and present dominance of the 
surrounding I^ake must be recognizad. This formerly stood at a level 
much above that of the highest ridges upon the island, as is clearly 
evidenced by the abandoned beaches on the north shore of Lake Superior. 
Such relations prove that Isle Royale was once a rocky i-eef in the 
lake, which, as the l^ake level was lowei-ed (it is quite unlikely that the 



136 MICHIGAN SURVEY, 1908. 

islancl has been materially elevated) became exposed as a wave-washed 
beach. These conditions are appix>xniinted to-da.v by the low outly- 
ing islands. The beach or shore is thus the original habitat upon Isle 
RoyalCf and in general, all others have been derived or developed from 
it. To discnse these as a truly genetic series would require that, these 
be described ximtiltnncoualij, as the differentiation took place. These 
habitats did not develop as isolated phenomena, but several developed 
at the same time, or abreast. Tluis as soon as enauffh of the land sur- 
face had become exposed so that its inequalities b^n to have an in- 
fluence, the ridges would be the parts best drained, and certain depres- 
sions would tend to nccuiuulate the drainage. This process would lead ■ 
to a simultaneous development or differentiation of the well, moderately, 
and poorly drained habitats. Almost all of the residual soil formed 
as the region was haseleveled was prabably cleared away by the glaciers; 
or later, as the waves fell from the island, by the pounding of the waves. 
Thus the relative absence of a soil must characterize all habitats. At 
what period life first reached the island in post-Glacial time is not 
definitely known; but it is likely that the pioneer vegetation of lichens, 
mosses and low herbaceous vegetation reached it soon after its ej-posure. 
If the biota reached the island about the time of the formation of the 
Algonquin beach, which, i-oughly speaking, may have l»een nt about 
the present elevation of 475 feet above the Lake surface, it has since 
spread upward and do\i-nward from that lei-^l. The composition of 
the initial societies is not liable to as much variation at the later ones. 
Thus if the Herring Qulls returned to the region at this early period 
of the exposure, they were probably the pioneer birds; but if only 
at a much later date, still other species might have accompanied tbeui. 
While such variations as this may be ex|>ected, and due allowance must 
be made for them, yet there can be little reasonable doubt but that 
water birds and those frequenting open- places' tended to become the 
pioneers, and that later, with the development of a soil and forests, 
other associations of birds became established. 

There are at least five important factors which enter into the com- 
position of the past and present conditions which have moulded and 
are even now moulding the formation of the habitats upon Isle Royale. 
These five are:— first, past climatic changes; seconrZ, the local topogra- 
phy; third, the falling lake surface; fourth, dynamic tendency of the 
vegetation; and fifth, the habits and structure of the birds. With these 
guiding principles, let us now turn to certain details of the resultant 
succession. 

a. The Aquatic Association and Hahitat. 

Tlie expanse of Lake Superior, the irregular shore line producing 
coves, the inland water bodies and streams, together furnish an ex- 
tensive and expansive area of habitat. The cutting of the Lake waves 
enci-oaches ui)on the land habitat, and the deposition by them elsewhere 
i-auses minor extensions of the land habitat (as at Kock Harbor where 
a sand spit furnishes a nesting site for a Kingfisher). Inland the 
encroachment of the vegetation tends to restrict the water areas, as the 
falling Lake level has, in the [Mist, tended to increiise the land habitat. 
These processes must l»e recofmized in order to grasp the dynamic ten- 
dencies of the hahitat. 



ECOLOGY OF ISLE ROYALE. 13? 

The characteristic aquatic society la composed of the Herring Gull, 
liOOD, American and Hooded AfergaQsers, and ttie Pied-billed Grebe; 
mainly fish eaters and scavangere. Other species, of greater inland 
tendencies, are attracted by the flsh food, as the Eagle, Osprey and the 
Kingflsber. The GuIIa show a decided preference for the great Lake, 
and the Loon for the inland waters. The presence of tiie Kingfisher 
was influenced by the harbor with its attendant sand banks and bars. 
As all these water bodies near Isle Eoyale freeze over in winter, the 
strictly aquatic birds must normally migrtLte to secure food. Of course 
none of these birds nest in the open waters, but on the island beaches 
(Gulls), near the moutfas of streams, and inland in marshy places; but 
all, as a rule, nest near the water. The" very young soon attend tbeic 
parents, and are thus in the water at an age when many laud birds are 
yet helpless in tlie nest, thus confirming their aquatic habits and habi- 
tat. During migrations many other species frequent this habitat. 

Where Isle Royale now- is, once rolled the open Lake; and it is not 
improbable that as the islaaid appeared the Herring Gull was oue of the 
first species to discover it. Such a bird might even reach the island under 
climatic conditions of the Ice Age, for the species now ranges far north 
along rlie shdre of the Arctic Sea. A sjiecies of such extensive chronolog- 
ical and geographical range will tend to give much stability to suc- 
ceeion. The present range of the Mei^ansers and the Loon is not so far 
north, and for this reason they may have arrived under milder climatic 
conditions. But if the island l>ecame exposed under mildpost-Glacial 
conditions, all of these species may have arrived at much the same time. 
But even with the chances for such variations the general succeesion 
seeiiifr to have been initiated with the aquatic association as the pioneer 
society. 

In following the genesis of the habitats and associations from this 
point onward, divergence and differentiation becomes so marked that 
it is impossible to develop all lines abreast. A linear treatment be- 
comes necessary, and therefore certain general relations are liable to 
become obscured unless specifically mentioned in advance. 

The aquatic and beach habitats possess a mariied tendency toward a 
zonal arrangement. From the Superior beach the transition is through 
open or ^mb zones into the climax forest. The topography of the 
island with its longitudinal ridges and valleys form a dominant factor in 
impressing this zonal structure upon the biotic associations. The series, 
— from the water, through the beach, open and shrub marginal zone, 
into the climax forest, — may be considered as the genetic vegetable suc- 
cession. They change simultaneously and are due to the same general 
cause, — the falling Iiake surface, which transforms the water area into 
beach, the beach into forest margin, and forest mai^n into the climax 
association. But as mentioned, it is manifestly impossible to discuss all 
these transitions at once, and each ecological unit must therefore receive 
separate genetic treatment. 

This tension line or marginal zone between the Lake and the forest 
shows such a wonderful diversity and complexity in its conditions, 
that several plant and animal associations are formed within this zone. 
With its onward march there are simultaneous changes in several asso- 



138 MICHIGAN SURVET, 1901. 

riationn which, while tliey will vary iu their cfaafajjoK, yet all tend to fon- 
verge iu harnionv with the dominant factors. Tliene. conditions migrate 
or radiate from the hijfliest land. On the other hand, the inland marginal 
zoneB, which border the Bnialler water bodies, inigi*ate inwardly; anil 
being closed areas, tend ti> become extinct. This mar^imrl zone, parti- 
cularly beyond the njiper beacii, forms one of the most interesting and 
complex conditions found upon the island. It is not an ecological unit, 
but is comjKised of several of them. This is where most of the eon- 
fusion arises in actual field work of habitat studies. 

b. The Shai-e ant! .Ifirsft Axuoriation nniJ Habitat. 

As the area of the islands ex[>anded and the shoi-e line was leujitheD- 
ed, the habitat for sbore birds inci-eased : but the stee|» and rocky shores 
M-ere unfaivorable for the development of Iteachett because loose ro<k, as 
tools for the waves, was limited in amount. The local character of the 
shingle and gi'avel to-day found in the various coves clearly indicates 
their local origin; and much the same conditions have obtained iu the 
past. On acconnt of ttiese conditions, tbe sandy loaches ore very con- 
spicnously absent. The dynamic tendencies of the beach are those 
whi<;h cause the extension or restriction of the aquatic and beach 
habitats, supplemented In' the drift whicJi is tossed upon the sbot«. 
A\Tiere there is shallow water, and mud accumulates, favorable condi- 
tions are furnished for invei-tebrate food for birds. Inland, the numer- 
ous lakes, ponds and marshes furnish shoi-e conditions which tend to 
become extinct through drainage or overgitiwth of the vegetation, except 
in those |>arts of the larger lakes where wave action tends to scatter 
such accumulations as rajiidly as formed, or to pi-event its formation 
altojjether. 

Although observations on this subject are <|uite limited, yet it seems 
fairly safe to consider the Sp<itted and Solitary Kandpii>erH as char- 
acteristic birdx of this association. TT|H)U such a i*ocky coast, sandy and 
gravelly iH-nches are r|uite exceptional and are conflnetl to protected 
coves. Additional diversity is produced where small streams enter 
tlicHe coves and produce deltiia 

Little is gained by sharply segi-egating the marsh and shore birds, 
althougli the marsh birds show a preference for conditions better 
represented or correlated with tO]K>graphically older coasts, pro- 
tected and inland conditions. Attention should be directed, how- 
ever, to the signifliant fact Ihat smTcssions initiated with such 
diversity will produce a variation in the composition of the associa- 
tions. Also that so far as possible these variations should be -coo- 
sidei-ed couipanitively and synthetically in reconstructing and antici- 
pating successions. 

The American Bittern, T.«Bser Yellow-legs, Swamp Sparrow and 
Marsh Hawk belong to this society of marsh birds. As in the case of 
the aquatic association, these birds generally nest in close proximity 
or entirely within these shore or marsh, conditions. Still other species 
frequent this belt to feed, as it is aa open area; but their presence is 
mainly conditioned by the adjacent shrubs or forest. The very limited 
number of s|>edes it) the aquatic amd shore associations is worthy of 
particular mention. 



ECOLOQT OF ISLE ROTALB. 130 

The Yell«>w-leg8, Spotted Sandpiper, Bittern aod Mareh Hawk range 
far to the north, even to the Barren Grounds, and thus Buggest chances, 
as in the case of the aquatic aMociation, of an early arrival and suc- 
cession upon the island. 

With the growth of the island, there has been a oorre«ponding ex- 
tension of the outer and inner shore habits, although the eDcroaching 
vegetation has had a marked tendency to restrict the area of the inland 
habitat. The dominant enYironmental inSuences in this habitat appear 
to be, 1, the physical character of the shore and beaches; 2. the dynamic 
forces of the water bodies and sti-eams; 3, the encroachment of the 
1-egetation; 4, the downward migration of the shore; and 5, the habits 
and structure of the birds. 

As a general rule, we may say tliat the beach of the outer laice 
tends to be- succeeded by either the bog or upland associationa, and 
those inland by the bt^ nssociation. 

c. Bog-forest Association and Habitat. 

As just stated the outer coast or an inland one may develop into 
a marsh or hog habitat or association. lu the bog. the Tamarack, 
Black Sprace and Arixw Vitie are the pioneer trees in traosformin*; 
the open marsh into a forested one; while upon the outer rfiore the 
alders and aspens tend to precede the conifers as a general rale. 
From the bog forest the transition to the Balsam-White Spruce forest 
may be perfectly continuous, and thus there will be a series character- 
ized by the dominant conifers. In places Arbor Vitffi may form the 
dominant swamp forest, but this is only a variation in the conifei: domi- 
nance. With improved drainage or the accumulation of ve^ietable 
debris, these habitats become converted into the Balsam-spruce climax 
foreet and hence the environmental dynamic tendency. 

As the forest encroaches upon the open bogs the Tamarack, Black 
Spruce, Arbor Vitie, Cassandra, Labrador Tea and alders are accom- 
panied by birds characteristic of this early stage; such as the Eed- 
breasted Xuthatoh, Yellow-bellied Flycatcher, Golden-crowned Kinglet, 
Cedar Waxwing, Chickadee, Canada Jay, White-winged Crossbill. 
Where alders abound the conditions are favorable for the Redstart and 
the White-throated Sparrow, But later, as the bog conifer forest 
becomes continnons and dominant, the ^'axning. Redstart and White- 
throated Sparrows diminish in numbers and finally disappear. Still 
later, as the swamp becomes eliminated with the development of the 
climax forest, the Yellow-bellied Flycatcher will also become excluded. 

This is perhaps the simplest suceessioo from the water to the climax 
forest, via the bog forest. This series is very perfectly presented in 
all stages and has an extensive range. The number of species In the 
association is rather large when compared with the preceding asso- 
fiattons. 

d. Aspeniinh Association and Hah'tat. 

This series develops from the beach and the waves fall from the 

ridges or low rock surfaces and leave the bare ex[)anBc«. As the rock 

disintegrates, decomposes, and humus arcumulates, a soil is formed, 

mainly in deprefwioiiR or nt Hie buses of the ridges, and from these 

19 



140 MICHIGAN SURVEY, 1M8. 

it tends to encroach upon the open places with a zone of Jack Pine, 
aspens, or White Birches. These areas are largely strips along the crests 
of ridges or small park-like openings on rather level rock. In no case are 
these single areas large, so that the habitat is only extensive in the 
aggregate With the presence of the open aspen and birch woods, the 
following society is likel; to be characteristic : — J unco. Oven Bird, Red- 
eyed Vireo, Chipping Sparrow, White-thnmted Sparrow, Flicker, Cedar 
Waxwing, Wilson's Thmsh and the Chickadee, As the deciduous trees 
are replaced by the open encroaching ccmifer forest, the Song Sparrow, 
the Nashville, Myrtle and Black-throated Green Warblers and Wilson's 
and Olive-bflched Thrushes, which frequent the forest margins, increase 
in abundance. The Oven Bird has an extensive northern range from 
Labrador into the Yukon Valley and may well have been a very early 
pioneer upon the island as the aspens and birches were probably the 
first broad-leaved tree arrivals. From the above it is seen that this 
means an extensive vai'iety, but as the dominance of the climax foi-est 
encroaches this number again becomes reduced. 

I'he composition of the society varies somewhat, depending upon 
the surroundings, as proximity of the present shore or distance fi-otu 
it. Many of these openings are continuous with the present beacli. 
It is not improbable that this was a prominent society whenever the 
waters fell rapidly from the island between rather stationary levels. 
This has beoi a society decidedly on the decline with the eocroachment 
of the forest. 

Probably this association varies considerably in its composition, and 
bas done so in the past; but its main features are fairly constant. 
These variations seem likely, through the influence of openings pi-o- 
duced by fires which, when extensive, may have caused a new equilib- 
rium among those species frequenting openings. 

The Burned Area Association. 

This phase should perhax>s be considered as supplementary to the 
aspen-birch association just considered. A fire brings about a reversaT 
of conditions through the destruction of the forest, and in some, cwses. 
a part of the soil as well. As there are all degrees of extent and com- 
pleteness in this process, thei-e is a corresponding \-ariation in the 
detiiHs of the resulting succession, at least in its early stages. Jt is 
on'y when there is a very complete destruction of the v^etatlon that 
the continuity with former occupancy is wholly broken. 

The easily inflammable character of these conifers, even when in a 
green conditicm. makes it likely that natural causes, such as lightning^ 
or marsh gas (cf. Penhallow, '07), may have been influential. The 
proximity of the gas supply and the conifers is of interest as this may 
influence their liability to flre and thus to this sort of reversal of con- 
ditions. Thus liability to flres is rather characteristic of the r^ion, 
and man's influence has tended merely to reinforce rather than to intro- 
duce this feature. Thus it seems probable that fires have been a factor 
in supplementing the natural park-like openings. In addition to the 
burned areas found upon Isle Boyale, other limited open areas are due 
to cultivation and are kept open. 

The birds characteristic of the more open situations are the Sharp- 



ECOLOGY OF ISLE ROTALE. 141 

tailed Gnrase, Song and Chipping Sparrows, Flicker, and the Pnrple 
Finch. The Grouse is a Plains form, is near its eastern limit, and is 
pertiaps a late arrival npoo the island. The other specieft are wide 
ranging in the Canadian coniferoas forests bat are not of sach northern 
range' as the agnatic and shore associations. There is nothing in their 
range to sn^jest their arriral earlier than the forest association. Tak- 
ing all the birds of the openings together, it is not improbable that 
they arrived at about the same time as those of the forests, hxit fre- 
quented different situatione, — tte forest kinds occupying the slopes 
and drier mlleiys, and the others the openings. 

e. The CUmax Association or Formation anA Eahitat. 

The climax Rssociafion should not be considered in such a way h» 
to lead one to think that it is distinct from the other associHtions. It 
belongs to all of them as the end of their series under existing biotic 
and environmental conditions. Thus the aquatic association, througli 
the bog conifers, is transformed into the Balsam-spruce association ; 
and from the beach through the aspen-birch association again to the 
balsams and sjvnces. The -climax association is the conditimi of ad- 
justment toward which all societies move under the present conditions. 
For this reason the earlier stages, conditions and associations of the 
climax have been outlined in the preceding dlscussicm. 

In the dominant forest the dense shade prevents an extensive ground 
cover of herbaceous plants; and although Ground Hemlock is abundant 
locally, yet in places the forest floor is quite open and free from lower 
shrub growth. The remarkable preseri'ation of trails or roads throngh 
such tracts shows clearly how slowly changes take place. Such a. 
habitat must be relatively equable in its temperature and moisture 
relations. 

Gec^raphically speaking, the primary characteristic of the climax is 
its relatire atahility, due to a dominance or relative equilibrium pro- 
duced by the se^-ere environmental and biotic selection and adjustment 
throughout the process of succession. 

At this point attention itbould be called to the fact that dominance 
is a resultant of an equilibrium produced by neutralizing or overcoming 
other forces and influences. We may think of the process of successioa 
as a stream of forces whose development may be <'ompared with the 
transformation of a drainage line, — such as, for example, that of a 
rivulet into a creek, and then into a river. The stream and the char- 
acter of the ground mutually infinence each other and the course fol- 
lowed is a resultant of the mutual adjustments. The stream is deflected 
by one condition and then another, just as succession varies with local 
conditions; yet the water continues to run down grade and se^s an 
eqnilibriom, and similarly, biotic succession continues on its course de- 
flected here and there by local influences, yet forever tending toward ii 
state of biotic equilibrituii. The dominance of th^ climax society or 
formation, considered as a pro<-eBB rather than a product, has much in 
it that is analf^^oos to the dominance produced by the process of base^ 
leveling. 

The characteristic birds of the climax forest are: — the Chickadee^ 



142 MICHIQAN SURVEY, 1906. 

Golden-crowned Kinglet. Red-breasted Nnthatch, Canada Jaj, Downy, 
Hairy, Arctic Tliree-toed and Pileated Woodpeckers, and the White- 
ninged CroBBbill, Here again the aasociation becomen email in \-ariet^' 
of species and comparable with the small society which must have be«i 
associated with the complete dominance of the Lake waters. TIiub there 
has been a development of diversity from simplicity, with later a return 
to simplicity. To tliese birds of the forest should also be added those 
species of general distributiouv as the Eagle, Swift. SwnJlows, etc., a 
class of birds whose predaceous, insect feeding and wide ranging habits 
make them particnlarly difficult to properly asBooiate. A careful etudy 
of this class of birds will be necessary before they can be satisfactorily 
correlated with their proper avian associations. 

But let us not overloJ* the fact that even this dominance is only 
7'elative, for since the Ice Age even tliis entire formation has migrate*! 
northward, and a true succession has been produced with its attend- 
ant changes in the conditions and in the composition of the associa- 
tions. Just as upon Isle Roya]e a definite dynamic trend was given to 
tiie complete environment by the falling Lake surface, so ia the post- 
fllacial northward migration there was a northward migrating climate. 
These conditions determined that on the north aide of thia immense sue- 
cession or migration habitats and associations were developed which 
are comparable to those attending the downward marcli of the Isle 
Boyale beach ; and eveji today, by passing from Isle Koyale to the tree 
limit with its zone of aspens and birches, one may find representatives 
of the various kinds of associations which in all probability moved north, 
just as today in passing from the forest to the rocky beach balsams 
and »pruce are encountered before the aspens and birch. If, however, 
this is only another case of convergence and not at bottom the same or 
a comparable process, we are then certainly far from an tmderstanding 
of even the general nature of the problem. 

3. Internal Factors. With the idea of succession, as exemplified 
by Isle Hoyale, let us turn to other factors which influence the internal 
relations of the birds within an association or society, because such 
relations are also necessary to an intelligent understanding of suc- 
cession. Bome of these general relations have been outlined, but certain 
others are needed which have been well expressed by Breivster ('06, p. 
fi2-fi3) : "Many if not most birds show a nmcked preference for breed- 
ing in certain regions, throughout which they are more or less evenly 
and generally distributed, but within which their munbers do not seem 
to increase beyond fixed maximum limits no matter how carefully tlie 
birds may be protected or how sncccKsful they may be in rearing their 
young • • • J iiavp observed — as, indeed, who has not I — that few 
birds — excepting those which, like Swallows, Terns. Herons, ond Gulla, 
are accustomed to nest in colonii-s — tolerate very near neighbors «-f 
their own species during the season of repi'odnction. At its beginning 
each pair takes pofisession of a definite Inict of wootliand. orchard, 
swamp or meadow, which the male is ever on the alert to defend against 
Irespassera of his own kind aud sex, although he often seems quite 
willing to shai-e liis domain with birds of other and perhaps closely iv- 
lated species. The rxteut of the aiva thus monopolii^ed varies exceed- 
ingly with l>ird« of dilTercnf 8i>ecles. M\ apple orchard which suffords 



BCOLOaY OF I8LB ROYALB. 143 

SQfflcieat room for — let ua siay — two pairs of Yellow Warblers, two 
pairs of orioles, three or four pairs of Chippies and four or five pairs 
of BobiDB, seldom or uever hnrborfi more tlian a sini^le pair of King- 
birds or crested Flycatchers. • • • As a nilf, the species which 
roam wer the most ground in the course of tlieir daily wanderings 
claim and maintain the broadest presen-es, while those of sedentary 
habits often content themsches with very modest freeholds. Whatever 
the extent of the domain, the birds who occupy it as a sypimer borne 
evidently regard it as exclunively their own. The readiness *and celerity 
with which trespassing birds are accustomed to retire when attacked 
or wen merely threatened by the established tenants, has seemed to me 
to indicate that the claims' of temporary ownership are respected by 
all right-minded birds. • " ' In my opinion the desire for exclu- 
sive possession so conspicuously shown by the male, and often by him 
alone, is usually the direct result of sexual jmlomy. This, as is natural, 
makes him intolerant, during the breeding season, of the near presence 
of rival males. If his concern were chiefly in respect to the food supply, 
it would be equally manifested at every seoflon and towards all birds 
who subsist on the same food that he and his mate require— wWch Ik 
certainly not the case." 

The tendency of pairs and species to space tlicmgdves and to become- 
fclatircli/ sedentary is thus a cbaracteristic condition in an associaticm,. 
and is an important element in an understanding of succession because' 
it shows the internal organization and habit with which an invader 
or pioneer from another association has to contend. As Dixon ('97, 
p. 91 ) has pointed out, this spacing tendency is an important factor in 
the extension of range of species and is intimately related to the loca- 
tion of nesting sites. These facts clearly show that both these internal 
influences and the environmental ones must be distinguished if we wish 
to determine the relatii-e influence of each and their bearing on suc- 
cession. The above quotation from Brewster clearly shows that iu 
general not only a gi-eater number of birds can live in a given area, 
but also that they can live closer together, if they vary in kind. Then 
again, within the association there are marked differences in habitat 
preference. Thus in tlie forest there Are those birds which nest in 
the trunks or among the topmost branches of the trees, or even upon 
the ground ; and these are differences largely distinct from the spacing 
of the pairs of the same epecles. These influences must be recognized 
among the dominant influences within the association, and upon which 
much emphasis must be placed. 

4. Environmental Factors. Then in addition to these internal fac- 
tors, there are the dominant physical factors. In the following discus- 
sion primary emphasis will be placed upon succession as found in the 
Xortheastem Biotic or Conifer Center, because suc<'esHions at other 
centers with different biotic components and other dominant physical 
conditions must possess a certain amount of individuality, in addition 
to those features common to succession in general. The dominant biotic 
tendency or dynamic trend of this center, as a resultant of all internat 
and enpironmental infiticncrs. is for the conifer biotic association to 
encroach upon all other soeUticn and habitats and to become the donii- 
nant or tiniversalhf diatribiitcd association. Thus, in general, all 



144 MICHIOAN SURVEY. 1908. 

habitata produced by local influences tend to bec<Hne transformed into 
the dominant biotic association or formation. In general also, amaJl 
bodies of water are rapidly encroached upon by iun'ash, vegetation or 
drainage, and tend to become extinct and forested. All other openings, 
aa the rocky ledges and ridges or burns, are encroached upon as soil 
accumulates or fires are prevented, and the forest biotic association 
spreads over the entire area. 

From Bnctw^lations it will be seen that our knowledge of the causes 
and condititffs of succession must largely result from the study of 
these local environment it or habitats and thevr biotic succession, because, 
where dominance is establislied the succession is almost completely 
obliterated. Each minor habilal and society is to be looked upon an 
simpl}/ a stage, more or less temporary, in the onnxird taave toicard the 
■domiruint or climaa: association. Thus in the marshes, birch or aspea 
woods, rock openings and ponds may be "original" conditions which 
are becoming cumulatively transformed in the diredtion of the final 
dominance of the climax biotic type. 

Tbe relatively slow rate of change in many environmental processes 
anfl tbe relative stability of the climax biota,' is doubtless the basis 
for the current view that such conditions are relatively constant or fixed ; 
but that change and not constancy is the normal and usual condition in 
nature is quite evident upon a moment's reflection. Almost every (me no- 
tices these changes after an absence of a few years from a region. Thus 
intimacy tends to blind us to changes unless a habit of giving attention 
to them is deliberately cultivated. For this reason some find it almost 
impossible to recognize environmental changes or to comprehend thrir 
significance. Jt is therefore of practical value to recognize clearly 
under what conditions changes may be most readily perceived. There- 
fore the importance of the study of local influences is emphasized, and 
the necessity recognized of distinguishing the dominnnce of geographic 
and relatively stable conditions or formations as contrasted with those 
due to iocai and often relatively changeable conditions. Then among 
these changes we must distinguish those which are mere fluctuations 
and .those which are iadicati\-e of the true progressive succession. 
This is mainly aocomplished by attention to general relations and the 
subordination of minor details. 

5. Environimenlal and Associational Convergence. At the present 
imperfect stage of ecological development, comparison must furnish us 
the most important and general clues to the processes of succession ; and 
undoubtedly thin method must long remain as our main guide on ac- 
<tount of its comprehensive applicaftion and the magnitude of tbe prob- 
lem to be solved. It is tlierefore desirable that the limitations of the 
method shouJd be'clenrly borne in mind. It is often assumed that tbe 
implied successions of a given place are the same as those which have 
developed at that place in the evolution of the present climax. But 
as we positively know that many different causes are able to prod/uce 
the same or very similar results, such conclusions must be received 
with due caution. Tlrat the dominant geographic conditions tend to 
override local influences seems very fairly established because diverse 
local or original conditions are transformed into the climax or domi- 



ivCoogIc 



ECOLOGY OF ISLB ROYALE. 146 

vant type. This clearly ehows that in time di^-erse local iafluencea bare 
flowed into the general environmental tread or current and have become 
a part of it. There is thus a very strong convergent tendency. By 
convergence is meant the independeOt production of the same kind of 
asHDciation from 'diverse starting points or habitats and associations. 
Quite minor ecological units may show similar but temporary con- 
vergent tendencies in their succession. It is therefore not surprising 
that any marked environmental dominance will tend to produce simi- 
lar or convei^c^it results, even in local areas. Under such circumstances 
similar associations or societies may be independently and repeatedly 
formed by the selecting environmental influences, such as, for example, 
are found in the numerous small lakes scattered throughout the conl- 
feiious forests. This convergent phenomenon is certainly a fertile 
source of confusion throughout all phasea of science. Perhaps the best 
guide through such a labyrinth will be to clearly bear in mind the rela- 
tive valne of general and local influenoes, and watch with an "eternal 
vigilance" for convergent results due to diverse causes. This con- 
vergent phenomenon is particularly liable to occur in the case of en- 
vironments produced by reversible physical conditions. It should 
fm-ther be stated that a study of these problems from a genetic and 
dynamic point of view will aid in recognizing such results. Under such 
circumstances attention is primarily directed toward the dominant 
causes and conditions of change rather than to the stages, products, 
and results produced by tbem. Convergence thus viewed is the result 
of several causes and should be considered a product rather than a 
process. This same distinction may be made for all societies, associar 
fions and formations. Comiergent plienomena are thus partioularljf 
liable to confuse wherever products rathei' than genetic prooesaea receive 
primary emphasis, 

6. Succession and Envirottmental Evolution. The relation of suc- 
cession to general biological problems is very intimate. This opens up 
a very extensive field which is only mentioned to indicate its general 
relation to succession. The facts of succession and evolution must ever 
remain far in advance of our knowledge of their causes. If, however, 
one tnms to the standard evolutionary treatises and searches for a dis- 
cussion of the evolution of the environment, as correlated with animal 
evolution, only the most general, or the elementary and superficial 
phases, are as a rule discussed. To be sure, certain papers and treatises 
take up special phases of tlie problem, and the broadest phases are 
treated by the geologists; but none of them seem adequate as a com- 
prehensive treatment of so important a subject. Sucoession, broadly and 
genetically considered (dynamic rather than static), is a phase of en- 
vironmental evolution. 

7. The Relation of Succession to Organic Evolution. Mention baa 
been made of the releitiou of succession to environmental evolution, but 
its relation to the organic evolution of birds should also be indicated. 
The mutual relations of organic and environmental evolution have been 
and will continue to be the battleground of biological thought for an 
indefinite length of time. Here lies the tension line between the two 
main schools of biological interpretation. 

One school maintains that all causes of ei'olntion are internal, aiid 



146 MICHIGAN SURVET, IMS. 

that tbe enrironment is odIt n conditifMi, not a cause. From thia point 
' of view the fundamental causes are internal and therefore environmeiital 
conditlouH can only indirectl.v influence evt^lutlon tfaroogh the weeding 
out of those forms not in harmony with ttie conditions; and hence it 
has a elective rather than an oriifinative influence. Prom this point of 
view succesBion aod environmentnl evolution citn contribute nothings to 
the elucidation of the causes of organic evolution, though they may to 
an understanding of the eelection produced by the succession of condi- 
tions la which organic evolution has taken and is tailing place. In 
harmony with this point of view, sacceesion, iMvadly treated, should 
furnish a fundamental method of treatment for the process of selec- 
tion, and the detailed principles of its worliing. This would certainly 
be an important advance because natural eelection has frequently been 
reproach^ for its indefinite methods and lack of definite treatment. 
Succession from this point of view is primarily related to tbe Dar- 
winian factors of evolution. Xo doubt this ia one reason viiy Darwin 
himself put sucJi high value npon the 'study of ecological relations of 
animals, t. e., their relation to their complete environmeait, or their 
struggle for existence. 

If, however, all causes are internal and not directly subject to ex- 
twual influences, they must be beyond experimentation to a correspond- 
ing d^ree. Under such conditions evolution becomes a descriptive 
■ rather than a causal science, and all that investigation can do is to 
describe the succession of forms produced by these internal causes. 

On the other hand -tiie rival school maintains that both internal and 
external conditions may be real cavaes of oi^;anie evolution. This is 
thought to be brouf^t about by the direct or indirect influence of tbe 
environment vpoD the germ cells, by environmental selection, or even 
by both combined. From such a point, of view the environment may 
thus be either a cause or a condition of organic evolution, or both. From 
such a standpoint the evolution of tbe environment receives increased 
importance, as under such conditions organic and environmental e\'olu- 
tion are cansnally related, and thus intimately correlated. Viewed thus, 
.environmental evolution is more than the description of the succession 
of conditions, but may be explanatory as well. 

The particularly significant feature is that environmental evolution 
and biotic succession are of great value and can cMitribute either to 
the causes or conditions, or to both, of evolutionary advanoement. 

VI, SosiE Principi.es of Succession. 

By succession is meant the progressive change (,== adjnstm^it) in 
the composition of the associations at a giveji place. If a swamp be- 
comes filled with dute sand, the birds charax^teristic of the swamp will 
be replaced by those of the dunes, and thus succession is initiated. But 
in addition to changes due to local influences there are those pro- 
duced by very extensive or geographic influences, as in the case of a cli- 
matic change. Attention nfaonld also be directed to the fact, that biotic 
succession is only a particular phase of the general law of change 
which we see operating wherever a complex of forces are tending to- 
ward a condition of mutual adjustment. That succession is a process 



ECOLOGY OP ISLE ROTALE. 147 

whicb, from its very nature, muet be as extensive as m-e the caiueH of 
change does not appear to have been clearly recop^ized by all students of 
biotic succession. For this reason there are certain principles of suc- 
cession which are well establiBhed in other sciences, but which have not 
been applied to biotic succession. In human society, for example, there 
are many institutions whose formation, development and perpetnatiou 
clearly illustrate the laws of succession which also apply, not only to 
plants and animals, but to geologic phenomena as well. It is not at 
all Borprising therefore that under these various guises their common 
features are easily overlooked and even denied by some students. 

In t^ study of the animal environment some knowledge of the gen- 
eral principles of succession, not worked out in detail for birds 
but already well established elsewhere, ought to be suggestive anri 
possibly valuable in the study of avian Bucces8i(»i. Though such geoer- 
aJizations are primarily of a provisional and suggestive character, yet 
investigation should be stimulated rather than retarded by them. Bucb 
descriptive characteristics and principles are stated briefly in a form 
convenient for testing and criticism and should be useful as are cri- 
teria in the study of ge<^aphic origin. So far as known to the writer 
only two authors have attempted to formulate principles of biotic suc- 
cession, and these have been limited to plants. The first is by the Dan- 
ish ecologist Warming ('ilfi, Oekol<^i8che Pflanien Geographie, pp. 3611- 
361), and the second by Clements (*05), whose treatment merits special 
attention. Cowles ('01) has done much to put the idea of succession 
upon a goietic basis. In the present outline only those features and 
principles are mentioned which are thought to be of a more or less 
general chairacter, and those particntaily applicable to animals. This 
list needs to be greatly prolonged, and the interrelations of these char- 
acteristics must be determined as well as their relative value and appli- 
cation to various ecological gi-oups and in diverse regions. The fol- 
lowing suggestions can only outline the problems involved. At this 
stage, differentiation is particularly desirable. Processes and pro- 
ducts bear the same names and muet be understood accordingly; thus 
the proceasee of dominance lead to ibe product dominance. Dynami- 
cally considered, the process is primary', but used in a structural sense 
such terms i-efer to products, 

1. Starting with any given 9et of environmental conditions and or- 
ganisms, theee become a cause and condition of futui-e changes. All 
changes are cumulative and form a continuous series or process. 

2. Ko sharp line can be drawn between cause and conditions in 
succession as their relations are often reversible. A cause at one time 
may be a condition at another, and vice versa. 

3. The formation or association itself must be considered as an 
essential part of the complete environmeu't, and rfiould be so under- 
stood when reference is made to the environment, cf. Xo. 1. 

4. A given formation in its dominance tends to encroach upon all 
minor habitats and associations. These minor habitats tend to be- 
come cumulatively changed convergently toward the climax environ- 
ment or formation. This Is a process of eliminating diversity and thus 
establi^ing dominance. 

5. Where complete environmental and biotic adjustmeni -has taken 

20 ,,,X.oogTc 



148 MICHIGAN SURVBY, 1908. 

pliioe, the domiuaDce of the biotic formation i» most complete. This 
ina,v be considered a geographic or environmental optimam. This, in 
general, implies complete BucccsBion and the dominance of the climax 
formati<m. 

n. From the standpoint' of causes and processes, the succession of 
societies and formations is the expression or result of the environ- 
mental process moving toward an equilibrium. 

7. The lack of a uniform rate of succession throughout large areas 
is the rule, on accouut of the slowness with which extreme conditions 
are transformed into those of the average. 

8. The slowly changing extreme conditions tend to preserve many 
of tlie most important early stages of conditions and succession; hence 
the utility of these belated changes in validating snccession as deter- 
mined by the comparative method, 

9. Other things being equal; the slower the succession the greater 
the chances for rariation in the details and composition of the societies. 

10. The formation or climax society is only the most conspicuouB 
ctise of convei^nce. reached by all routes and successions, at a given 
environmental center. 

11. The succession of societies within a formation is liable to be 
more stable in its main features than the composition of its societies. 
Probably the general features of such a soccessioa most nearly ap- 
proximate that which the region passed through in the development 
of the formation. Adams, '05, p. 67. 

V2. Pommtions of different geographic centers will vary in their 
doiinnant dynamic tendencies, yet open (nnforested) formations will 
have certain features in common, as will also forest formations. Thus, 
not only will the compositions of the societies vary, but also the climax 
formations and their dynamic trends. 

13. The stability of the climax environmental factors and their biotic 
formations is only relative. They may themselves migrate or change 
by a pr(^;reB6ive succession in the direction of the dominant environ- 
mental trend. This migration involves a true succession, as is well 
illustrated by changes and successions attending the Glacial influencea 
and the elevation of the Coastal Plain of the United States. (Cf. Adams, 
'05). 

14. The stability of dominance is due to a complete biotic and 
enWroumentul adjustment brought about by the repeated selections of 
the preceding succession and resulting in a "pure culture." Dominance 
may be likened to the static social condition of China or to a monopoly. 

15. Succession is a form of complete or entire environmental selec- 
tion, certain species or associations receiving an envirwi mental ap- 
proval while others are excluded. This is a particular and extensive 
form of natural selection. Successional selection in its broadest ecolo- 
gical aspect includes the evolution of the organisms, particularly as 
members of associations in their most intimate environment. 

Hi, Any association not a climax is in unstable equilibrium and in a 
condition unfavorable to its permanence. The climax society is in a 
state of biotic and environmental equilibrium. (Cf. Warming, No. 6 
and Clements V; also cf. No. 13, 14). 



3y Google 



ECOLOGY OF ISLE ROYALE. 149 

17. Widespread physically uniform conditionH favor n domiDaot 
biotic formation. Climate may neutralize topographic diversity, or 
topography the climate. Baseleveliog and other geological processes 
which favor the production of uniform conditions will favor dominance. 
(AdamB, Amer. Nat., ,15, p. 842). 

IS. From an evolutionary standpoint the ealier stages of succession 
ore liable to be struggles with the physical environment; later, in the 
intermediate state of "storm and stress," the competition is most di- 
verse and intense, and may thus be a fertile source of adaptive changes 
and individual adjustments, through se\'ere selection; and finally in 
the stage of dominance, the competition is also biotic and physical, but 
under relatively simpler conditions. Permanence of new characters may 
be favored by habitat isolation and thus fuvor polytypic or divergent 
evolution. , 

19. In RuccesHion the adjustments and modifications of species may 
be accomplished by a change from one society to another as well as 
by indiTidual modifications or adjustment within the society. 

20 Pioneer invaders, except in social species, are generally isolated 
«ud increase progressively with dominance. Cf. Warming, No. 1, 

21. Species and individuals in the early stages of • succession or of 
societies are relatively few, increase in the intermediate stage, and 
are again reduced in number with dominance and in the climax society. 
Cf. Warming, No. 2. Clements VI, (3, B). 

22. The species of open (unforested) formations are only pioneer 
societies in forested formations and vice versa. Cf. Warming, No. 5, 

23. The less sedentarj' species, those less inclined to regularly re- 
turn to old nesting sites, and young birds tend to become pioneers and 
thus extend the breeding range. Cf. Warming No. 4. Dixon, '97, p. 
Kl. 

24. Pioneers generally come from near bv and from similar condi- 
tions. Cf. Clements III, (3). 

25. Extension of range takes place mainly at the unoccupied mai^n. 
This may mean unilateral or radiate extension. Cf. Clem^its V, (5). 

26. The succession from the aquatic aBsociation to the forest is 
probably an ancient one. In this there is a general succession from the 
leas to ihe more specialized kinds of birds. Cf, Warming No. 2; Clements 
VI (4). 

VII, SoMH Adyantaoes op a Knowlbdob op th^ Laws op Succession. 

The study of snccession implies a detailed knowledge of the field 
relations of birds, and as this hasr received so little attention as a sub- 
ject of special study, it is perhaps worth while to mention briefly some 
of the practical and scientific advantages which we may reasonably 
expect will result from the development of this phase of ini'estigation. 

The onrrent discussions of environments are generally very frag- 
mentary and chaotic, and the careful study of bird habitats and suc- 
cession will greatly improve this phase of ecology. Here is a field of 
study in need of distinct recognition as a subject worthy of detailed 
investigation, in addition to those lines already current. When once 
this field is developed, then and only then will it be possible to in- 



ISO MICHIGAN SURVEY, 1908. 

telligently diaruBB tbe evolution of avian enviroiiiuents and to cor- 
relate them with the evolution of birds themselvcB. It is qnite prob- 
able that one of the main couditiong whieh prevents a more rapid ad- 
vance along evolutionary lines is in a large measure diie to the almost 
utter failure to anahze dvnami <-an v environment al i-mnpl eieg . Suv- 
ceseion, studied in its broader aspects, should greatly aid in the for- 
mulation of the laws governing the "struggle for existence," which is 
frequently condemned for its indefinite charactei'. 

From another ]>oint of view there aPe very important reasons for 
urging extensive studies of this character at a relatively early date, 
because tbe encro achmenta of civij ization, which by the destruction of 
the foreetB, the di'ajuage oi tiie Tand, irrigation, fanning and grazing 
of the grasslands, are rap idly deBtroying original envi ronmental c on- 
ditions before they ai-e studied ecologloallyT^ivitiCll ffT WUrope nas al- 
ready gone through this stage of demolition, and it is only to new and 
relatiraly unmodified countries that we can loiA for an adequate state- 
ment of these pi-oblems and their relations in their original and pri- 
marily evolutionary and developmental form. It is not improbable 
that the next generation may wonder why Bome subjects, the investiga- 
tion of which might have been delayed, have received detailed attention, 
while others equally or perhaps even more important have been almo^^t 
ignored and must forever remain unknown because of this n^lect to 
secure the "vanishing data." (C'f. Haddon, '03.) 

Such ecological studies may be expected to have a valuable reflex in- 
fluence upon the naturalist himself. ^Ve may hope that the future re- 
visor of a group of birds will consider a knowledge of the field relations 
of his specimens as an essential qualification, just as at the present 
time a large aeries of specimens is held necessary. Fifty years ago a 
limited series waa considered no disqualification, just as to-day the 
lack of a knowledge of their ecological relations is not so considered. 
Perhaps our ideas of relative values must change. In this connection a 
statement from Tristram ('94, p. 472) is to the [Ktiut: — "The closet 
aystematist is very apt to overlook or take no count of habits, voice, 
modification and other features of life which have an important bear- 
ing' on the modification of Bpecies. To take one instance, the short- 
toed lark {Caliindrella trachydactyla) is spread over the countries bor- 
dering on the Mediterranean; but along with it, in Andalusia alone is 
found another species. Cat. baetidOj of a rather darker color, and with 
the secondaries generally somewhat shorter. Without further knowl- 
edge than that obtained from a comparison of skins, it might be put 
down as an accidental variety. But- the field naturalist soon recc^-; 
nizes it as a most distinct species. It has a different voice, a diOei*-: 
ently shaped nest; and, while the common species breeds in the plains,; 
this one always resorts to the hills. The Spanish shepherds on tfaeJ 
spot recognize their dietinctness, and have a name for each species." ' 

Many cxtunples of similar character might be cited to show the 
scientific value of a knowledge of the environmental relations of birds, 
and a nmraent's reflection will show that the problem of succession is 
only a small part of the genei-al problem of environmental relations 
of plants and animals. Attention has already been directed to the 



relation whi<h this general subject bears to evolutionary )>roblem8. [ 



ECOLOOT OF ISLE ROYALS. 151 

It is not at all aalikely that ancceBBJo n i» very closely relaW ^"^ ■f""" 
of the ca oflcg of bird migratioiij mnl lllftt Tt'ith advapce in this subject 
much ligjEir would be tnrown upon migratiou. Migration is doubt- 
less another ilhistr ation of convergent phenomen a. In all probability, 
migration has originated not only independently in very diverse kinds 
of birds, but perhaps repeatedly, from different causes, even in the 
same group. The causes of migration must l>e numerous, varying with 
different eeolc^eal groups, which appear to be the true natural units 
for study and comparison. Thus the comparative etudy of migrations 
of different kinds of associations, as formations and societies, should 
lead not only to a better understanding of the various aesociationa, but 
should also contribute to tlie genial subject of migration which seems 
to have shown a tendency toward stability in the current methods of 
study. It scarcely seems probable that with the diverse formations in- 
habited by birds, and with their ecological diversities there should be 
only a few causes of the phenomena. 

To keep pa ^e with auccesaions animals must either adjust themselves , 
change their habitat, or migrate. Fi-om suck reJations ii is eviaent that 
varfoUH SUpinwWI "ellV IruuiHPL lUi responses must be tested primarily 
irithin the amioriation avd cnvirotinient to icftir/i the animal normalhf 
helongs. To this class belongs protective coloration and allied phenom- 
ena. To be of fundameotaJ value, the influence must have some perman- 
ence and this may be sought in the dynamic trend and dominant influ- 
ences of different associations. It is difficult to conceive of other more 
reliable methods of ai»i»roach to such problems. 

In addition to the scientific value of this line of investigation, there 
are important economic npplic.Ttinna ^ ttu lawn ofn y-jini e nvimnmpnt 
This is parti culafiyTPTtH-onorestry and agriculture. The forestry prolj- 
lem is continually becoming more important, but the relation of bird 
life to forests and forest succession has. received little attention. As 
agents for scattering seeds of trees and shrubs, birds ni-e very important. 
Here is where the interests of the avian ecol(^st and forest ecologist 
overlap. The student of bird life will wish to know how a region is to be 
reforested, and what smceBsion of bird life will attend the auccession 
of the forest .as reforeslnli on Piogi'esses. On the other hand, the 
forester will wish to know how birds will aid or retard him in the 
process of reforestation. Then, in guarding or protecting the forest, 
what help can be aecuitd from birds with regard to insect pests? These 
are only samples to show that here is a field which, as time advances, 
will become of more and more importance, and that these problems 
will eventually call for specially trained men to handle them. 

In connection with forestry and agriculture we have quite exceptional 
conditions for extended experimental studies in hird succeswion as re- 
lated to forest succession, crop rotation, etc. The relation of birds 
to agriculture appeals to a much larger number of people than does 
their relation to forestry. There are several reasons for this; first, 
because more jwraons are interested iii farm and horticultural crojwi 
than in forests; and second, because birds an* soon attracted in such 
large numbers hy the food supply of grains and fruits which these crojis 
80 greatly increase, that the extensive destnii-tion hy birds readily at- 
tracts attention. And while we hear much of the great reduction of 



1B2 MICHIGAN SURVBT, 190S. 

certain species of birds in parts of the coantry, it is not at all im- 
probable that with the destruction of the forests (which wm* dense and 
dominant and tended to limit the abundance of many species frequent- 
ing the open), and the increase of food in cultivated fields, there has j 
been an increase in the total nnmber of birds, even in spite of the I 
great numbers killed bj man. 

But to the i^ase of succession with which we are primarily concerned, 
almost no attention has be«i given, in spite of its fundamental rela- 
tion to crop rotation and the corresponding avian succession attend- 
ing this. Indeed there seems to be a very decided need of a thorough 
investigation and discussion of the general principles underlying alt 
these economic problems, that they may be brought into harmony with 
the advances made in some other phases of ecology. 



» Rbpebexcbs. 
Adamt, r iian c. 

1905. The Postglacial Dispersal of the North American Btota. BIoI. Bull., 9, 

pp. 53-71. 
Beyw, G. E., AUiwn, A., uid Kopman, H. H. 

1906. List of tlie Birds of Louisiana. Part 1, The Auk. 23. pp. 1-15. 
Bray, W. L. 

1306. Distribution and Adaptation of the Vegetation of Texas. Bull. Univ. of 
Texas. No. 82. 
Btewiter, W. 

1906. Tbe Birds of the Cambridge Region of Massachusetts. Mem. NuttalT 

Omilh. Club, No. IV. 
Brown, R. A. 

1907. A Study of the Birda of the Overflow, Bast of Ann Arbor, Michigan. 

Bigbtb Ann. Report Mich. Acad. Scl.. pp. 162-174. 
Bonis, F. Z. 

1901. A Sectional Bird Census. The Wilson Bull., N. 8.. 8. pp. 84-103. 
Clements, F. E. 

1905. Research Methods to Ecology. Lincoln, Nebraska. 
Coville, F. V. 

1893. Botany of the Death Valley Expedition. Cont. U. S. Nat. Herbarium, IV. 
Coville, F. v., and MacDougal, D. T. 
1903. Desert Botanical Laboratory of the Carnegie Institution. Carnegie In- 
stitution. 
Cowles, H. C. 

1901. Plant Societies of Chicago and Vicinity. Bull. Oeog. Soc. Chicago, No. 1. 
1901. The Phyaiograpblc Ecology of Chicago and Vicinity; a Study of the 
Origin, Development and Classiflcatlon of Plant Societies. Bot. Gaz., 
31, pp. 73-108, 145-182. 
Dixon, CbM. 

1897. The Migration of Birds. London. 
Fcf nald, M. L. 

1907. Soil Preferences of Alpine Plants. Rbodora, 9, pp. 149-193. 
Forbes, S. A. 

1907. An Ornithological Cross-Section of Illinois in Autumn. Bull. III. St. 
Lab. Nat. Hist., 7, pp. 305-336. C^r» -nl > 



ECOLOGY OF ISLE ROYALB. 153 

Frottingham, E. H. 

1907. Notes on the Ulctlgan Forest Reserve. Bfshth Rep. Mich. Acad. Sci., 
pp. 167161. 
Guiong, W. F, 

1903. The Vegetation of the Bay of Fundy Salt and Diked Marshes; an Ecologi- 
cal Study. Bot. Qaz., 36, pp. 161-1S6, 280-302, 349-367, 42$-4e&. 
190E. The NaMent Forest of the MIscou Beach Pl^n. Bot. Oaz.. 42, pp. 81-106. 
Gilbert, G. K. 

1885. The Topogntphtc Features of Lake Shores. Fifth Ann, Rep. U. 8. Oeol. 
Surr., pp. 69-123. 
Gr«T, A., and Hooker, J. D. 

1881. The Vegetation of the Rocky Mountain Region. Bull. U. 8. Geol. and 
Oeogr. SUFT. of Terr. (Hayden), 6, pp. 1-77. 
GulliTer, F. P. 

1899. Shoreline Topogr^hy. Proc. Am. Acad. Arts and Scl., 34, pp. 149-258. 
Haddon, A. C. 

1903. The Saving of Vanishing Data. Pop. Scl. Ho., 62, pp. 222-229. 
Harper, ILM. 

1906. A Phytogeographlcal Sketch of the Altamaha Grit Region of Uie Coaatal 

Plain of Georgia. Ann. N. Y. Acad. Scl., 17, pp. 1-3G9. 
Harvey, L. H. 

1903. A Study of the Physiographic Ecology of Mount Ktaadn, Maine. Univ. 

of Maine Studies, No. &. 
Jacob*, J. W. 

1904. The Haunts of the Golden-Winged Warbler. Gleanings, No. 3. Waynes- 

burg, Pa. 
Johnson, K. S. 

1907. Mean Monthly and Annual Relative Huniiillty Charts of the United States. 

Rep. South African Assoc. Adv. Scl., 1906, pp. 161-168. 
Jadd,8. D. 

1902. Birds of a Maryland Farm. Bull. No. 17, Dlv. of Biol. 8ur. U. 8. Dept. 

AgT. 

LeConte, J. L. 

18G0. General Remarks upon the Coleoptera of Lake Superior. Agasslz, "Lake 
Superior," pp. 201-242. 
HcCreary, 0. 

1006. The Ecological Digtributioo of the Birda in the Porcupine Mountains, 
Michigan. Mich. Geol. Surv. Ann. Rep., 1906, G6-67. 
HerrianitC. H. 

1890. Results of a Biological Survey of the San Francisco Mountain Region 
and Deaert of the Little Colorado, Arizona. N. A. Fauna, No. 3, Dlv. 
Ornlth. and Mammalogy, V. S. Dept. Agriculture. 
1S99. Resalta of a Biological Survey of Hotmt Shasta, California. N. A. Fauna, 
No. 16, Dlv. Biol. Surv., U. S. Dept. Agriculture. 
Oibom,H. F. 

1902. The Law of Adaptive Radiation. Amer. Nat., 36, pp. 353-363. 
Palmer, W. 

1900. Ecology of the Maryland Yellow-throat, and Its Relatives. The Auk, 17. 

pp. 216-242. 

PenhaUow, D. P. 

1907. A Blazing Beacb.^ Pop. Scl. Mo.. 70, pp. 657-664. 
np«r, C. V. I 

1906. Flora of the SUte of Washington. Cont. U. S. Nat. Herbarium, XT^>^IC 



154 MICHIGAN 8URVBT. I»08. 

Pound, R., and Clemento, F. E. 

1900. The Phytogeography ot Nebraska. Bot. Surr. ol Nebraska. General 
Survey, 1. 
Kidgwari R> 

1874. The Lower Wabash Valley, Considered In Its Relation to the ntunal DIb- 
trfcta of the Bastem Region of North America: with a Synopals ot Its 
Avian Fauna. Pro. Boat. Soc. Nat. Hist., 16, pp. 304-332. 
18S9. The Omltbology ot llllnola. III. Nat. Hlet. Suft., 1, Springfield, 111. 
Ruthven, A. G. 

190S. An Kcologlcal Survey of the Porcupine Mountains and Isle Royale, Michi- 
gan. Geol. Surv. Mich., Ann. Rep-, 1905, pp. 17-5E. 
Stejneger, L. 

1903. A Reply to Recent Strictures on American Biologists. Science, N. S., 
19, pp. 371-376. 
Townsend, C. V. 

190G. The Birds of Essex County, Massachusetts. Mem. Nuttall Ornithol. Club. 
No. 3. 
Transeau, B. H. 

1903. On the Geographic Distribution and Ecological Relations of the Bog Plant 

Societies of Northern North America. Bot. Gaz., 3S, pp. 101-420. 
1905. Forest Centers of Eastern America. Amer. Naturalist, 39, pp. 875-889. 
190&-'O6. B<«B and Bog Flora of the Huron River Valley. Bot. Gaz., 40. pp. 
3&1-3T&. 418-448; 41, pp. 17-42. 
Tristiam, H. B. 

18B4. Field Study in Ornithology. Smith. Rep. for 1893, pp. 465-486. 
Watlcina, L. W. 

1900. Michigan Birds that Nest in Open Meadows. First Report of Mich. Acad. 

of Scl., pp. 66-75. 
"Whitford, H. IT. 

1901. The Genetic Development of the Forests of Northern Michigan; A Study 

in Physiographic Ecology. Bot. Gaz., 31, pp. 289-325. 
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Young, R. T. 

1907. The Forest Formations ot Boulder County. Colorado. Bot. Gaz., 44, pp. 
321-352. 



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ECOLOGY OF ISLE ROYALE. 



THE COLEOPTERA OF IBLE BOYALE, LAKE SUPERIOR, AND 

THEIR RELATION TO THE NORTH AMERICAN 

CENTERS OP DISPERSAL. 

BY DR. CHA8. C. ADAMS. 

1. Introductory Xotc. 

The beetles secured ia 1905 by the Museum expedition were collected 
during July and Augnst by rarions members of the party. We are 
indebted to Prof. H. F. Wiekhani, of the Univwsity of Iowa, Iowa City, 
for the determiDatioD of moat of the species ; the Temainder were deter- 
mined by Mr. E. A. Schware, of the U. S. National Museum, through 
Dr. L. O. Howard and Mr. E, S. Titos. To these gentlemen we wish 
to express our appreciation for these favors. Mr. A. B. Wolcott has 
kindly furnished certain records of distribution, as indicated in the text. 

The field notes and collections were largely made by Dr. H. A. 
Oleason; some were made by the writer; and specimens were also col- 
lected by B. F. Savey. The geographic range of all the species taken 
is giren in some detail, as a basis for geographic conclnaions. Time 
limitations have prevented a full discussion of these. 

Tlie geographic relations of the fauna have been discussed in more 
than customary fullness. The entire subject of the geographic rela- 
tions of the North American beetle fauna had to be gone over; and 
as the work advanced, it became evident that a general account of these 
fauna! relations was desirable from a standpoint sontewhat different 
from that generally expressed. Undoubtedly many important papers 
and statements have been overlooked, so that it will be desirable for 
others to further extend this treatment of the subject. 

Attention should also be called to the fact that in the past studies of 
the distribution of insects hare been largely irresiiectiTe of their 
habitats, associations and such ecological relations. This has been 
one of the many defects of distributional studies, as it is of the present 
study. It is for this reason that an attempt is made to briefly discuss 
the habitat relations and successions of beetle associations. Life 
history, food habits and other ecological phases need detailed investiga- 
tion and discussion so that all these phases can be related to the causes 
and conditions which affect distribution. The economic advantage of 
a knowledge of the laws of succession of beetle associations has largely 
been neglected, but ultimately must become one of the main general 
principles in much economic practice. This will probably become more 
conspicuous when forests assume a higher value, and the relation of 
beetles to reforestation, etc., demands careful attention. 



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168 MICMiaAN SURVEY, 1908. 

77, Notes on the Habitat Relations of Beetles. 

The beetles collected by onr party were tabulated by Stations to 
determine if mailed babitat preferences were evident. The tabolatioD 
BbowB that at most stations only a relatively few species were found. 
At those atatioDB where the largest variety was found, as at our camps, 
the condittODB were exceptional. The occurrmce of the dowers of Uie 
Cow Parsnip attracted many species. It is quite evident that tliese 
flower frequenting species did not breed there, so that in a strict sense 
they do not belong to these open areas, as the breeding places of in- 
sects must furnish the only substantial basis for the determination 
of insect habitats. In many cases only provisional habitats can be 
assigned with our presait incomplete knowledge of life histories. In 
many species the larval and adult habits are very different, particular- 
ly with regard to their food. For this reason error is very liable 
to occur and caution is necessary. In the present provisional dis- 
cussion the haunts frequented by the adults have been primarily utilized. 
This is an nnfortunate limitation, but it is hoped that this will not 
confuse the main feature of the problem. 

On the JAke Snperior beach (I, 1) the following 14 species were 
found: Caloaoma frigidum, Bembidium {five species), Rhantas hinota- 
tut, Anatis 15-punctata, Macropogon rufipea, Corynibites medianas, 
Podabrus diadetna, MaUhodes niger, Sericea vespertina and Lep- 
utura chrjfsoooma. It is evident that some had been washed up 
by the waves {Calosoma and Anatis) while others normaJly frequent 
sandy beaches {Bembidium) or the beach pools (Bhantus), while still 
others were here because of the open character of the beach and the 
proximity of the forest, Hayward ('97, p. 37) says concCTning the 
habitat of BembidiuTn: "Most of them are riparial in their habits, 
occurring under stones and refuse near the water's edge along streams, 
the shores of ponds, or on the seamoss, while a few occur alm(»t any- 
where." 

The clearing about the Light-house (I, 7) had the greatest variety 
of beetles found at any station. It included 37 out of the 89 species 
found at< all stations. There were several circnmstances which com- 
bined to make this number large. The greater opportunity of those 
about camp to make collections; the season of the year (July); the 
presence of the Cow Parsnip in large numbers (which acted as insect 
traps, and upon which beetles congregated in sach numbers that they 
were easily brushed into the cyanide bottles in large numbers) ; and 
the jffesence of logs, stumps, brush, etc. The flowers were a very con- 
spicuous factor, and on these Leptura chrysocoma gathered in large 
numbers. The open space was favorable to the Carabids, the flowers, 
for the Coccinellids, Elaterids, Buprestids, Trichias and the Cerambycids. 
The two latter frequent also the logs and stumps. The wandering, 
trauip-like existence of these adult wood-boring beetles should not con- 
fuse one as to the true habitat of the immature stages which is in the 
forest. The surrounding forest was mainly composed of White or Paper 
Birch, Balsam and Spruce. 

At the camp on Siskowit Bay (Y, 3) somewhat similar conditions 
were found to those at the Light-house (I, 7), but there were fewer 
flowers, more cut timber, furnishing logs, stumps, brucdi; a log shack 



BCOLOQY OF ISLB ROYALE. ISO 

WEB very thoroughly infeeted with beetles and their Hymenopterons 
parasites. The BtnToiinding forest was largely White SiHiice and Paper 
Birch. Xylotrechus was particularly abundant at this place. 

Other open places, as those bordering the beach (V, 2) or the small 
openings on the ridges {I, 2], produced, in addition to the wandering 
flower feeders Bupresth atriata and Mordelleatiita acapularis, a few 
Carabids, as Rarpalus megacephalua and Pterostichua femoralia. 

In the Balsam, White Spruce and White Birch forests (I, 3 and V, 
4) the Staphylinids, Orophoema, Boletobiua cincticolUa, the Erotylids, 
Tritoma and the Carabids, Calathita and Blechrua, are characteristic. 
The moist conditions which favor fleshy fungi show a marked influence. 
Here in the forest, of course, must also belong a great number of wood 
and bark boring beetles, which our limited collecting found assembled 
in the sunny openings on flowers. 

Along the Desor trail (III. '04), through the hardwood forest of 
Yellow Birch, aspen and Sugar Maple, two other Staphylinids were 
found, Quediua fulgidus and Tachinua memnoiua and the Scarabaeid 
Oeotrupea hlackburtUi. 

If now we turn to the lakes and bogs, a very different kind of beetle 
life is found. On the surface of Siskowit Lake (V, 6) were found 
Oyrimta minutua and picipea and in the water-lily margins of ponds 
and lakes were found (III, 5 and IV, 3) Donacia proximo, cincticomia 
and Oalentcella nymphaeac. In the tamarack and arbor vitae swamps 
(I, 4, 6 and V, 5) the following species of water beetles were found: 
Baliplua ruficoUia, Bf/droporus triatia, Agabua congener and Scutop- 
terua Jtomii. 

These may seem very elementary and commonplace observations, but 
the principles which underlie the correlation of certain (even comioon) 
species and their environmental conditions are very generally ignored 
by students of local faunas, except for collecting purposes. To know 
the exact habitat of certain species in one locality does not by any 
means prove that tbe subject is exbausted for other localities. ' 
No general ecological treatment of our beetle fauna has been attempted, 
not even of the smaller groups, such as families or genera, or even 
for a local area. The nearest approach we have to such woi^c is found 
in certain economic papers, devoted to insects affecting some particular 
plant. Here is an excellent field for investigation. 

Before leaving the subject of habitats, attention should be called 
to certain publications which are of particular use in the study of tbe 
life histories of insects in these northern forests. The first is Packard's 
"Forest Insects," and the second is Felt's "Insects Affecting Park 
and Woodland Trees," 

A few suggestions are added as to methods of ecological collecting 
which may aid similar 8ur\'eys. When the time for a survey is limited 
some system of ecolt^ic trapping will prove of great advantage. Thus 
for aquatic beetles traps, like those planned by \eedham, may be very 
useful; and still others are needed for the ground fauna and those 
frequenting trew and shrubs. Sweeping and beating as usually prac- 
ticed, while securing many species, certainly produce little ecological 
data. It may be suggested that systems of trapping may be devised 
which will contribute much valuable ecological infonnation. 



160 MICHIGAN SURVEY, 1908. 

///. The 8ifCce8tion of Beetle Asaociattona. 

'nie subject of BUcceasioQ ia a relatively new- one in entomology. The 
only other paper treating of beetle succession known to the writer is 
that by Shelford ('07). Our points of view are very eimilar but have 
been independently conceived. Broadly speaking succession means the 
change, in time, of the insect life at any given habitat or place. 
Our aim is to note the changes in the composition of the beetles 
found associated in a given breedinf/ habitat or region. The method 
is first to determine whait species of beetles are associated or found 
together in the same habitat, and then to determine their mutual and 
environmental relations, so that their laws of change may be determined. 
Habitats and their associated insects have ver>' rarely been considered 
«a worthy subjects for special study. Even ia very excellent local 
lists, but little attention is given to this subject. This is well exemplified 
by Wickham's Bayfield list. In one case he says : "A peat-bog of several 
acres ia extent also proved very productive of peculiar species." But 
unfortunately he does not indicate the kinds. 

In spite of the lack of a detailed study of the problem of beetle suc- 
cession, however, certain general relations are apparsit. We will onlj 
attempt an outline of the problem as found on Isle Royale. This in- 
volves an idea of the history of the island as the Lake formerly stood 
at a much higher level, which as it fell exposed I^le Royale. We are 
thus given, as a natural starting point, the Lake shore and beaches. 

1. The Lake Shore and its Beetle Associations. 

Topographically the shore is very diverse in its character, and all 
stages are to be found, from a cliff to a low rocky shore and on to the 
gravelly and aondy beach. When the shoi:e lies at a low angle, so that 
beach pools are developed by the waves and rain, certain water beetles 
as Rhantus binotatus at Tonkin Bay ( I, 1) and Scovill Point 
(IV, 1) find a habitat. Upon topographically older beaches, where 
gravel and sand have accumulated, various species of Bembidium are 
to be found, as previously listed. 8uch a sandy beach often contains 
a mixed lot of beetles, and may contain examples of a large number of 
species from all habitats, which have been tossed up by the waves. 
It is probable that many of these come from a considerable distance. 
Upon the upper parts of such a beach, where soil accumulates and 
annuals grow, certain flower beetles, as Leptura and Trichias are liable 
to be found feeding. In such a soil may be expected Carabids, as 
PterostichAis fcmoralis was found upon the heath beach (V, 2) on the 
south shore of the island. 

The transition from the upper beach to the rock openings is often a 
gradual one; all stages of which were found preserved. 

2. Rock Opcningn and Astuciatcd Bcctlcn. 

These park like rock openings and open oak ridges furnish a transition 
from the beach into the forest conditions. They are characterized by the 
absence of soil or the presence of only a shallow one. and by the rein- 
deer moss and heath plant society, liie shallow soil and low open v^- 
vegetation favors the continuation of some of the Carabids found upon 



ECOLOGY OP ISLE ROYALE. 161 

the apper beach, as Pteroatichus, Under such conditions were found 
P. femoraKe and Harpalas megacepkalus (1. 2; V, 2). The flower 
feeders also continue to maintain their position, bnt the Bembidiumi 
have lai^ly been eliminated. 

These open sunny spots, sorronnded by dense shady forests, in their 
attractivenees for insects, remind one of electric li(^ts where insects 
congregate in sndi vast numbers. As representative of these condi- 
tions the great variety of beetles found about the camps should be 
recalled. 

3. Lake, Pond and Bog Habitats and Aasociationa, 

From the park-like rock openings let us turn to another series of 
open habitaits: those which are initiated by inland water bodies. The 
vater beetles to be found in the open lakes were not given special at- 
tention but there can be no doubt as to their existence in such places, but 
in general we may say that water beetles increase in number andi hind 
nearer the margins and in shallow water. Here the Oyrfnidt (V, 63)» 
Donaoia (III, 5), and Oalerucella (VI, 3) abound, while farther in- 
land at the bog mai^ns and in the bogs occur Haltplids and Hydro- 
philids, Saiiplut, Eydroporus, Agahvs and Soutopteraa (I, 4, 6; T, 6). 
Even this brief series outlines the main features seen in the transition 
from the strictly open water conditions and species to those of the bog. 
But this succession may even be safely carried a st^ farther, as i» 
clearly shown by an important obBcrvation by Wickham ('97, p. 126) at 
Bayfield, Wisconsin. He states that "l%e Water Beetles were not: 
found in such abundance as I had hoped from a perusal of lists fronk 
northern localities, and of those named in the present report a great 
part were taken not in water but under moss in damp spots — a pecu- 
liarity which I have noted in some species of Agaibua collected on a 
previous trip to Alaska." Under such conditions as this the bog aaso- 
oiatlon of w»ter beetles may even be able to spread beyond the bogs 
and invade the forest, a change of habitat which has been recognized 
among plants but has been largely overlooked by students of animals. 
It is only by the detailed study of habitats that the significance of such 
facts can be understood and the peculiarities of succession determined 
for different regions. 

There is thus seen to be a very perfect transition from the bog 
forest into that of the balsam and spmce, and the dominance of coni- 
fer trees clearly shows that the beetles frequenting such forests will be 
on the increase as the aquatic association of beetles declines in domin- 
ance. The methods and detailed order of this succession awaits in- 
vestigation, but it is clearly dominated by the forest succession. 

Jf. The Forests and Their Beetle Associations. 

The gradual and perfect character of the transition from the bog 
to the balsam-spruce forest has just been indicated. This is paralleled 
by a corresponding transition from the park-like openings and the bare 
ridges to the forest. These liabitats change as rapidly as soil, humus, 
and shade increase at the mar^ns of the openings; and as shrubs and 
young trees encroach upon the open. Thus as the hogs fill up, and as the 
soil increases on the rock surfaces, both habitats tend to become trans- 



162 MICHIGAN SURVEY, 1908. 

formed toward and invaded bv the balsam-Bpnice forest conditioDB and 
asBOciation. Here is a clearly defined convergeiLt tendency, the exposed 
ridges and the water basine both tending to become forested. Corre- 
sponding with these environmental changes are corresponding ones in 
the beetles. The Carabids, as previously mentioned, continne in the 
homns from the rock openings as the wuter beetles may in the damp 
moss. As the vertical extent of the forest increases and the forest 
crown migrates upward, the intervening trunk, bark and branch habitat 
for beetles enlarges and the leaf eaiting inhabitants of the forest crown 
rise upward. This crown fauna retains, or rather continues SMne of 
the characteristics found at the mai^nal zone, with which it retains 
direct continuity. The mai^nal zone of trees is likely to be birchen (yel- 
low or white) or aspens, in the rock opening succession, and conifers 
in the b(^ series, a feature which Influences the beetle fauna. With a 
dominance in the forest of Balsams, Paper Birch and White Spruce, the 
beetles (and many other insects as well) are likely to be much in- 
floenced, not only by the plauit food, but also by the physical con- 
ditions associated with the forest. As one plant or forest societ? 
replaces another, the unfavorable oonditiona of the declining society 
may be expected to favor insect injury as it is well known that in 
general vigorous trees suffer less from sncb attacks than those which 
are defective. Uoder such circumstances as this Insect injury may 
be a useful index of succesBion, as well as a factor hastening it. Under 
such circumstances the climax of insect abundance or dominance 
may la^ behind the climax of the development of the plant society 
upcm which it depends. Insects may also initiate a plant succession, 
through a period of extreme abundance by doing damage to the food 
plant, thus permitting the invasion of other forms. If, however, the 
hardwoods, the yellow birch-sugar maple forest, is the climax type, then 
the balsam-spmce-paper bircb association will be succeeded by it in 
time; and a farther change in the beetles may be expected. But here 
also, as in the coniferous forest, a dense forest stand appears to be 
unfavorable for the abundance and variety of beetles (as is generally 
the case with many other aiuimals). This scarcity of beetle life in 
the dense forests of the I^ake Superior region has been commented 
upon by LeConte ('50, p. 201) as follows: "The whole country being 
still almost in a primitive condition, the specimens are equally dis- 
tributed throughout a lai^ space; the woods will not therefore be 
found very productive to tiie collector. In fact nearly all the species 
were adjacent to small streams ; or else were driven on shore, particu- 
larly on sand beaches, by the winds and waves after being drowned in 
the lafca" 

Throu^ont the preceding discussion the intimate relation of the 
beetles and the v^etation has been assumed. There seems to be 
a good reason for this. TJlke ('02, p. 3) has well expressed this dependence 
as follows: "Now, as about half of all the beetles depend upon plants 
for their food, the greater variety of food plants the larger we find 
the nomber of species of beetles." In this we alsA see why so few 
species (relatively) are found where a climax plant society has become 
dominant, because such societies are, as a rule, composed of but rela- 
tively few species. At the same time it is seen why at an intermediate 



ECOLOGT OS* ISLa ROTALB. 168 

stage, with a diversified vegetation, we may expect the greateBt variety 
of beetles. From sncb relations as these it foiloiws that a knowledge 
of plant Bocceesion vill famish a very important tMlflis for the study 
of beetle micceBsion. 

While theae remains hare been primarily intended for beetles, it 
is eqnally evident that they have a mnch more extended application 
to other plant feeding InsectB and certain other invertebrates as well. 

IV. The Oeneral Characteriatics of tkc North American Beetle Fauna. 

In attempting to form some idea of the general faonal aflinities of 
the Isle Boyole beetle fauna, the literature was searched for a gen- 
eral account of the distribution of North American beetles. As no 
recent comprehensive account of the subject could be found, various 
general statements and generalizations were compiled. On account 
of their soaittered occorreace in the literature and their value and 
saggestivenesB to stndents of other groups of insects, it has been 
thought desirable to assemble and publish them. An effort has been 
made to quote only the more important statements. Several of the 
old^* statements by LeOonte, before he accepted the theory of evolu- 
tion, are omitted. A similar selection has also been exercised in the 
case of some other writings. The main aim has been to bring to- 
gether the most comprehensive generalizations which have been made 
upon our beetle fauna, so that they may have f^'eater utility, furthea- 
extension, and revision. Supplementary data from other groups of 
insects has lai^ly been omitted, although this should be given due 
weight in a comprehensive study of this subject. The references should 
he consulted in connection with the compiled abstracts. 

1. Compiled OcneralizatiotM on the Fauna. 

Carpenter, W. t. 1875, pp. 539-542 : "The principal and most inter- 
esting resnlt obtained from the study of this collection, is the demon- 
straticHi of the fact that the alpine insect-fauna of the Elocky Moun- 
tains, is nearly identical with that of Mount Washington (New 
Hampshire), Labrador, and Alaska; and that insects which are found 
upon mountains at great elevations will likely occur in a much higher 
latitude at a less elevation. 

"Insect-life, with the exception of the grasshoppers, is more abund- 
ant in the foot-hills than the plains near the foot of the monntains. 
An altitude of about seven thousand feet appeared to produce the 
greatest variety of species." p. 540. 

Cockerell. T. D. A. 1893, pp. 305, 306, 309, 310-311, 312, 313-314. 
315, 316, 317, 319-320-322. 

"The Insect fauna of the mld-alplne zone of Cueter County [Colorado] pre- 
sents some elements which are sufficiently diverse: but taken as a whole, It 
le a natural fauna, helongl&i; to a well -defined region, and henr^e available 
for comparison with other like faunae. It Is. indeed, truly characteristic of 
tbe mld-alplne, that besides Its ordinary elements. It contains spcclee coming 
up from the sub-alplne, and down from the high alpine; but altbougti it thus 
liftppene that Junonia coenia and Famaaaiui sminthevt have been taken In 
tbe same zone, It does not follow that either are truly characteristic of It, 
«r that they belong to tbe same fauna. All faunal lists contain such excep- 



164 MICHIGAN SURVEY, 1908. 

tlons or devlRtlons from the average; but vhen, as la the case of Colorado 
as a whole, there Is do imltorinlty about the range of the various Bpecfea. and 
the majority do not occur throughout the territory. It la Impossible to treat 
the region as containing a single fauna." p. 305. 

"The mld-alplne zone, as I have defined It,* extends from about fl,500 feet 
to 10,000 feet. It Is essentially the zone of oak-ecrub (Querent undulata) and 
quaking asp (Poputua tremulotaea) . Its most chaiacteristfc conifer Is Pintu 
ponderoia var. icopjilomm, but with the higb-alplne zone It shares Picea enget- 
manni, with the eub-alplne, Finua eAutia, and Junijierui virginianut." p. 306. 

"Among the Coleoptera It will be notlcad at once how many of the species 
are boreal extending to Canada {tens. lot.) and often to the New England 
States. The Soutbem element Is but alight althougb distinct II looked for; 
and there la also a fair number of species endemic In the Kocky Mountains. 
The Tenebrlonldae, characteristic of the Western prairies, are fairly numennn. 
The Coleopterons fauna, as a whole. Is strikingly distinct from that ol tta« 
HIsslaslpiri region and the Eastern States generally, except aa regards the 
boreal element. Mr. WIekham has publtsbed a list of the beetles found In the 
vldnlty of Iowa City, and on comparing It with the present list. I was astonished 
to find how few were the species common to both. This result Is brought 
about In large measure, no doubt, by ,the different character of the forests — 
those of Iowa ctmtalnlng a great variety of deciduous trees, those of Colorado 
mainly cwklfers, with very few deciduous spectee. Thus. It happens that not 
one species of Cenunbycldae Is Common to the Custer County and Iowa Cltjr 
lists, although six species are common to our district and the much more distant 
State of New Jersey." p. 309. 

"The high -alpine zone in Custer County extends from 10,000 feet on tho 
Sangre de Crlsto range to aummtts of the mountains (Olbb'a Peak, wroni^ 
called Qlbaon Peak, 13.729 feet; Horn's Peak. 13,447 feet; Humboldt Peak. 
14,041 feet, etc.). A list xit the blgb-alplne species eo f ar as observed. Is given 
In "Can. Gnt." 1890. Although the number of records Is not great, Uiey show 
that the high-alplne and mid-alplne lonee are sufficiently distinct." p. SIO. 

"Of the high-alpine Coleoptera, 2G species are recorded, and a S6th may 
be added, namely, Coccinejfo trifatoiata L., from near the Ulcawbw Mine In 
October. It extends to Canada, Lapland, etc. Of these 26, seven ganera ar« 
not mld-alplne, namely, OrtoAacliiui, Diclielonycha, ChrytobotKria, Zeugophora, 
Athout, Daaytet and Glyptina. Eleven of the species are wanting tn the mta- 
alpine collections." p. 310. 

"These statistics would undoubtedly be altered by further research, but I 
do not think they can be without significance. That the high alptne and mtd- 
alpine fauna are largely of different derivation seems to be proved by the large 
proportion of generic difference. Thus. 23 distinct species of Hymenoptera In- 
clude no less than IG genera; and eleven Coleoptera Include eight genera. 
The high-alplne, therefore. Is not, as regards Its peculiar features, derived tram 
the mid-alplne or lower; contrasting In this respect with the hl^-alplne of 
Ecuador, which 1b bo derived. 

"The affinltlee of the hl^h alpine not being with the mld-alplne, they could 
only be with the far North. Alberta being a suitable region for comparlsoa, 
I wrote to Mr. Thomas E. Bean. asXing him to tell me how many of my high- 
alplne species occurred with him. He moat kindly replied, giving me the fol- 
lowing interesting Information: 

"Of the ColeopterH. be finds at Laggan f>oIoptus loteralu, FodabTvt lateraHa, 
Orsodachna atra. Cicindela longilabris, Adoxxu viti», CKrytobothris trinervia, 
Coccineila tranMvertoguttata. Trichodei ornatus. Scmaeopi pratensit and Cor- 
delia icutellaria.- He adds: ''That Is a good sprinkling concldering that I de- 
rive the facts from a small lot I sent Mr. Fletcher several years ago, pre- 
sumably the commoner species.' * • • The timber line at Laggan Is at 
7.000 ft. p. 311, 

■'Thirty-Blx Coleoptera were found and Identified In the sub-alpine zone, and 
of these twenty-two, or nearly two-thirdB, were not found In the mld-alplne. 
These include the following eleven genera not found in the mid-alplne: Ptlyo- 
phagus. Batyle, Dityhta. Badisler, Serica, Diabralica. Tomicv». Polyphylla, 
Euryomia, Listnis and Dfamaris. Of the thirty-six species, only one, Hippo- 
damia lonvergens. was observed to range up to the hlgh-alpine. 

• Use '■ EnWiDological .Vews." 1893, p, 303, 



ECOLOGY OF ISLE ROYALE. 166 

"Thus, to both Coteoptera and Orthoptera, the difference between the two zones 
Is seen to be very marked, not only as to epecieH. but also as to genera, ebow- 
Ing that we have to deal with distinct fauna, p. 312. 

"So far as I am able to judge, the suppression of the central region Is en- 
tJrely Juatlfled, but I cannot agree as to tbe propoeea Sonoran region. An 
analysis of the Insects ot tbe Colorado Rochy Mountains shows that the blgh- 
alptne and mld-alpine elements, although sufficiently distinct, are both essential- 
ly boreal. If we follow Dr. Merrlam's arrangement, It appears that the hlgh- 
alplne Is truly boreal while tbe mld-alpine belongs to the transition reslon, 
containing a considerable number of strictly American types. The sub-alplne, 
on the other hand, is southern or Sonor&n. 

"Dr. Horn has kindly given m« his opinion as follows; 'My IdeM of the 
distribution of the Coleoptera In the mountainous region of Colorado, which 
Is a good center of the Rocky Mountain chain are as firilows: The blgta region 
seems to have been populated from the Canadian throngb the H. B. T. reslon. 
A collection made above 8,000 feet in Colorado is atmoat identical with cne 
made In tbe Lake Superior region. The same fanna runs down to N. M. and 
ArlEono, and again appears, mixed, of course, tn the Mexican Mountains. 

" 'The sub-alplne region Is one that continues from Washington to New Mexico, 
as shown by such striking forms as Brgate; MelanopttUa miranda, Iphthimut 
»«Tratut, Oaieniea externa. Caloioma Junalum in varleUea. 

"'The lower region, foothills, etc.. Is a mixture of New Mexico forms with 
those ot th« Eastern United Btatea, with some peculiar forms allied more to 
the southern regions. 

"'California Is a peculiar n«l(Mi, and. In man? resp«cts, allied to Europe (la 
Seneral), I think Calltomla sntwiles us with more species of genera pecniiar 
to Europe than does the Eastern region.' (In litt., July 14, 1S92.) 

"According to the tacts now recorded It seems that there 1», flnitly, a drcum- 
polor and strictly boreal element; secondly, a boreal but modifled or Canadian 
eJement; and thirdly, a southern element belonging to the arid portion of Dr. 
Merrlam's Sonoran region. I do not thluK any distinct &unae except these 
can be recognised, and the central region acoordSngly falls. But there la, 
sprinkled among the w^lnary types, a dittinct element of endtmic tpeclet, to 
which I shall refer later. There also seems to be a tew surviving fragments 
ot an ancient fauna, of which AntAracopteryr is a good example. 

"There seenu to be a amati California element, but tbe species falling under 
this head are perhaps rather Southern than properly Callfornian. pp. 313-314. 

"The resemblance between the Colorado fauna, and that of the Mlsslssti^i 
basin and further East, always, excepting the boreal element that comes from - 
the North, is very slight Indeed. Tbe great plains to the east of the Rocky 
Mountains have been as much a barrier as the sea would have been. p. 314. 

"A Method for De/lning Faunal Regions. It appears from a consideration 
of what has been written on faunal regions, that it would be desirable If some 
rules could be laid down, so as to leave tbe matter less to the discretion of 
the Individual writer. It would require a good deal ot research to determine 
what rules could be litid down, that would work, but as regards insects, at 
all events, I have thought It possible that the following rule might answer for 
secondary faunal divisions: 

"Any (ICO districts shall 6t regarded as in the same jecondorv fownol division, 
if the number of species common to both exceeds the number of genera in com^ 
mon. p. 315. 

'Equigeneric Areas. For minor divisions, to be used In relation to partlcu:ar 
groups, I have devised what may be termed equigeneric areas. 

"Eguigeneric areas are areas throughout which the genera of the group under 
consideration are identical, • 

"These areas are sometimes large, sometimes small. When two genera over- 
lap, the r^on where they both occur, however small, mokes a separate equ- 
generlc area- This might be thought a dlsadvantt-ge; but really. I believe it 
to be an advantage In the method, since It Is Important to reco[;iiIze these 
Intermediate or overlapping areas, p. 316. 

■'Origin of the Rocky Mountain Fauna. The numerous fossils of Colorado 

bear testimony to the fact that tbe region of the Rocky Mountains has In tbe 

past been peopled by a highly remarkable and numerous fauna. This fauna, 

however, does not appear to be ancestral to that ot the present day. Nor has 

22 



166 MICHIGAN SURVEY, 1908. 

the present fauna any specla] c<Hinectton with that of the blgli reglona to the 
tar South—the Andes. In order to arrive at Just conclUBlons, ft will be need- 
ful to consider tbese points In some further detail, 

"Alpine InsecU of the Andes. The recently- published 'Supplementary Ap- 
pendix' to Mr. Whymper'B work on his travels amoDget the Andes of Ecuador, 
containing an account of bis captures, Includes some very valuable Information 
about the Inaectfi of high altitudes In that country. The late Mr. H. W. Bates 
has written the Introduction, In which the following passages occur: 

" 'It there had been any distinct element of a North Temperate or South Tem- 
perate Coleopterous Fauna on tbe Ecuadorian Andes the collections he made, 
InexhauBtlve though they may be, would have shown some traces of It; but 
there are none. A tew genera belonging to temperate latitudes, though not 
found in the tropical lowlands, do Indeed occur, but they are forma ot almost 
world-wide dlstrlbutliHi in similar climates, and there Is no representative of 
tlie numerous characteristic and common genera of the North or South, Even 
tbe Northern genera, more or less abundantly found on the Mexican highlands, 
are absent. 

" 'One feature of tbe fauna is of great interest. It Is the occurrence of apterous 
speclea of genera which at lower levels are always winged. 

" 'It seems to me a fair deduction from the facts here aet forth that no distinct 
traces of a migration during the llletlme of existing epecles, from North to 
South, or vice versa, along the Andes, bave as yet been discovered, or are dow 
likely to be discovered,' 

"Going through the list of insects taken at high altitudes in Ecuador, the 
following points may be noted. There are four new species of Pterottichua 
from 12.000 feet upwards, but they represent a new subgenus. There Is not a 
single Amara or Harpalvt, pp. 31T-31S, 

"The Glacial Epoch. It can readily be Imagined that such a state of aSalrs 
[Prestwick's account of the Amer. Ice Age] would lead to the destruction of 
a large part ot the fauna, tbe remainder either surviving along the northwest 
ccoat-llne, or going southward to the Gulf States and Mexico. The eastern 
fauna, with which we are not now particularly concerned, would largely sur- 
Tlve, owing to there being a considerable area of unglaciated territory avail- 
able. This, Indeed, baa been the case. The Callfomlaa fauna would survive 
in port to tbe north, and also In lower California and the western coast region 
of Mexico. But the fauna of the central region would be almost annihilated, 
because the warm winds being cut off by the coast ranges, the country would 
become extremely cold, even far down into the higher lands ot M^lco. Tbe 
avid region where not actually glaciated would be a frozen deeert, and tbe 
migration of the fauna southward would be far from easy. 

"In the eastern province the species ot the moist Northern atates would And 
little difficulty Id migrating southward into the equally moist Southern States. 
The Isotherms would shift southward over moderately uniform country. In 
the centr:^ refjion, bowever, this would not be the case. There Is no place 
available to the South, except tbe molster coast line, and tbe Interior uplands, 
which latter were undoubtedly glaciated. Tbe great plains between the Rocky 
Mountains and the Mlsaiaslppl would have made an Impaaaable barrier for moat 
species, preventing migration In that direction. 

"But, It may be urged, at so-me point to the southward the mountains or 
central uplands would cease to be glaciated, and why should not migration take 
place Into the neotropical region. That it did not take place at all events 
beyond the isthmus, la evidenced by tbe facts above quoted from Mr. Whymper's 
'Appendix;' and tbe reason of this no doubt is, that the Isthmus itself was 
submerged, and all connection between North and South America cut off. This 
question of the submergence of the Isthmus ot Panama has been fuUy dis- 
cussed by various naturalists, and need not be enlar;:ed upon here. 

"It is Impossible In the present paper to give more th^ji this bare outline 
of the subject, but I believe the conclusion Is justified, that the central region 
fauna was practically stamped out during the glacial epoch; and that tbe 
present fauna Is derived from the boreal faunae which survived to tbe mst 
and to the west, and the southern fauna which survived in Mexico. This view 
seems to be supported by a consideration of tbe present distribution of species, 
as well as by geological evidence, pp. 319-320. 

"Post-Olacial Developments. Excepting the remnants ot the am^eot fanna. 



amaeot fanna, 

, Google 



ECOLOaY OF ISLB ROYAIlE. 167 

ftll the strlctlr endemic el«ment In tli« Rocky MountaioB ta of poit-gJacial 
aigtn — that ts, according to the view* bere »et fortli. ThlB means ft good deal, 
If It 1b actually the case, as I believe. Under certain clrcumatances, Bpecies 
ilerelop quickly, and we have, at least anumK insects and flowering plants, a 
gteaJ. array of new species coming Into existence. Such species are closely al- 
lied to species from which they sprang, and to each other, so as to give rise 
to much dispute as to their validity— as an example, cme may cite the genus 
Jlrffimni*, whl(± has been very productive of post-glacial species In America. 
In such a case It matters little whether we term all these diverse Forms true 
species, or eulHipeciee or races, — but to lump them under a common name ob- 
scures the facts, and leads us to Ignore one of the moat Interesting phenomena 
that are presented to a zoologist, pp. 320-321. 

"Bpeciei- Forming Area*. It Is well known that the genera commonly accepted 
oj-e unequal In value, but most of those whose validity could not be queetloned, 
sre evidently of considerable antiquity. 

"But the curious -thing Is, that these wide-ranging genera are not equally 
productive of species over their whole areas, p. 321. 

"Among insects, Arg^nrUt and Colitu, and several genera of Koctuae, exhibit 
strong species -forming tendencies in the Western States of North America. 
Catocala. In the Eastern States, has a tmt BtKmg species -forming area. And 
BO on In many other Instances niilch will occur to the reader. This phenomenon 
Is a most remarkable one, since It' affects chiefly old and almost cosmopolitan 
genera, and does not occur In the same districts In all the genera. Two cosmo- 
politan genera, as we have seen, may have their species-forming areas on op- 
posite sides of the world. It would teem. Indeed, as if there were causes at 
the bottom of It, that we do not yet understand." p. 322. 

Fall. H. C. and Cockerell. T. D. A. 1907. pp. lGO-151, 1G2-1G3; "Comparing 
the beetles ai New Mexico with those of Colorado, one Is struck by the large 
amount of difference In the lists. Colorado haa not, of course, the important 
and characteristic Middle Sonoran element, but the higher elevations are con- 
tinuous from north to south, and one would expect a practically Identical fauna. 
Botanical investigations, however, hare revealed striking differences in tha 
plants of the northern and soatbem Rocky Mountains, and a degree of ea- 
4)«mlclty among those Inhabiting the mountain ranges which Is quite nurprlslng. 
The oaks {Querent) are abundant in New Mexico, and have a luxuriant develop- 
ment as far north as MSnltou. Colorado, and even beyond. But at Boulder, and 
north of Denver, generally, they are totally absent. On the western slope they 
go farther north, and one species Just enters Wyoming; but there are none 
at all in Wyoming, with this exception, and none in Montana. This alane would 
«xplain the northward limitations to the distribution of the numerous species 
of Coleoptera which are attached to the oak, and various similar cases could 
be cited. It appears probable that the oaks were driven south during the 
Slaclal period, and owing to the nnsultablllty of their seed for being carried 
treat distances, have been unable to recover their lost ground. Under these 
circumstances, the ample powers of flight of certain of the oak feeding beetles 
are of no service for promoting migration northward of the slowly moving 
line of oaks. pp. IGO-lGl. 

"It will be noted that New Mexico shows a greater proportitm of non-Colorado 
genera than species; or. In other words, the species found In New Mexico but 
not in Colorado are more likely to be of non-Colorado genera than In the 
reverse case. This Is explained by the fact that the desert fauna In nearly ail 
groups is rich In peculiar genera, but these are represented so far up as New 
Mexico by comparatively few species. On the other hand, the boreal fauna, 
-so strongly developed in Colorado, Is largely characterized by the abundance 
«f species of circnmpolar genera. 

"In Colorado the eastern plains region has been little searched for beetles, 
.and the corresponding region of New Mexico Is also poorly known. There Is 
no doubt that the plains will furnish many species additional to the lists and 
most of these will doubtless be common to both. The Following are characteristic 
eastern species which are known to reach New Mexico, but have not yet been 
found in Colorado; Scariteg gublerraneut. Clivina bipuatulata. Clivina ferrea 
Atpidogloiaa tubangulata, Panagaeng fasciatus. Tachvs xanthopna, Fteroattchui 
.»^/i, Dyruutes tityua, Anomaia undttlata, AHndrUi teres. 

"The New Mexico list contains over 135 species, Indicating that the eastern 



168 MICHIGAN SURVEY. 1908. 

fauna is really cToealng the plains to Bome extent, and not only, reaching vb by 
way ol tbe nortbern mountains. There are etrong reasons tor' believing that 
a considerable part of this migration Is recent, and has been assisted Involun- 
tarily by man, Thia affords, of couree, a strong argument in (avor of the 
speedy exploration ol western regions, in order that ttkelr original fauna may 
be ascertained before it Is unduly oontAminated by introduced forma. Fortunate- 
ly For the naturaliBts, the desert vlll not quickly or easily accommodate alien 
elements, but It Is quite otherwise In more ordinary localities: and as Perkins 
has sbown in tbe Hawaiian Islands the result may be destrucUoc as well as 
confusion. 

"Tbe number of species common to New Mexico and Southern California, but 
not known from C<^Drado, is over ISO, indicating a wide-spread southwestern 
fauna; but In general, the spedes of tbe Southern California caaat region are 
. not tboae of the Rocky Honntalns. 

"We And over 30 names of New Mexico species listed from the Lower Rio 
Grande, but not in the Colorado, Southern California or District of Columbia lists. 
Such for example: Cincindela cirpumpfcta, Cindela »evera, DyicMriut ter- 
minaliM, PMlopAttga vtridtcollit, Selluomorpha ferruffinea, Oode§ oupraevM, 
Iichiodontus ferreua, LwUtu texantu, Agnjua otbt^ftdiu, Jiattinocenu textmva. 

"The following are examples of characteristic touthern genera which readt 
New Mexico, but do not enter Ci^rado: I'halplus, Hoioiepta, SanOalua, Thrim- 
copvff^. I^ciu, PhuioHa, AphonUlet. Btrategiu, Allorhina, Derobrachut, TylosiM, 
Dinirohiat. 

"Because of tbe eonsptcuous place which these southern genera occupy in 
tbe fanna, an entomolosist arriving from tbe north or east la very likely to 
sasume that the Middle Scmoran of New Mexico contains precisely tbe same 
elements as the Liower Sonoran of Arizona Just as It haa been assumed tbat 
Florida Is typically West Indian, becaun its numeroue We«t Indian genera 
attnhct attention, and the alMence of innumerable West Indian types is not ao 
readily ohwrvcd." pp. 151-1E3. 

Hamilton. '94 a. pp. 406-416. Cf, also Fauvel 'S9. Hamilton gives the fol- 
lowing llsta of BpeclcA Indicative as to their nativity: 

1. Speclea equally native In North America and in northern Asia not yet 
otaerved as occurring In Ehirope — 19 qteciea. 

2. Species native In North America and Northern Asia occurring In Europe — 
277 species. 

3. Species native in North America and Europe not at present known to 
occur in northern Asia — 50 speciea. 

4. Species probably Introduced Into North America now acclimated occurring 
In Europe, and those marked witb a ■ likewise In Asia. Many of these are 
cosmopolite, or becoming so, through commerce — £16 spectee. 

B. Species cosmopolite or subcosmopollte. 

Horn, a. H. 1872. pp. 383-384. "As is well known to all collectors, varlou* 
species of Eleodet occur In great numbers In alt parts of tbe west of our con- 
tinent, and the speclea themselves occur over a wide range of territory, and 
are not limited, as might be Interred from their apterous condition, to ragiona 
of small extent. As we pass from east to west over a given line, we find 
variations of average temperature, and of course great differences in eJtltude. 
These two causes, combined with, of course, the botanical cbanges. bave tended 
to produce variations from a given type to a greater or leea extent Elvodea 
obscuTa Say affords a beautiful llluetratlon of tbe extent to which this diver- 
gence may be carried. As a general rule I And. not only in Eleodea. but also 
In many other genera, that Che higher the elevation or the colder tbe climate, 
the rougher and more deeply sculptured Is the species. Tbe smoother forms of 
E. obgcura may therefore be expected !n the southern regions In which It occurs; 
for example, var. di»per»a is New Mexican, elytra with scarcely any tiTLOes 
of atrlae; var, obscura, elytra distinctly sulcata, but not deeply. !e from Colorado 
and Southern Idaho. As we advance to tbe west the elytra are more deeply 
sulcate, as in var. arala, while var. sulcipennij, from nearer the Pacific Coast. 
has deeply sulcate elytra, with very convex Interspaces. The same variation 
of sculpture occurs In Calosoma luxatum, Say. wblch starts In Colorado with 
comparatively smooth elytra, until In Vancouver we And the elytra covered with 
tines of KranuUr elevation.^, forming the variety known as C. pemelioidea. 
Walker, The two extremes of each series above noted appear to differ widely 



ECOLOGT OF ISLE ROYALE. 169 

from each other, and to be sDtltled to rank ae a distinct species. In the fore- 
golng remarks reference only has been made to variatloiiH within apectflc limits. 
The same law appears to hold between different species. In the genus Omtia 
the most roughlr Bcnlptured species occurs In Washington Territory, {O. Deieanii 
Befche) and the amoothest (O. laevii, Horn) from near Vlsolla, California. The 
object of the preceding remarks Is to explain what appears to be a law of 
variation for our western slope, and thus cauee the nnnaeess&rr multiplication of 
species, founded on slight characters, to be avoided. 

"9pecles everywhere Id our fauna appear to be distributed on ilnes of country 
presenting as nearly as possible similar meteori^oglc conditions. Thus many 
Oregon forms extend southward Into California, sradoalty seeking a higher 
mountain habitat as the region becomes warmer. Two species Illustrate this — 
Tragotoma Harrttil and Pftry^an-opftlltM collarit. Both extend tbelr habitat 
from Maine to California following the cooler regions westward from Maine 
throneh the Canada and Red River region, thence northward nearly to Sitka. Prom 
the latter point southward to Oregon both occur at ordinary level, and rising 
as a more southern region Is reached until at the latitude of Vlsalla they occur 
only a short distance below the anow-llne, at an altitude of from ten to twelve 
tbouaand feet. p. 383. 

"As might be expected each new region vlaited yields new MeUHdae of the 
genera Bpicavta and Lytta; in fact, each species of Attragalut has Its peculiar 
Lytta; and whenever any of that genus of plants Is found In flower, an ac- 
companying visitant may always be looked for." p. 384. 

LeConte, J. L. 1850. pp. 239-Z3S*, 240*: "First, the entire absence [in Lake 
, Superior region] of alt those groups which are peculiar to the American con- 
tinent. Thus, there Is no DIcaelus, no Paelmachus among the Carablca; the 
Brachelytra are represented only by forms common to both continents. Amcmg 
the Buprestldae is no Brachys; In the Scarabaeldae, the American groups (except 
Dlchelonycba) are completely unrepresented; In brief, there Is scarcely a genus 
enumerated which has not Its repreaentstlve In the Old World, p. 239. 

"Secondly, the deficiency caused by the disappearance of characteristic forma. 
Is obviated by a large Increase of the members of genera feebly represented 
In the more temperate r^ilons. and also by the introduction of many geneni here- 
tofore regarded as confined to the northern part of Europe and Asia. Among 
these latter are many species whlch^ can be distinguished from their foreign 
snaio^es only by the most careful eiamlnatlou. p. 239.* 

"When a species In one district is paralleled by another In a different region 
80 closely allied that upon a superficial glance they would be regarded as the 
aame. Theae are called aruiiogouK apeciej; e. g.. the Ollsthaert, Spondyll, Bem- 
hidla, Helophon, etc., etc., of the preceding catalogue, as c(»npared with European 
species. 

"Where several species In one region are represented by several others of 
the same genus, which perform a similar part in the economy of nature, without, 
however, displaying any further afllnlty to each other. These are called equivalent 
apeciea: e. g,, moefc of the species of Clclndela, Brachlnus, Clytus, Donacla, etc., 
of America, as compared with those nf the eastern world, p. 239.* 

'■Nctw'thFtaDdiHB this snproilniatlon to a uniform, 9u^n^ct1c Pti^ilprd. "-e atlll 
find In these tmreal regions, a prevailing character of North American fauna — 
the extreme paucity of CurcuHonldae. The Donaclae too, although numerous, 
do Dot afTord any prominent nareilellsm." i>. 240*. 

LeConte. J. L. 1851. pp. 249-250, 251, 252, 253-254. "The first fact observed 
by the collector [In California], is the very small number of species which 
can be obtained at any single locality. Day after day be meets with a continual 
repetition of a few commoa forms, with an occaaloca] admixture of rare species; 
so that at the end of two or three months a single locality will have furnished 
hfm with about ZOO species of Coleoptera, and a rather leas number of other 
orders. It will be here remembered that the contrary is true of the eastern 
port of the continent, where each locality furnlehes a large number of species, 
extending over a large area, and represented by comparatively few Individuals. 

"On removing to another locality, the same thing Is again observed, with this 
difference; tbe species of the first place, even the most abundant, are replaced 
by others, many of which are true representative species, approaching as closely 
as those of Eastern America and Ehirope; while others belong peculiarly to their 



170 UICHIOA.N SnRVET, 1&08. 

own district, and ftre without sjiy repreeent&UveB in the other parts of the 
country, pp. 249-250. 

"It must Iw observed that the locallttes east of the Sierra (Vallecitas, Colorado 
and Olla) show more reeemblance in their productions than the maritime regions 
of California: the desert nature of the country undoubtedly iH-oduces this effect, 
by preeentlns conditions unfavorable to animal life; yet even In this uniformly 
aterlle tract, great differences are observed among the smaller species whlcb 
abound only In moist places, p. 250. 

"The flrat point worthy of notice in this list la the extremely small number 
[compared with Europe] of Scarabaef, Elaterldae and Longtcornia: this might 
have been predicted, as these Insects derive their food for the most part from 
large jdants. The Curcullonldae and Chrysomelldae are not In the same pro- 
portion as in the more wooded countries. The saprophagous Coleoptera. with 
the exception of Htsterldae. are almost wanting: and these latter are not Id 
larger prop<»-tion than with us. Thus the only effect, so far as observed. Is 
the paucity of species In tribes tor which the country alForda but little food. 
The Staidiylini and Carahica bear the same proportion to the whole, that they 
do with ub; while the deficiency caused by tbe small representation of the tribes 
mentioned above, Is made up almost entirety by tbe Tenebrlonldae, wbicb, as 
is well known, are bnt slightly develoi>ed In E^astern America. The Malachldae 
tiK also in larger proportion than in other parts of the continent." p. 2G1. 

"The Tenebrlonidae, from being the group most characteristic of the country, 
might be supposed capable of giving us the most certain data with regard 
to the law of dlstrlbutJon. The great majority of tbe genera of this tribe are 
apterous: and (tf those which are not apterous, all the genera found in Cali- 
fornia are coamopolitan (Phalerla, Platydema, Helops. Uloma, Tenebrio. TJpIs, 
etc.) , except Blapatlnus. which again occurs in tropical America. Of the 
apterous genera, only three are found In eastern temperate America: two ot 
these are peculiar, and one (Noeoderma) which exists in Calltomia is also 
found in Brazil. Of this group, there are in CalifCH-nla about 23 genera, of whlcb 
6 or £ extend into the tropics. 

"The Histerldae, though not in undue priqwrtlMi, exhibit a peculiarity: they 
nearly all belong to the genus Saprlnus, which. In Eastern America and Knrope, 
forms scarcely one-fourth of the group. 

"Thus the only manner in which the insect fauna of California approaches 
that of Europe, is in the great abundance of apterous Tenebrlonidae. But in 
this respect it does not differ from a large psj't of South America and by 
the very form of these Tenebrlonidae, which bear no resemblance at all to those 
of Europe, the greater relation of the Californlan fauna to that of the rest 
of America is clearly proved. It will be seen, too, that the resemblance to 
European forms in tbe other tribes is only Indirect, proceeding solely from 
universal or zonal forms, while the greater relation le again with the rest 
of America. It will moreover be seen, that while the stronger relation of the 
fauna Is continental, yet a sufficient number of individual peculiarities are 
Introduced to prove that tt conHtltutes a system of Its own, bearing no relation 
to that of Eastern America, except the slight continental resemblance proceeding 
Indirectly through the tropics, pp. 251-262. 

"The principles shown by the preceding analysis may be expressed briefly as 
follows: 

1. Catirornia constitutes a peculiar zoolc^Ical district, with sufficient relation 
to the other districts of America to prove that It belongs to the same continental 
system. 

2. This zoological district is divided Into several sharply defined sub- districts, 
having a very cloee resemblance to each other. 

As tbe same mode of distribution obtains In the group of Islands adjacent 
to the western coast of America, we are led to believe, 

3. That the local distribution of a small number of species is tbe characteristic 
of the eastern Pacific region, as the extensive distribution of a large number 
Is the prevailing feature of the Atlantic. 

4. The genera occurring in. but not peculiar to. this district, belong to two 
classes: either they occur on the Atlantic slope of both continents, or they 
ere peculiar to America, and are also found within the tropica." pp. 253-25*. 

1859. pp. III-V. "Before proceeding to consider tbe special material usedlntho 
preparation of this memoir, it will be proper to give a short sketch of the 



BCOLOOT OP ISLE ROYALE. 171 

general results thus I&r obtained regardlns the geographlcKl dlHtrlbutfon of 
Coleopterous insects In the terrttor; of out republic. 

"The whole region ol the Unlteil States 1b divided by meridional or nearly 
meridional It nee Into three, or perhaps four, great zoolt^cal districts, dls- 
tfnenlBbed each by numerous peculiar genera and species, which, with but few 
exceptions, do net extend Into the contiguous districts. The eastern one ot 
these extends from th« Atlantic Ocean to the arid prairies on the west ot 
Iowa, Missouri, and Arkansas, thus embracing (for conveDience merely) a 
narrow strip near the sea-coast of Texas. This narrow strip, however, belongs 
more properly to the eastern province of the tropical zoological district ot 
Mexico. 

"The central district extends from the western limit of the eastern district, 
perhajiB to the mass ot the Sierra. Nevada of California, including Kansas, 
Nebraska, Utah, New Mexico, Arizona, and Texas. Except Arizona, the entomo- 
It^cal fauna of the portion ot this district west of the Rocky Mountains, and 
in fact that of the mountain region proper, is enttrelv unknown; and It is 
very probable that the region does in reality constitute two districts bounded 
hy the Socky Mountains, and southern contlnnatlon thereof. 

"The western district Is the maritime alone of the continent to the Facttlc, 
and tliua includes California, Oregon, and Washington territories. 

"These great districts are divided Into a number of provinces, of unequal 
size, and which are limited b; changes In climate, and therefore sometlmea 
distinctly, sometimes vaguely defined. 

"The Atlantic district may be divided Into: 1, a northern province, Including 
Maine, Sastern Canada, Nova Scotia, Newfoundland, etc., and extending weet- 
wardly from Lake Superior to Lake Winnipeg and Western Canada, which fades 
Insensibly Into the great Arctic district; 2, a middle province, limited weetward- 
ly by the Appalachian chain, and extending to Southern Virginia; 3. a western 
province, including Minnesota and the States of the valley ot the Mississippi, 
as far as the State of that name; 4, a southern nrovlnce. including the States 
south of Virginia and Kentucky; G, a subtropical province. Including the point 
of the peninsula of Florida; 6, a subtropical province, including the sea-coast 
of Texas. 

"The Central district, aa far as known, may he thus divided: 1, a northern 
province, comprising the regions north of the Missouri, the plains of the Sas- 
katchewan, etc.; 2, a middle eastern province, divided Into two subprovlnces, 
including: a, Kansas, and Nebraska; b, northeastern New Mexico;' 3, a south- 
eastern iH-ovtnce, Including Texas, with the exception of province six of the 
Atlantic district; 4, a southwestern province. Including the upper part of the 
volley of the Gila; and 6, a south-southwestern province. Including the lower 
Gila and Colorado. The unexplored portions of this district will Indicate middle 
western, and northwestern provinces, or perhaps the necessity ot constituting 
with them and the southwestern province a district to be called the Interior 
district. 

"The Pacific district may be divided as follows: 1, a hyperborean province, 
consisting of Sitka and the neighborhood; 2, a northern province, including 
Eastern Oregon and Washington; 3. a middle province. Including Cali- 
fornia probably as far south as Santa Barbara; 4, a southern province. 
Including California from Santa Barbara to San Diego, extending to the crest 
of the Sierra. Southern, or lower California Is also, perhaps only In part, a 
province of this district;* but, as yet, no collections of magnitude have been 
received therefrom. Other provinces will, from the peculiar method of distri- 
bution of species In that portion of America, be defined when more full collec- 
tions are mode, but at present cannot he indicated. 

"At the north, the Atlantic and Central districts seem to merge Imperceptibly 
together, about the valley of the Athabasca, and Winnipeg rivers, and finally 
to disappear In the limited Arctic fauna; the hyperborean province of the Pacific 
district also tares into this Arctic fauna, without, however, losing Itself so 
perfectly In the northern provinces of the other districts. We have thus evi- 
dence that the American Arctic distrkt may be divided into two provinces, an 
eastern and a western, 

• " A revr ipedea, collecled by John XantuB, Esq.. at C>p« San J.ucaB, though all new, Indlntte a 
girsler resembhtnn to Ihe EauDS ol the lonei Colonido, than to that of marUlme CaUlomla l. IIUb 
province may therefore bo (ound evBntu«Hy to belong to .the interior distrlol." OvIC 



172 MICHIGAN SURVEY, 1908. 

"At the south, the Atlantic dlBtrlct merges through Florida Into the Carlbbeao 
tropical province, aod through maritime Texas into the Mexican lower eastern 
province. In the same direction the Central district merges Info the Mexican 
upper or central province, and the Interior district, towards the Guir ol Cali- 
fornia, Into the Mexican western province. Regarding the southern afllllatl<MiB 
of the PaclSc district wo know ^solutely nothing; scarcely a single species 
found at San Diego had been found In Mexico. 

"The method of distribution of species In the Atlantic and Pacific districts, 
as already observed b7 me In various memoirs, Is entirely dlfFerent. In ttie 
Atlantic district, a large number of species are distributed over a large extent 
of country; many species are of rare occurrence, and In passing over a distance 
of severaJ hundred miles, but small variation will be found in the species 
obtained. In the Pacific district, a small number of species are confined to a 
small region of country: most species occur In considerable numbers, and in. 
trardllng even one hundred miles. It la found tttat the most abundant speclea 
are replaced by others, In many Instances very slmflar to them; these small 
centers of distribution can be limited only after careful colIectlooB have been 
made at a great number of localities, and it is to be hapeA that this very 
Interesting and Important subject of inveetlgation may soon receive proper at- 
tention from the lovers of science on our Pacific shores. 

"In the Central district, consisting, as It does to a very large extent, of 
deserts, the distribution seems to be of a moderate number of species over a 
large extent of country, with a considerable admixture of local species; such 
at least seems to be the result of observations In Kansas, Upper Texas, and 
Arizona." pp. III-V. 

1860. pp. 2.4. "The distribution of species In the northern part of the region 
which furnishes the materials for this report [Pacific R. R. Report], presents no 
remarkable phenomenon. As In other northern lands, certain tribes like Adephaga, 
Staph yllnidae, and Elatertdae assume a greater predominance in the fauna, from 
the fading out of the groups more characteristic of warmer climates, while a 
greater number of species are found common to both continents. Of these latter, 
about one-half are found on the Atlantic slope of America, while the other half 
have not yet occurred there. 

"The number of species occurring on both sides of America is also largely 
increased In these northern regions, but with the exception of Epiphania comutiu 
and PriognatHua monilicornia, the genera of such speclea are distributed on 
both continents. 

''On proceeding southwards to Oregon (and Washington Territory, wblch Is. 
for purposes of convenience, always Included when Oregon Is referred to In 
these pages), similar phenomena may be observed, though on a diminished 
scale. The species of the eastern continent, not found on the Atlantic slope 
of America, have entirely vanished, and of the species common to both sides 
of both continents, but four remain. The number of species common to the 
Atlantic and Pacific slopes oF America has greatly diminished, and among 
them Haplocltile pygmaea, Ligyrtta gibboaus, Alaus myopa, and Jficrorhopala 
vittata are the only representatives of America genera. 

"Finally reaching California, the species common to the two continents are 
reduced to SUpha lapponica and Dermestea vulpinu*. the species common to 
Atlantic and Pacific America have not diminished absolutely in number, but 
from the more complete and copious fauna known to their relative proportion 
is much lessened. Among them, however, are found but few which extend 
their range to ttie Atlantic States proper, while the greater proportion are not 
found east of Kansas. Of American genera, Ambljichila cylindriformia, Lacftno- 
phorus eleganttilus, and Earmetopon a (rum are found In Kansas, or New 
Mexico, while Ligyrua gibboaua and two apeclee of Diabrotica also extend to 
the Atlantic," 

"In Russian America the genera seems to follow to a certain extent the 
course already pointed out of the species, that is: the genera common to both 
continentB have a much greater relative proportion, and among them a by no 
means insignificant part have not yet been found In Atlantic America; but 
as some o( them are ctiaracterlstic of high northern latitudes, there Is reason 
to believe that the number will be reduced by more thorough explorations in 
Labrador. Newfoundland, and the regions near Hudson's Bay. 

"Of genera confined to America, but six or seven occur In Russian America; of 



ECOUXJY OF ISLE ROYALE. 173 

tbese but three. PiiBtodactyla, Eplpbanls. and Prlognathua. have been <1etected 
on tb« Atlantic slope. Prlatodactyla might. Indeed, be for the present excluded 
from the list of peculiar American genera, for two reasons: 1. a certain number 
of epeclee classed by Dejean. with Agonum, and remarkable for having but 
two dorsal punctures, are In reality Prlstodactylae. and until the species ot 
Siberia are thorouglily revised, we are warranted In supposing that some of 
tbem may also be Included: but, 2, tiecause the distinctions between Calathus 
and Prlstodactyla. as ob»ierved by Lacordalre. are hardly sufflclent to warrant the 
retention of the latter genus. 

"In Oregon the eastern genera, n'ot found in the Atlantic States, have dimin- 
ished in number, but among them occurs Caillslhenes. which is Found In Kansas. 
The number of American genera has largely Increased, even with our limited 
collections; ot them 14 are found In the Atlantic States, 2 In Kansas, while 
3 are peculiar to Pacific America: of the 14 found in the Atlantic States. 
Haplochlle, Dlchelonycha, Anelastes. and Alaus are the oniy ones not found 
within the tropics. 

"In California the Renern of the eastern continent have increased absolutely. 
from more eitenslve collections, over those found in Oregon, but do not attain 
the same relative proportion as those found In Russian America; among them 
Is one, Tryssus. a genus heretofore known only from Madagascar, and Is thus 
far the sole representative of the tribe of Scarabaeldae, to which It belongs 
on this continent. 

■•The number of American genera has greatly Increased, partly by the ad- 
dition of genera found within the tropics, and partly by the Introductlou of a 
few peculiar genera; the most remarkable addition, however, is that of eighteen 
genera of Tenebrlonidae, of which but two. Noeoderma and Blapstinus. extend 
Into the Atlantic States, while only four others extend into Kansas or New 
Mexico. The genera found in the Atlantic States, and not la the tropics, are 
Thai plus, Axinopalpus. Dlchelonycha. Anelastes, Perothops. and Melanactes. 

"Another fact ot great Interest is the distribution of species within narrow 
limits observed in California. I am not able to exhibit the results In a tabular 
form, as collections have not been made with minuteness at a sufficient number 
of localities to give any definite results, but I can merely state my own ex- 
perience, that but few species occurred at more than one place, and call at- 
' tention to the fact that, In every collection made at a fresh locality, a large 
proportion of new species is found, while In Oregon, at points equally distant 
from each other, a greater uniformity la seen. 

"The analysis, therefore, conducts to the same results announced by me. 
in 1S51, at the meeting of the American Association for the Advancement of 
Science; the fourth proposition was, unfortunately, announced In too absolute 
terms, as the only two genera then known to me. Tbalplus and Ailnop&lpus, 
were not considered na of sufficient importance to modify the result. Thalplus, 
Indeed, is to closely allied to Diaphorus, that we may well expect some o( the 
species ot the latter genus to belong to It, while Axinopalpus is by many en- 
tomologists not separated from Dromius. The other four American genera com- 
mon to California and Atlantic America, not found in the tropics — Dlchelonycha, 
Anelastes, Perothops. and Melanacies— upon which I am now obliged to modify 
the assertion, were subsequently obtained. 

"The four propositions mentioned by me in the essay mentioned are: 

1. California constitutes a peculiar zoological district, with sufDceient rela- 
tion to the other districts of America to prove that it belongs to the same 
continental system. 

2. This zoological district Is divided into several sharply-defined sub-districts, 
having a very close resemblance to each other. 

As the same mode of distribution obtains in the groups ot islands adjacent 
to the western coast of America, we are leii to believe — 

3. That the local distribution of a small numlwr of species is the character- 
istic of the eastern Paclfle region, as the extensive distribution ot a large 
number Is the prevailing feature of the Atlantic basin. 

4. The genera occurring In, but not peculiar to, this district belong to 
two classes; either (with the exception of Ergatesl they occur on the Atlantic 
Slope of both continents, or. If peculiar to America, they are (with the few 



exceptions above noted) also found within the tropics." 
This paper is accompanied by four tables as folloi 



.CocH^le 



17i MICHIGAN SURVEY, 1908. 

I. Genera Common to the Eastern and Western Continents. 

II. Genera Peculiar to America. 

III. SpecleB Common to the Atlantic and Pacific Slopes of the Continent. 

IV. Species Pounrt In Russian America and In the Eastern Continent, not 
Introduced and not Found In Atlantic America. 

1862. p. 336. "Some of' the more conspicuous and peculiar species are des- 
cribed below: enough has been stated to show that the aiBnItles of the fauna 
[of Lower California] are with that of the region extending from the Colorado 
Desert across to the Rio Grande valley, thereby confirming the results obtained 
by Prof. Balrd and Mr. Cope from the study of the vertebrata collected by Mr. 
Xantus. 

"The limited tiumber of speclee oF these two classes precludes the possibility 
of the occurrence of many new rorms In the region here treated of; but in the 
number of peculiar species of the much more extensive class of insects seen 
!n Mr. XantuE' collections, we recognize that lower California constltuteB one 
or more provinces of the Interior district, aa denned by me In the Introduction 
to my synopsis of the Coleoptera of Kansas and New Mexico. 

"The preponderance of Tenebrlonidae, both in genera and species seen In the 
fauna of Upper California and Arizona, has here been partially deetroyed. Tbe 
genera which survive are, however, such aa are already known from the last 
mentioned region. None of those peculiar to maritime California have as yet 
occurred." p. 336. 

1878. pp. 447-448. "The elevated interior region of North America presents 
peculiarly favorable opportunities for the study of some of the most interestlog 
questions connected with geographical distribution of animals and plants. 

"If the materials at our handa be. as Indeed they yet are, a very scanty 
representation of the organic forms now living in that part of the continent, 
they are, at least, sufficient to Indicate the direction in which Inveatigatlons 
should be pushed, In order to arrive at definite and final results. 

"The peculiarly favorable circumstances to which 1 chlefiy refer at present 
are dependent on the following points in tbe development of the region: — 

1st. The gradual enlargement of the land-surface at the expense of tbe 
circumambient seas during the latest Mesozolc periods. 

2d. Tbe gradual elevation of the middle of the continental mass during post- 
Cretaceous times, so as to greatly modify the climate In respect to both moisture 
and temperature. These changes have been ao gradual, that we may say with 
certainty (excluding the local eruptive phenomena, which were more numerous, 
but not remarkably different from those of tbe present age) there has been 
no great or paroxysmal disturbance destructive of the land -surface In the 
elevated plains east of the Rocky Mountains since the deposition of our early 
Cretaceous strata (Daliota Group). 

3rd. While, during tbe Glacial epoch, the valleys of the mountains were 
filled with glaciers of moderate size, and the line of permanent Ice streams 
and fields brought to a much lower level, there was an absence of the extensive 
ice sheets and fiooded areas, which in Eastern America destroyed entirely the 
terrestrial organized beings of the former period. 

"It must be inferred from the first and second of these premises that tbe 
new land exposed by this gradual development of the continent received Its 
colonies of animals and plants from the conterminous older land-surfaces in 
various directions, and that the subsequent elevation of the continental mass, 
by which the moisture was diminished, caused a later invasion of the territory 
by those genera and species which are characteristic of arid regions. 

"We may also conclude, from the third premise that the glacial displacement 
of species In the Rochy Mountains has been much less than in Eastern America, 
and that a very small area would be left bare of life on the return to a normal 
temperature; consequently, the previous occupants of the higher mountains 
would again return to their former domain, increased by refugees from the 
circumpoiar continent of temperate climate, driven southward by tbe Increas- 
ing cold, 

"Such being the case, it ought to be possible, with well-prepared lists of the 
lnse<'ts of the plains and mountain regions, by comparison with Hats of tbe 
local fauna of other zoological districts of the continent, to ascertain, with 
reasonable probability, the invaalona from different directions by which, in the 
first place, the newly emerged land was colonized; and., in the second place. 



ECOLOGY OF ISLE ROYALE. 175 

the modifications, either in distribution or In structure, which have aubsequently 
occurred. 

"I h&ve on an other occasion' expressed my belief that the study of the 
distribution of existing Insects could give much information concerning former 
topographical and geographical chajigea in the surface of the earth. I then 
gave several examples to sbow how the distribution of species peculiar in their 
habits and structure conflrmed what was already known by geological investiga- 
tion of tbe gradual evolution of the middle part of the continent. I will now 
advance the additional thesis, that we may obtain somewhat deflnlte Informa- 
tion of the sequence, extent, and effects of geological changes in tbe more recent 
periods by a careful study ot tbe insect fauna in Its totality." 

1878b. pp. 470-471. Includes lists oC Florida Coleoptera: 

1. Florida species also found In the Antilles. 

2. Common to Florida and Mexico and partly found in Texas. 
?.. Common to Texas, Arizona, and southern California. 

4. Anomalous common to Florida and South America. 

5. Bistrlbutlon of anomalous species. 

Hurray, A. 1870. pp. 7, 8, 11-12, 32.33. 3G-37, 38, "The position I am about 
to maintain then is, that, subject to modifications to be afterwards mentioned, all 
the Coleoptera in the world are referable to one or other of three great Htlrpes. 
These three no doubt originally sprung from one stlrpe, and acquired their dls- 
ttnguishing features by long-continued isolation from each other, combined with 
changes In their conditions of life. But now we have three, and only three, 
great strains, sometimes Intermingling with each other, sometimes underlying 
or overlying each other, and sometimea developed into new forms, but always dls- 
tlorulshable and traceable to one or other of tbe three sources. 

"These are — 1, the IndoAfrlcan stirps; 2, tbe Brazilian stlrpe: and 3, what, 
for want of a better name. I shall call the microtypal stlrpe, in allusion to tbe 
general run ol the species composing It being of a smaller size, or, more strictly 
si>eaking, not containing such large or consplcucnis insects as tbe others. It la 
not altogether a satisfactory name, because the stlrps does contain some large 
species, and it is not peculiar to it to abound In small ones. But, taken as a 
wb(de, its ingredients are smaller and more modest in aK>earance than those 
of the others. Tbe fanna and flora of chit own land may be taken as its type and 
standard, pp. 7.8. 

"Tbe iDdo-African stlrps. as Its name Impllee, Inhabits Africa south of the 
Sahara, and India and China south of the Himalayas, also the Malayan district, 
tbe Indian archipelago, and the New Guinea group. This range is less modified 
by the general Introduction ot foreign elements than that of tbe next stirpa. 

"The Brazilian stlrpe Inhabits South Central America east of the Andes, and 
north of the River Flatte, and furnishes, moreover, a large share In the constl- 
tntloB of North America, but has also received in return a very perceptible tinge 
from the microtypal stlrps. 

In the microtypal stlrps I include the fauna of Europe, Asia north of the 
Himalayas. Baalern North America, bo tar as not modllled by the Brazilian element, 
and, what has lees of this strain, the whole of North west America, California, 
part of the Mexican fauna, Peru, Chill, the Argentine Republic south of Tucuman, 
Patagonia. Tlerra del Fuego. Polynesia, New Zealand, and Australia, p. 8. 

"Let'uB now turn to the three great stlrps, and pass each of them in review, 
trace their course, and determine their limits. 1 shall begin with the micro- 
typal Btirpe (with which we are most familiar). It is the most extensive of the 
whole, being distributed over the whole world with tbe exception of the In- 
dian, African, and Brazilian regions; and even they, from various exceptional 
causes, have a greater or less tinge of it In their faunas. It contains some 
minor faunas, and these, again, a. number of subfaunas. The Buropeo. Asiatic 
region Is one of these minor faunas, and of it the Atlantic islands, the 
Mediterranean, and tbe Monoglian are subraunas. Taken as one fauna, 
the Buropeo-Aslatlc extends from the Azores esst to Japan, the whole 
of that vast space being inhabited entirely by the same type and, for the most 
part, by the same species, a few only dropping off here and there, and being re- 
placed by others of the same general character, p. 11. 

1. TnuiB. Am. Assoc. Adv. Science. 187S, Detroit. Prealdenl 's address, [f/, l,f Ccaitf , 76.1 



yCoogle 



176 MICHIGAN SURVEY. 1908. 

"The Buropeo- Asiatic Beetle- fauna' does not stop even at Japan; It paBses over 
Into North America by Behrlng's Straits, or rather, I should say. It Is found In 
North America on the other side of BehrlDg's Straits. In Russian America we 
have a fresh crop of Europeo- Asiatic form, genera and species; and here another 
noteworthy circumstance presents Itself. It. is generally taken for granted that 
there Is a unftorm homoeeneous arctic fauna which estends all around the arctic 
circle. It Is so, and It Is not so. It Is bo on the large scale, but not so on the 
small. The arctic fauna Is subject to the laws of spreading by continuity and 
stoppage by barriers just the same as any other fauna. I have elsewhere endeav- 
ored to show that the mammalian fauna of Greenland la Europeo -arctic as dls- 
tlnguBhed from Amerlcano-aretlc. I maintain that the homogeneity of a fauna 
depends on other causes than uniformity of condition of life within its limits. 
I cannot doubt that if there had been an isolated communication between the 
£ndo-African districts and the North-Pole, we should there have had a fauna 
related to and developed out of that fauna, and wholly distinct from the other 
faunas ot the arctic regions. It is continuity of soil or freedom of intercommuni- 
cation which has produced the present uniformity of fauna in the arctic regions; 
but were minor interruptions exist, or old barriei-s or conditional equivalent to 
a. barrier formerly existed, there are also subdivisions In the character of the fauna. 
and in the position of these minor divisions we see the operation ot these laws 
and are able to trace the existence and former position of the barriers. Thus 
we find two minor subfaunas In Arctic America, an eastern and a western 
one. Two causes may have produced these. One of these may have been 
the sea which. It can i-carcely be doubted, formerly existed between the 
Oulf of Mexico and the Polar Sea, in the Hue of the Missouri and Mackenzie 
rivers; another may have been that the ground now occupied by one at these 
subfaunas was under water at a later period than the other, so that it was 
peopled at a different date from it. Probabiy both contributed to produce the 
present arrangement of the subfaunas to the east and west of the Mackenzie Kiver. 
That there was a barrier there, and that that side was still supplied with the same 
general type (though with mjnor devlatioas). is to be explained by their having 
received their species from the same general stock, but coming to It from dif- 
ferent directions, the one from the east, the other from the west. That the minor 
differences to which I allude are. in the case of North America, to he referred 
this cause, and not to mere gradual Increase of variation arlaln;; from Increase of 
distance, seems to be a legitimate Inference from the fact that while the whole 
of the north of North America, without exception, belongs to the Europeo- Asiatic 
type, there are a number of European genera which occur in North-east America, 
and not In the North-west, and a few tchich occur in the Xorth-west, and not in 
i/orth-east America, pp. 32-33. 

''Returning to the Asiatic terminus ot the microtypal stirps, let us now en- 
deavor to trace its further couFse. The genus Blaps. which Is a characteristic 
feature in the Coleopterous fauna ot Central Asia, will furnish us with the means. 
It may be taken as a representative case applicable to other species also, although 
it is the most striking instance which occurs to me. Upwards of 100 dltfereot 
species ot Blaps, out, of a total ot about 150. have been described as Inhabttiag 
the country between Southern Russia, Mongolia, and Mantchourla. Now If we 
cross to California in continuation of the same line we have not Blaps, but we 
have Blaps's brother and he baa been a twin. We have Eleodes. Its perfect 
counterpart and representative; and it is to be observed that while the faciei 
of the si^ecies actually inbablllni: California is entirely that of Biaps. a number 
4>t species which are found in Kansas and on the eastern flanks ot the Rocky 
Mountains have a somewhat different facles: and I should add that the suppoal- 
-tion that these are stra^Klers from the Calirornian shores is strengthened by the 
fact that the genus does not occur to the east of the Missouri; other Heteromeroua 
forms, reminding us of Mediterranean and Asiatic species, occur in California, 
and the whole of the north. west of America has a greater preponderance of the 
microtypal stirps than perhaps occurs east of the Rocky Mountains, pp. SG-ST. 

"Next step to the south of Calltornla comes Mexico. It also is lari;ely supplied 
with Eleode»: and although some of the showiest and finest non-microtypo! Col- 

■■ ' 1 Has unulilc in my *(li-UKrniililinil l>i.«rilnuion of MummuiN' lo pili.[it Dr. ttclalerN t*rmlnoloKy 
Of Pulup.i relic, Ntoarctfr. Ai.. I)i'('jii.si- ui- rliil noi atntv iii the ncleiil nnil 1 mils oC oiir n^lonii: iiixt 
now, of rourw. In tlilN piinpr 1 iiiii iIIH Ifss iln so. 'an a nrllii'ItMl i-rfi-n of mv hyiiu*bF»is. if it be 
Bounil. niu»t be lo mill furtliiT lireak down tliclr limlti^ niiU iksiruy i\kii iiolldily.'- 



ECOLOGY OF ISLE ROYALB. 177 

eoytera In the whole world come rrom Jlexlco. they have no bearing on tbiB part 
of my inquiry; for they cotae From parts of Mexico which are In direct com- 
manlcatlon with another stirps, the rich Coleopterous fauna of Brazil and Vene- 
zuela: and the vast multitude of small European-looking species which occur 
on the high lands and western side. la quite suEBclent for my purpose. The col- 
lectiotta made by TruquI In Mexico sbow this thoroughly microtypel character 
in a very marked way, Staphyllnidoua genera, eueh as Falaffria. HoTiuilota, ftc., 
abounding. Mexico, being a sort of halfway houEe between Europe and Australia, 
might be expected to contain species both from the north and the south which 
have got thus far. Eleodia Is an Instance of this from tbft north, Phitonthua 
another; both reach as far as Chili, but not Into Australia. Zophera*. on the 
other hand, is an instance of a species wbicb occurs in Australia, and runs up 
into Mexico, where It Is In strength, and goes even a little further. Mexico may, 
indeed, have been its starting-point, but the connexions and relations of It and 
the allied genus Notodendron decidedly indicate a separation between the eastern 
and western type of both; and the weateru type extends into Australia and New 
Caledonia." p. 38. 

Schworr. E. A. 1888. pp. 166-167, 168-170, 171-172. "After a study of this 
peculiar fauna of Key West which I also found on many other localities farther 
north and which constitutes the eemitroplcal fauna of Florida, I have come to 
the conclusion that it is entirely of West Indian origin, and that the region I shall 
hereafter circumscribe as Semltropical Florida does not contain any endemic 
forms. In other words, the distinctive fauna of Southern Florida Is a permanent 
colony of West Indian forms, much more numerous In species than it has 
hitherto been supposed; the number in Colcoptera alone amounting, accord- 
ing to a very low estimate, based upon my collection, to at least 300 species not 
yet in our catalogues, pp. ieG-167. 

"Before entering on a discussion of the character and extent of this West Indian 
colony in Florida it seems worth while and Instructive to give a glance at the 
Bouth-western extremity of North Ameiiica where our fauna comes also In contact 
with a semltropical fauna. The great faunal regions known as Nearctic and 
Neotropical are connected or divided by the Central American fauna which from 
the nature of the conditions participates in the characters of Iwtb regions, but 
Is more nearly allied to the latter than to the former. It is again divided lata 
the fauna of the Central American continent and the Insular fauna of Central 
America, more commonly called the West Indian fauna; ttaeee two faunal regions 
t)eiag related to each other in the same degree as Is the fauna of our Atlantic 
slope to that of the Pacific slope. At the zone of contact between the North 
American fauna and that of Mexico the conditions are as follows: The ocean 
current along the PaclHc coast of North America runs from north to south, thus 
facilitating the spread of more northern ' species southward. It loses its force 
and disappears before reachfng southern California and thus the Nortb American 
fauna along the coast does not come into contact with that of the Mexican coast. 
On the mainland we find between California and the largest portion of Arizona 
on the one side and Mexico on the other, a broad tract of the most barren and 
sterile* country which proves to be a moat effectual barrier between the two 
faunal regions. Farther east, and more especially along the Rio Grande, a complete 
intermingling of the two faunas takes place In such a way that species of ail fam- 
ilies participate In this intermingling. It is thus impoaslble to decide whether a 
collection of insects comes Irom Texas or the State of Tamaullpas, or whether 
it comes from southern New Mexico, from south-eastern Arizona, or from Sonora. 
The Morrison collection, for Instance, has been distributed among North American 
entomologists as coming from south eastern Arizona and la worked up in the 
'Blologia Contrail Americana' as coming from Sonora, Mex. pp. 167-163. 

"In looking for the original home of this colony of West Indian insects and 
plants we have been hitherto too much accustomed to consider the island of Cuba 
88 the only place from which tbls immigration has taken place. In the task ot 
determining my South Floridlan Coleoptera it was found over and over again 
that these Immigrants may have been described not only from Cuba, but from any 
other of the West Indian islands, or from the Central American continent south 
of Yucatan, or even from Columbia and Venezuela — in other words from all 
parts of Central America which come under the Influence of the Gulf stream. As 
can be seen from any physical atlas, the warm equatorial current enters the 

'BiolQ(!ia Cenrrali-AmiTirGnj,' " Trans. Amec. Ent. S^C 



178 MICHIGAN SURVEY, 1908. 

Caribbean Sea througli the Windward Islands and attaining by this contractlMi 
a considerable velocity forms the Gulf Stream which flows between the southem- 
moet chain of the West Indies and the Leeward Islands and etrlkes the Central 
American continent, flowing northward along the coast. Deflected by the project- 
ing peninsula of Yucatan, the stream turns eastward and reachea the coast of 
Cuba and the southernmost part of Florida. Thus the West Indian colony of 
Insects In Florida may come from any part of this vast area swept by the Gulf 
stream, although the largest proportion comes of course from Cuba since this 
Island is the nearest to Florida. This Immigration by the aid of the Gulf stream 
explains the following Interesting phenomenon In geographical distribution. We 
have seen that Insects from the coast of Central America aouih of Yucatan may 
occur In Southern Florida; but the same species often had the power of exteudlng 
their geographical distribution northward on the Central American mainland 
through Mexico, thus reaching the south-eastern limits of the United States. 
Certain species may occur, therefore, in the United States, In Western Texas 
or South-eastern New Mexico and In Southern Florida, being however, 
^sent In the intervening Southern States, viz: Eastern Texas, Louisiana, 
Alabama, Georgia, and Northern and Central Florida. This curious distribution 
"has never been pointed out so far as I am aware but can be exempllfleil by nnm- 
■erouB species, not only among the Coleoptera but also other Orders of Insects. 

"The distance between Cuba and Florida Is not very great, the current of the 
Gulf stream Is very swift, and logs and other debris swept out to sea from the 
rivers of Cuba may reach the coast of Florida within three or tour days; from 
Yucatan in about double that time. It is evident that within that short time ail 
^stich insects may safely be carried from the West Indies to Florida which. In the 
Imago or preparatory stages, live under bark, or within the wood of tre*«, or 
within seeds and similar sheltered conditions, or whose eggs are firmly attached 
to trees and covered with viscous liquid. But It is evident that this sea voyage 
la too long tor all such Insects as do not live In such sheltered positions. As a 
consequence, all odephogous Coleoptera, further all those living under old leaves, 
in the ground, In very rotten wood and similar places, and finally moat of the 
Chrysomelidae which lay their eggs either onto the leaves or in the ground are not 
brought over from the West Indies. There are, therefore, no West Indian Cara- 
bidae, Lampyrdae, Btaphj/Unidae and other rhypophagous Clavlcorn families and 
very few West Indian Scarabaeidae and ChTyiomelidae to be found In Southern 
Florida.* This Is a meet character Istic feature of the semltroplcal Coleopterous 
fauna of Florida, strikingly contrasting with the state of atlatrs In the south- 
western extremity of North America. I have stated before that along the Texan 
and New Meilcan frontier there Is a perfect intermingling of the North and Central 
American faunas so that It Is Impossible to decide whether a miscellaneous col- 
lection of Coleoptera comes from Western Texas or the adjacent parts of Mexico. 
A miscellaneous collection, consisting only of about 100 species but made pro- 
miscuously in semltroplcal Florida can at a glance be distinguished from a similar 
collection made In Cuba or any other part of the West Indies. Further, the pecu- 
liar composition of this fauna at once precludes the assumption that any agencies 
other than the current of the Gulf stream could have been active In assisting the 
Immigration from the West Indies, pp. 1S8-170. 

"Most of the more southern Keys are covered with semltroplcal forest, i. e. 
forest covered with composed of West Indian trees, while, as I stated before, the 
true Floridian fauna and flora are almost entirely absent. These Islands are, there- 
fore, by no means favorable to a study of the relation of semltroplcal to the true 
Floridian fauna. However, a stay of a few weeks on the shores of Biscayne Bay 
fully sufficed to settle this question. Here, as well as on the mainland tartlier south 
and the northernmost Keys (Key Largo and Elliott's Key) the Floridian flora 
largely Infringes upon the semltroplcal forest and rednces the same to smaller 
or larger Island-like patches lying close to the shore or occupying similarly isolated 
patches on the shore of the Everglades and the few Islands In the Everglades. 
The bulk of the mainland Is covered by pine woodst with an undergrowth com- 

" > The absence of freeh naler In the coral regloii of the keys and 1h« mainland south of Miami 
River necesailates the nljaenci! of iiytitcidae and mosi other aqunclc or semi aquatic funlUes. Even 
the Everglades and [he HTers draining the same at the northern «id of Biscayne Bay aeem to be al- 
most destitute of aquallc Coleoptera. 

" t While it is true tliat the pine of Soulbrm Florida, Pi'nus Cubeiuii. is also of West Indian orl- 

Sn. its distribution in Florida is quite dlffprtnt from in* tfst of the serailropical tlMa Uid Its Intro- 
ictlon Is pTldt'ntly of very andcnt date. Its fauna do>.!i not difttr from that of the Yellow Pine, 
(P. ralaf.ris)." 



ECOLOGY OP ISLE ROVALE. 179 

posed almost entirely of true Plorldlan planU. There are further vast stretches 
of what la called 'the pralrla.' I. e. land quite recently formed, partly by the accuinu- 
latlan of seaweeds swept asbore by the waves, aud partly by the advance of the 
Mangroves. This prairie is covered witb the same herbaceous vegetation wblcta 
we see la similar places in Central Florida and does not contain a single semi- 
tropical plant. Even the hammock Is Invaded by several Florldian trees: the 
Live Oaks, several Palmettos, the Hackberry and others make their appearance 
and. oo higher ground we find plenty of Pertea carolineniis. Now on all these trees 
In the pine woods and on the praJrle, In short wherever there Is the Florldian 
flora we meet the true Florldian Insect fauna whereas the semitroplca) fauna Is 
coi'fined to the semltroplcal forest.* This fact once recognized, It becomes evi- 
dent that the northward extent of this fauna is identical wltb that of the semi- 
tropical forest, a fact fully borns out by subsequent experience." pp. ITO-lTl. 

"I desire to emphasize here once more as one of the principal characteristics 
of this Bora and fauna, that north of the Everglades they nowhere appear Inland 
but always close to the shore. Even along the inner bank of the Indian River 
there are— or rather were— hut a very few spots covered with semltropical forest, 
viz: on the mouth of the St Lucie and Sebastian Rivers, at the southern end of 
Herritt's Island and perhaps some others; but they are now mostly destroyed by 
cultivation." p. 172. 

1890. pp. 1S6-1ST. 

"The mountain ranges In America run in the direction from north to south, 
and the colonies of circumpclar Insects upon their summits have thus been able 
to preserve their connection and specific Identity with the arctic forms; whereas 
In Europe, where the mountain ranges run from east to west, the alpine colonies 
have generally undergone changes and, by Isolation, lost their specific identity 
with the arctic species. There Is, therefore. In the Old World an abundance of 
distinct alpine forms, none of which are Identical with North American species; 
while we, on our high mountains, have but few, If any, alpine, but more arctic 
forms, pp. 18S-187. 

"Among the strictly circumpolar Coleoptera the predaceous famlltes predom- 
inate over the phytophagous families; the Carabidae. Dytiaddae, BtaphyHnidae, 
and Coccinellidae are well represented, the Cliryaomelidae and Rbpnchoptiera are 
tolerably well, and the Ccrambycidae and Etateridae are poorly represented. The 
Bitprestidae are absent although this family contains numerous boreal species 
In every region. The phytophagous Bcarabaeidae do not, or barely extend into 
the arctic regions; the coprophagous 8carat>aeidae (Apliodiua) are well repre- 
sented there, still none of them (with the exception of Aphodiua ruflpes, which 
doubtfully belongs here) is on the list of circumpolar Coleoptera," p. 187. 

"Species not Belonging to the Circumpolar Founo.^This division compriaea 
endemic species of probably intratropical origin, which have spread, by natural 
dispersion, into the temperate zone of North America." p. 1S7. 

1890a. pp. 170-171. 

"Turning now to the bulk of the species in the list [St. Augustine, Florida] we 
find that they consist of the usual admixture of more or less widely-distributed 
species and true Florldian forms, the proportion being but little different from that 
of the other localities, e. g., Crescent City, Enterprise, Taropa. . , But the St, 
Augustine list contains another element, viz: species belonging to the faunal 
region lying directly north of eastern Florida and comprising lower Georgia, the 
lower Carollnas, and eastern Virginia. This Is an ill-dehned region with very 
few. or no, peculiar spectes, and only characterized by a certain combination of 
a number of southern species. The existence of this faunal region will become 
evident to any one who, on a summer day, goes from here [Washington] down 
to Fortress Monroe, Va. The difference between the Washington fauna and that 
ot Fortress Monroe will then be found quite striking. Of this fauna I noticed 
about twenty species in the St. Augustine list not previously known from Florida." 
pp. 170-171. 

1901. pp. 1. 2, 3. 

"Still, southwestern Texas belongs, at least as far as the Insects are concerned, 
to the lower Sonoran fauna, of which it forms a marked subdivision, but wltb 
marked affinities to the austrorlparian region. 

* There la. in ■ddltion. Id Southern Florida a maritime fauna of Ktnitroplcii cturacter, but the 
.1 ■ — .J 1 — .i.„ .jmg (iboul 13 la Coleoptera) Is ao anal! that it Is hardlr worth 

^. itent la atUI uQcerlain but It la aate lo say that on the eastern coast 

hbeyond Uoaqiillo lalel at New Bmynia." 



ISO MICHIGAN SURVEY. 1908. 

"Collections made at Laredo. San Dl€go. Corpus Chrlsli and In the lower Nueces 
river valley prove that, with (ew exceptions, no tropical forms occur In that aectioo, 
and the trip on the stage from Alice to BrownBville shows that the character of 
the coiintr<' does not change southward until the black alluvial SO'II of the delta 
of the Rio Graude Is reached. Here, within the bends of the river, as well as 
along the various bacUwaterp and old river arms (resocaa) which dlsaect the delta. 
Isolated areas or strips of larger or smaller extent are covered with a dense forest 
having thick underKrowth of varied shrubbery and ft rich vegetation of lower 
plants, the like of which Is not Keen at any other place In Southwestern Texas. 
The forest jungles (In Florida tbey would be called hammocks) are the home 
of the semltropical irsect fauna of Texas, which, so tar as known to me, has, 
previous to the year 1S9&. never been Investigated hy any entomologist, since even 
many of the most abundant species are either entirely new or not yet recorded 
from the United States. If, confining myself to Coleoptera found by Prof. Town- 
send or mysuil near Brownsville, I mention the genera Agra. Daavdactj/lua, Phya- 
orhinus, Achryton. Onaphalodes. AmpMonycha. MegasceJis, Plectrotreta. Brachy- 
coryne, Liilronvchns. Polypria (quite a number of others are not yet determined, 
or u nd escribed I . no one can deny the existence of a semitroplcal insect fauna along 
the north bank of the lower Rio Grande. The number of species compoalnt; this 
fauna Is very lariie: In Coleoptera alone I estimate that, after proper exploration, 
between 300 and 400 specie will be added to our lists. 

As stated above, these semitroplcal thickets occur In Isolated patches In the 
lowest parts of the delta; wherever the ground is a little more elevated, the usual 
mesqulte and spiny chaparral, liberally Interspersed with Opuntlas. malre their 
appearance, and with ibem the general fauna of southwestern Texas." 

Scudder, 1895. pp. 27-28. 

■'The Post-pliocene deposits have proved the most prolific with thirty-two species, 
though here only seven families are represented, of which the Carabldae and 
Staphyllntdae. but especially the former, very largely predominate. The greatest 
Interest attaches to the interglaclal locality near Scarboro'. Ont,, which alone has 
yielded twenty-nine species,' and is the largest assemblage of Insects ever found 
In such a deiwsit anywhere. These clays have been studied and their fossils col- 
lected by Dr, G. J. Hlnde.t who sets forth the reasons why he regards them as 
interglaclai. tying as they do upon a moralnal tilt of a special character and over- 
tain by till of a distinct kind. The elytra and other parts of beetles found by him 
represent Ave families and llfteen genera; they are largely Carabldae. there tieing 
half-a-dozen species each of Platynus and Pterostlchus, and species also of Patro- 
bus, Bembldlum, Lorlcera ai<d Btaphrus. 

The next family In Importance is the Staphyllnldae. of which there are Ave 
genera. Geodrctmtcus, Arpcdlum. Bledius, Oxyporus and Lathroblum, each with a 
single species. Hydrophiltdae are represented by Hydrochus and Helophorus, each 
with one species, and the Chrysomelldae by two species of Doaacia. Finally a 
species of Scolytldae must have made the borings under the hark of a Juniper 
described below. 

"Looking at the Bsseml^lage of forms as a whole and noting the distribution of 
the species to which they seem to be most nearly related, they are plainly IndtgMi- 
ous to the soil, but would perhaps be thought to have come from a somewhat more 
nortliern locality than that in which they were found; not one of them can be 
referred to existing species, but the nearest allies of not a few of them are to be 
sought In the Lake Superior and Hudson Bay region, while the larger part are In- 
habitants of Canada and the northern Cnlted States, or the general district in 
which the deposit occurs. In no single Instince have any special afflnities been 
found with any characteristically southern form, though several are most nearly 
allied to species found there as well as in the north. A few seem to be most nearly 
related to Pacific forms, such as the Elaphrus and one each of the species of 
Platynus and Pterostlchus. On the whole, the fauna has a boreit aspect, though 
by no means so decidedly boreal as one would anticipate under the circumstances." 
pp. 27-28. Cf. Scudder '94. 

[Jlke, H. 1902. p. 3. 

"The appearance of northern and southern forms are here controlled [Wash- 

'■•This slati'mpnl includo tour sppcfcs (Hy. 
milani, and Brmhtdium IntamfMumi. found b\ 
Krlp, in riiiy Ix-ds v^ry slnillor lo those found 



ly Google 



ECOLOGY OF ISLE. ROY ALE. 181 

Ington, D. C-l by the chang<> of seasoas. ao in early aprlUR we may always evpect 
more northern types, wblle in mldBuntmer the southern ones predominate." 

VanDjfke. E. J. 1901. pp. 198199. 

''The California faunal region proper Includes practically all the lowlands of 
the State, the fertile valleys of southern California ani) the extensive valleys of 
tbp Sao Joaquin and Sacramento, the lesser valleys along the const and the foot 
hills bordering them. The fauna prevailing throughout Ihese portions are bo 
affiliated with Sonoran forms, narticularly towaril the south as to warrant the 
designation of such portions as Sonoran Bub-reglons. and by the extension of 
these forms into the foot hills where they have Interbred with Boreal types 
through a series of aged, genera characterlatlc of both parent regions have l>een 
evolved. Otnu*. Brenrms la cychrld subgenus). Mplriiis. Fromecognattiua. FJeo- 
roma, and Rosalia with others while more or lese related to adjacent northern 
forms probably developed from a rich circumpolar fauna under the Influence of 
adaptation to environment. Omut occurs rather generally throughout the state, 
and Metriut and Framecognathvi ilmllarly but less frcQitertly in the moisl timber 
^elt of the Coast Range, although an Alpine variety of Metriun is found In tha 
Sierras, and Brennua Is confined to the coast. Many other examples of restricted 
location could be given. In earlier periods California was more Isolated partic- 
ularly from the Sonoran region and northern Influences prevailed. Then such 
genera as Omua and Piecoma became first established. Subsequently a few sotith- 
»rn forms such as Contontia and Its congeners gained access. These constituted 
the old California fauna, but when the southern Isolation ceased, followed by the 
invasion of Sonoran forms, a new and later fauna was developed. This IbMry 
Is partially supported by the fact that in the Islands off Lbe coast and In certain 
still Isolated areas are faunas which are largely sui generii. and typical of the old 
California fauna above described." 

Wlckham. H. F. 1902. pp. 221-222. 

"The phenomena of distribution in Colorado are of much Interest. Within a 
radius of a few miles we may And assemblages of species representing at least 
three distinct faunae. The Qrst, that of the great plains surrounding the moun- 
tains, is marked by a great development of wingless or Imperfectly winged forms, 
probably largely Invaders from the south where we may suppose thnt the arid 
deserts first made their appearance and where this characteristic feature Is more 
In evidence among the beetles. Good examples may be found among the Meloldae, 
Tenebrlonldae and eplgaeal Rhyncbophora. Occasionally these forms leave their 
natural haunts and extend for long distances up the river valleys. Thus Eleodes 
may sometimes be met with at altitudes exceeding ten thousand feet. As we enter 
the timbered country on the higher foot-hilln and lower mountain aides, we en- 
counter a fauna which while not unmixed with species that have come up from 
the plains, shows a strong affinity to the life about our Great Lakes. Higher etlll 
— that is to say from about eight thousand to nine thousand feet, according to 
the exposure, presence or absence of near-by snow-fields and so on — we meet with 
many species of genera still more boreal in habits. We may mention Nebria with 
its many species, usually taken along the coldest mountain streams, the flattened 
Bembidia, and the large Aphodii. Above timber line the peaks sustain a few 
beetles which seem to be of arctic origin, left, probably, by the retreating Ice- 
sheets of the Glacial period. 

"I cannot agree with Prof. Cockeretl* who claims that the Glacial epoch would, 
for the time l>elng result in the almost complete extermination of the Insect fauna 
of Colorado and the ad.^acent table'lands. He assumes that the arid region 
'where not actually glaciated would be a frozen desert,' something which I think 
Is not Indicated by such geological evidence as we poeaess. The glaclatlon of Col- 
orado was apparently not particularly extensive. Neither does it seem likely that 
the western Ice-sheet went so tar south aa San Diego; at any rate the indications 
aeem to show that along the highlands of Southern California only the loftier 
mountains were glaciated at all. Today great glaciers exist In the immediate 
vicinity of well-wooded districts rich In animal life. The same phenomenon may 
have occurred during ancient times." 

1893. pp. 232-233. 

*' 1. That the fauna of southern Alaska Is less closely related to our alpine, 
northern Inland, or north-east coast faunae than Is that of the Stikine Canon or of 
Glenora. 



* TnUlMctfoni of tlie American Enlomolocical SoclBly. Vol. XX. p. 319. 



lyCoogle 



J82 MICHIGAN SURVEY, 1908. 

2. Tha,t the Stlkloe Canon fauna la more doeety allied to that of tbe Nortb and 
East tban 1b that of the coast, and about tbe same aa la that of Glenora. 

3. That the chief relations of all three are In the direction of Lake Superior: 
with larger lists this affinity might turn to the Rocky Mountains, especially in 
the case of Oleaora. 

Kegardlng the afflnltles of the faunae of the Coast, the Stlhlne Canon and 
Olenora among themselves we And: 

4. That one-sixth of the Coast species extend up to the Canon while only one- 
thirtieth reach Qlenora. 

6. That the last-named fauna is much more closely allied to tbat of the Canon 
that to that of the Coast; nearly one-fourth of the Glenora species are found also 
at the Canon while only about one-eleventh extend to the Coast. 

6, That the fauna of Qlenora Is apparently Ie«e related to that of the Coast 
than to that of the interior or the East. 

"Reference to tbe accompanying maps will throw some ll|:ht on the problems 
here suggested. Olenora Is on tbe Inside of the great Coast Ranges while the 
Little Canon is regarded by Dr. Dawson as marking the bead of the old salt-water 
Inlet that has been silted up. This would account for much In tbe distribution ot 
the species In question. The climate of the country above tbe Canon Is also much 
dryer and with greater extremes of heat and cold than on tbe Coast. Aside from 
the Influence of the barrier of the Coast Mountains interposed between faunae 
whl<4i mi^t tend to Intermingle, tbe change of plants consequent upon ditference 
in climate on opposite sides must also have Its effect on the Insects dependent 
on vegetation tor food." pp. 232 233. 

190G. p. '46. 

"My proposed explanation, correlating tbe briefly outlined geological history with 
tbe facts offered as to the distribution of the Insects [shore Insects of the Great 
Basin], may be summarized as. follows: 

1. The shore beetles under consideration are confined to the Great Basin or its 
immediate borders, and have, In general, no allies in other districts from which 
they could have been recently developed. This In itself is strong presumptive 
evidence that they are endemic, not Immigrants. 

2. Within the Basin, recent conditions are such that the present distribution 
cannot possibly be a matter of modern origin. The small lakes now remaining 
in the Basin are separated by great tracts of arid desert, impassable to beetles 
depending on a moist soil for their development and food supply. The nature of 
these Insects is such that they cannot be carried long distances, as eggs or larvae, 
on tbe feet of birds or other animals. 

3. Ancient conditions, as shown by the geological history through the Pleisto- 
cene, were favorable to tbe diffusion of shore-loving insects through the Basin, 
because of tbe much greater extension of the lakes In those times. 

4. The Insect most thoroughly studied, Cicindela echo, is entirely cmiflned, in 
Its present range, to the neighborhood of lakes, froni which their size and the 
presence of nearby sprlnga, may be presumed to have lasted in some form from 
a remote i>eri<id — even through times of severe drought. Other littoral forms 
follow the same general law, though some of them are less sensitive to local con- 
ditions, 

"Prom these facts, I think we can come to but one conclusion — the heetlee under 
consideration are types that have inhabited the Basin during the Pleistocene times 
vhen the shores oi the great lakes stretched over hundreds of miles of what are 
now desert sauds. As the lakes shrunk during times of drought, the Insects fol- 
lowed the retreating beaches. Those which attached themselves to bodies of 
sufficient size or permanence were able to suHtain their specific existence, while 
auch as were dwelling on the edRea of pools of a transient nature were extermin- 
ated altogether. Thus we have tbe phenomenon of discontinuous distribution, 
presented not by one species alone but by an entire assemblage." p. 46. Cf. 
Wlckham, 1904. 

2. Comments on the Preceding Qeneralizations and on the Literature 
(if Geographic Distribution, The American authors who have given 
special attestion to tbe study of the geographic distribution of onr 
beetle fauna are few iu number, but they are very repreaentatiTe men. 
First and foremost is l>r. J. L. LeConte, the most remarkable and 



ECOLOGY OF ISLE ROYALE. 183 

"exceptional" of American entonio)oj;ist». A man wlio, had he devoted 
himBelf to subjects of more general iaterest than insects or to more 
general problema would, in all pi-obability, bave been generally re- 
cognized as one of the greatest of American naturalists. Other students 
who have devoted much attention to distribution, although none have 
given as much attention to the general principles of the problem as did Le- 
Conte, are: Schwarz, Hubbard, Hamilton. Wickham and Cockerell. 
Then there are several authors of local lists which must furnish the 
basis for comparisons, but only in a few coses do the authors of these 
local lists attempt to discuss the general charactcrifitics of their fauna ■ 
or compare them with those of other localities. This is certainly an 
unfortunate omission, particularly so as. in general, the authors of 
Buch lists should be the most competent to discuss the main features 
of their fauna. Of the various local lists, two are to be particularly 
commended for the ecological notes which they contain: those by 
t^chwarz in I.'lkes Washington list, and those by Hamilton, in the Pitts- 
burg list. It is through the ecological influences upon distribution that we 
must expect the greatest advances in the future study of distribution. 
In this connection there should be mentioned the studies by Webster 
on the routes of dispersion of certain species, particularly those of 
economic importance. A very useful bibliograph,v of local lists of beetles 
has been published by Hamilton and Henshaw ('9l-'92). and still other 
recent local lists will be found in the bibliography accompanying this 
l>ai"er, although no attempt at completeness is made. 

liimited time has prevented a detailed discussion of the quotations 
as ori^nally intended, but in their pi-esent form they are much more 
iiccessible than when scattered. 

V. The Present Centers of Dispcrml of the Beetle Fauna. 

The general characteristics of the Isle Eoyale beetle fauna can onlv 
be appreciated through a comparison with other areas, particularly 
with those of boreal regions and the remainder of the North American 
continent. Only the major features can here be outlined. It has been 
thought desirable to consider the subjects from the standpoint of centers 
of dispersal, rather than from the current taxonomic standpoint because 
of the emphasis thus put upon the genetic side of distribution and its 
ecological relations. 

In a former paper, (Biol. Bull., 1902, 9, p. 122) the writer listed 
certain criteria which may be used to determine biotic centers of dis- 
persal and centers of origin. As is well known, centers of origin and 
centers of dispersal do not necessarily coincide, although all established 
centers of origin must be centers of dispersal. Centers of origin are 
very often difficult or impossible to determine with the present state 
of knowledge; and many are likely to reniain so indefinitely. Then 
there is the possibility-, or even probability, that some forms have origi- 
nated at more than one place, and independently. This certainly com- 
plicates the subject of origins, increases the im|>ortance of determin- 
ing them, and means that this method must be repeated in such cases, 
but not that such determinations are impossible. Centers of origin, 
either single or multiple, at once become centers of dispersal, and 
by means of disi>ersal new centers become established so that there 



184 . MICHIGAN SURVEY. 1908. 

may lie numerous centei-s of dispei-sal in wide ranjjfing fonns. It ^otild 
also he again stated tliat centers of diiii>erfial while not nece»uiril.v 
centers of origin, are likely to heconte aucli with age, jKirficularty if 
favored by diverse environmental conditions. 

It is desirahle to nnderstand clearly what in meant by criteria. As 
understood hy the writer, they indicate the kinds or convenient classes 
of evidence to which we may tura for suggestions and proof as to the 
origin and dispersal of organisms. Their value is largely relative, so 
that they vai"y much in value, and in their application to various groups. 
In some cases a criterion may have great weight, while in another 
taxonoinic or ecologic group it may have no value or so little as to be 
merely suggestive. Each case must be tested oti its own merits. The 
main advantage of criteria is the definite form in which they present 
the problems and in the deflniteness which it gives to such inquiries 
as to origin. The number of criteria needs to he gi-eatly increased 
by the formulation of those restricted to groups of peculiar taxonomic 
or e^oloijic clianuter. It should l»e clearly emphasized that it is the 
convergence of evidence from many criteria which must be the final 
test in the determination of origins rather than the dependence upon 
any supposedly absolute criterion. 

The development of criteria has l>eeu largely along taxonomic lines, 
because taxonomy has been based largely upon structural characters 
nitlier than upon the convergence of all kinds of nflinities and evidence. 
For this reason ecological criteria have been lai-gely overlooked. With 
their increase in number, certain origins and dispersals may be estab- 
lished which otherwise could not I>e determined. 

It should be understood that the breeding range only is of fundamen- 
t«i value in the use of criteria, in the determination of origins and the 
centers of dJspei-sal. Of course only natural dispersal is considered 
when criteria and natural centei's are involved. Dispersal as influenced 
by man has peculiarities of its own which have not yet befen carefully 
formulated. Sitei'ies introduced hy man may thus secui-e many new 
centers of disiiei-sal. 

Aside from historical and paleontologiciil evidence the following 
criteria may he listed as those which will probably be of value ia the 
study of beetles. They have also furnished the basis for the determina- 
tion of centers of dispersal and origin of the North American beetle 
fauna. 

1. Location of great or maximum taxonomic differentiation of a type 
or types. 

2. Location of synthetic, primitive or closely allied taxonomic forms 
or groups possessing convergent affinities. 

'i. Location of maxiinnm size of taxonomic forms or groups. 

4. Continuity and convergence of lines of dispersal. 

5. Dii-ection indicated by seasonal api»earanoe; vernal suggesting 
boreal or montane origin, and aestival as austral or lowland derivation. 

0. Direction indicated by continuity and directness of individual 
variations or modifications along highways of dispersal. 

7. Lo<ation where the succession of beetle associations or societies 
reaches the relativt; e<)uilibriuni of a climax association or formation. 



«ibyGOOgIC 



ECOLOGY OF ISLE ROYALE. les 

f*. Tiocation of dominam* and great abundance of individnals. 

9. Direction indicated by biogeographioal or ecological aflBnitieH. 

10. Location or least dependence upon a i-estricted habitat, except 
huinid t.vpCK in ai-id repioiis. and analogous cases, 

11. Location (when both a center of origin and dispersal) of maxi- 
mum ecological differentiation in habits, habitiits, food, etc. "Adai>- 
tivc radiation,'' in part, of Osborn. 

By various combinations ninny additional criteria may be produced. 
By sorting into grouiw most of the above criteria will readily fall into 
either a taxononiic or ecologic class. Biit it will readily be seen that 
no sharp distinction can be drawn lietween the two groups; and fur- 
ther, no particular advantage is gained by such a classiflcation. 

The neccBsarily condensed character of such formulations iiuikes 
fnrther expansion and discussion desirable, but certain criteria are 
so well known and easily understood that their discussion is not nec- 
essary iis in the case of No's. 1. •_' and i; the remainder will be briefly 
t-onsidcTed. 

;{. Maximum size. Tliis should be expected to apply to the lai^er 
taxonomic units as well as to the smaller ones. In certain families, 
genera, etc., there can be no question but that thia criterion has great 
value, althougli it might not apply to allied groups. The broader out- 
lines of the relationship must be borne in mind and should not out- 
veigli exceptional cases. This relation of large size and centers of 
<irigiu seems to be supjiorted in jwirt, by Murray's i7l), pp. T-Sj pri- 
mary strains of Iteetle descent. Two of the thi-ee strains, the Indo- 
African and the Brazilian centers, contain the largest beetles. But 
this entire subject needs critical study before its value and limitations 
can be fully understood. 

5, Seasonal distribution. .Uthough familiar with this criterion, 
it was, by an ovei-sight, omitted from my former list of criteria. The 
northern affinities of the venial flora have long been known. My atten- 
tion to this ovei-sight wnw called by my friend, Mr. A. B. Y\'olcott. 
Recently Ulke ('02, p. 3) formulated this, in part, for beetles. But it 
flhould i)erhaps i>e extended to include montane forma also, as the ver- 
nal fauna of the mountains may be ex|>ected to extend their breeding 
range downward, whei-e they will appear as vernal forms at lower alti- 
tudes. At the same time the fauna at lower altitudes might tend to 
spread up the mountains where they would occur at the height of the 
summer season. I do not know that this subject has been investigated. 

The late fall feeding habits and the lack of ability to resist low 
temperature on the part of certain si>pcies which are extending their 
range, may be indicative as to their direction of origin. Many plant 
feeding insects, acclimated to northern localities, tend to cease feed- 
ing some time before the fall frosts and are thus lietter able to resist 
low temperatui-es (cf. Ibichmetjew, 'iM. Zeit. wiss, Z(miI.. 4f», p. (JOttj 
than those which feed late and are well fed. Chittenden I'lll. \>. 74) lias 
iwi^fDized this general tendency, but has not correlated it with Jlacli- 
net Jew's results. 

It is highly ])i-oliabIe that there ai-e miuiy other seasonal plieno- 
mena which indicate, in a general but more or less dcflnite nianiier. (lie 
direction of origin. r 



186 MICHIGAN SUHVBY, 1808. 

6. Continuity and dirertne^ta of individual vamtions. The contiDU- 
ity and directness or definiteness of individual variations along routes 
of dispersal may give very definite information as to the direction of 
origin. This in perhaps not of universal application but parries much 
weight nnder certain conditions. For example, continuity of variations, 
as dwarfing or iiioreasing size, have a certain definiteness which clearly 
points in a limited numlter of directions, when correlated with highways 
of dispersal (cf. Horn '72, p. 383), This is particularly so when a route 
is of a restricted character, as a drainage line, or a vaJley. If these 
variations were entirely promiscuous along lines of dispersal, there could 
be no idea of direction ; hut by taking into consideration the entire range, 
as ooe is pterfectly justified in doing, continuity and directness clearly 
point in a given direction. Jt is mainlywhen the animals along a route 
are uniform or promiscuous tliat direction cannot be determined by tlie 
character of the variation. This criterion, as restated, like most other 
criteria should not be used independentlv. Compare Tower '06, pp. 
12-13. 

7. Geographical centers and climax asaoeiations. To apply this 
ecological criterion it is necessary to understand the principles which 
underlie the succession of beetle associations or societies. By a beetle 
association is meant that combination of beetles which occur asso- 
ciated in the same breeding habitat. As the environmental conditions 
upon which beetles depend chnnge, the beetles also change and thus a 
succession is produced. The same general principle holds for a beetle 
association. Thus as the conditions change the association also 
changes and a surrfsnioii of beetle associations is produced. When, 
however, a relatively complete adjustment or equilibrium is acquired, 
and changes become slight, a self perpetuating or climax association or 
formation has be<;ome established. Areas occupied by formations, 
through their abundance and dominance, become centers of disp^^al, 
although they art probably more productive or originative, at an in- 
termediate stage, before the dominance of the climax associaticm ia 
fully established.' 

Meml)ers, therefoi-e, of such climax associations may be expected to 
point in the direction of such centers as include their associated species. 
If such a i-elation is valid, the various characteristics of climax associa- 
tions will aid in the determination or location of centers of origin and 
dispersal. Such criteria may have more value in determining centers 
of dispersal than those of origin. This criterion will probably apply to 
secondare' societies, but with attenuated force. 

S. Dominance. This is a fundamental criterion in the determination 
of ecological associations. The tendency for certain associated speciea 
to obtain exclusive possession of any given area implies the abundance 
of individuals and their dominance. This idea is prevalent and funda- 
mental in ecologic studies. This is also a relative term, and like all 
other criteria, has its limitations. Dominance in a desert must In 
general have a diff'ereut meaning than in a humid area. 

9. Bicgeographical or ecological affinities. In its broader applica- 
tion tins criterion is applicable to general biotic relations and to large 
areas. It is one of the oldest criteria used in the determination of faunal 
and floral affinities. In some i-espects it is closely related to Xo. 7. This 



ECOLOGY OP ISLE ROYALE. 187 

criterion can be illuBtrated by reference to tlie Ajax Butterfly (/. ajax). 
The sole food plant of the Ajax larva ia the Pawpaw, a shrub clearly 
of tropical ori^n. The allies of Ajax are also tropical ; thus the asso- 
ciated biogeographic (plant and animal) affinities clearly point to the 
tropics. It is this combination of certain ecological relations or associa- 
tions which show biogeographic afflnitieei. Thus food and other habits 
and instincts become of special value. Here also belongs a large class 
of ecological relations, particularly those related to the succession of 
insect associations. The great dependpnce of insects, as a class, upon 
vegetation necessitates a close relation between the succession of plant 
associations or societies and certain species of beetles. If certain mem- 
bers of a biotic (plant and animals) association or society have certain 
geographic afBnities, others associated with them are likely to have 
similar affinities (cf. Horn '72, p. 384). This phase ia not identical with 
the idea (»f fauna! or floral affinities, it includes them and the relation of 
biotic aiaociation, particularly us members of a climax association or 
formation, when geographic affinities are to lie determined. 

This criterion is of very extensive application. It is really a group 
of criteria and not a single one, because associations include not only 
organisms in close proximity, but also commensals, symbiots, parasites, 
etc. Seasonal phenomena might well be included within this class. 

iO. Least dependence upon a restricted habitat. From the stand- 
point of animal associations this is a criterion which may be ex- 
pected to have a rather extensive application. Its most conspicious 
application is to that of dispersal. Outlying colonies tend to have 
a, limited or restricted range. At the same time such colonies are 
particularly liable to become extinct, as they are usually near the 
limit of favorable conditions. Uften beetles in such a location are 
dependent upon a single food plant, etc. This is true of the "boreal 
islands" in swamps within the glaciated portion of the continent. For 
example, members of the tamarack bog association, toward their southern 
limit, have very restricted or local range; but to the north, the bog 
forest conditions, as it were, spread from the bogs proper and become 
of extensive geographic range, as the water beetles invade the damp 
mosses (Wickham, '97, p. 126). The outlying tropical "islands" border- 
ing the Rio Grande, as described by Schwarz ('01) and Wickham ('97a), 
apparently illustrate the same phenomena. These restricted, attenuated, 
or isolated colonies, dependent upon special conditions, are clearly 
indicative that they are pioneers or relicts, which point toward 
the region where their range is spread out and becomes of geographic 
extent. But it does not follow that every isolated habitat has such 
■a. meaning. In general, a study of succession in the region will deter- 
mine to which class the colony belongs, pioneer or relict. 

There is an exception to this criterion in the case of semi-aqnatic 
or aquatic animals in an arid region. In such regions the springs, 
streams, and water basins are so limited in extent that their isolation 
is conspicuous; and yet these conditions may be very fai'orable to the 
formation, or at least preservation, of new variations and species. 
Thus an arid region may be particularly favorable, in a sense, to the 
formation of varieties and species, although individuals may not l)e 
numerous. In such cases the amount and kind of differentiation mitU- 



188 M[CHIGAN SURVEY. 1908. 

in the aren slionld carry more weiplit than abundnnoe of indmduals. 
But by the proper «-ori*elation of criteria, such cases will not be con- 
fusing. This sort of differentiation is well shown among beetles by 
Wickham, '04. '(15. 

This criterion evidently does not apply, at least in part, to the fannn 
now found in glaciated North America. This is made probable through 
origin elsewhere and a later expansion in the glaciated area as the Ice 
Age declined. 

11, Ecological diffei-entiation. Ecological and taxonomic diffeivntia- 
tion need separate recognition, althougii they are frequently not di:<- 
tinct because of their intimate genetic relaiions. With great taxonomic 
diversity, within a gronii. there is almost certain to be ecolojjic diver- 
sity; but generally much letw attention is given to the ecological diver- 
sity. Compare Xo. 1. 

The following outline of the centers of beetle dispersion must be 
considered provi.sional and suggestive, as it is a aubjei't which has 
i-eceived but little attention from the standpoint here presented. The 
preceding criteria, of taxonomic and ecolt^ic natui-e, have been given 
much emphasis in locating the present centers of dis])ers!il. Through- 
out this section references are given to significant pai)ers, hut this 
does not imply that the views here presented are ap]>roied by the 
authors to whom i-eferenoe is made. Tliese referencew also apply to 
the quotations already given in detail. 

1. The American Tropical Venter. From Panama northwartl to the . 
Mexican pUvteau is the main body of the tropical center. Xarn»«- 
elougatious extend coaKtwise ou each side of theplateau, andon theOulf 
Coast to the Rio Grande river, William Wickham '07a; Schwarz '(II: 
Toii-nsend '95, '37; Tower '06. Outlying colonies are found on the 
Pacific coast of Mexico and at the Southern extremity of Lower Cali- 
foi-nia; at the mouth of the Colorado river (Bchwarz) ; and in south- 
em Florida (I*Conte '7Sa; Kchwar?. '7S, "SS). The fauna of the \Ve«t 
Indian Archijjelago probably belongs «-ith this gi-eat coniiKwite center. 

A vast number of beetles are characteristic of this complex area. The 
Biologin Centrali-Americana devotes thirteen volumes to the descrip- 
tion of beetles from part of tliis area. \Yithin the United States the 
attenuated tropical element has Iieen most carefully studied and its 
faunal affinities detennined by Scliwar^ ("SSi and tt'iokham (!)7al. 

This tropical center is composed of several distinct units. This is 
an ancient center of origin, pi-eservation and of disjiei-sai. Tlie routes 
of dispersal into the T'nited States have lieen along both coasts of 
Mexico and via the West Indies. It was practically uninfluenced by the 
Ice Age. 

2. The Meriean Platcfiti ami the itonthirfHtem Dry Bencrt. This center 
includes the Mexiojin I'iateau; most of Lower California; the deserts of 
southwestern United States; the low landx of California; the Great 
Basin and the (ii-eat Tiains northward into Canada and east to the 
forests. I^Conte '.")1, TtO, 'fi(t, 'fiL'; VauDyke '01: Wickham '!)(!. 'OS, "tli, 
'Oii: Tower '0(5; Fall and Cockerel! "07, 

(Characterized by numerous desert species; winglesN Teuebrionidae 
(Hoi-n 71), and Cicindellidne. A given Iwality is characterized by a 



ECOLOGY OF ISLE ROTALE. 189 

limited number of species wliich are individually abaodant; aumerous 
local fannfle. Containa the characteristic desert fauna of Nortii America. 

lofluenced markedly by glaciation only at the extreme north, and 
poBBiblj in the Great Basin, but certainly by the great fresh water 
lakes formerly occnpyinp this basin (Wickham '04, '051. An old {pre- 
Glacial) center of origin, preservation, and center of dispereal. 

.^. The Southeaatem Humid Hardvood Fmegt Area. This area in- 
cludes much of eastern United States east of the plains (exclusive of 
southern Florida), and north to the Canadian conifers. Tlie Coastal 
Plain (and possibly the Mississippi Embayment area) may form a sub- 
<«nter through the influence of its conifers (Schwarz '90; Chittenden 
'00, '01). 

Local lists within this center: Summers '74; Schwarz '78; Ulke '02 j 
Dury '02, '06. cf. LeConte '78a. 

Characterized by the abundance of forest insects, particularly those 
infesting hardwoods; species of extensive range; few local fauna; a 
large number of species found in a given locality, often but few in- 
dividuals. 

An ancient centre of origin, preservation and dispersal. Glaciated 
on the north, and post-GlaciaJly repopulated, as was also the Coastal 
Plain and Embayment, with the elevation of the Coastal Plain. 

4. The Trantcontinental Conifer Area. This area includes the re- 
mainder of forested North America, and Includes all of the higher 
mountain ranges. Its relation to the Coastal Plain conifer belt has 
not been determined. 

Characterized by conifer feeding beetles. Cerambycids, Scolytids ; 
numerous Carabids and Staphyllnids, and thus shows a decided sutwrctia 
circumpolar affinity. Few endemic elements and local faunae. Ex- 
tensive range of species, except in the high mountains. Largely a new 
land surface through glaciation; largely repopulated at a relatively 
late date; apparently characterized by dispersal or diffusion of forms 
rather than for their origin or preser^-ation, except the mountain areas 
of the United States, which were areas of preservation and origin. 

The very different history of its eastern portion, with much more 
extensive glaciation, is to be contrasted with the moderate western gla- 
ciation. The differences in the character of the forests is also marked; 
the giant western conifers are to l>e constrasted with the smaller con- 
ifers of the Northeast, The Mackenzie Basin and the Plains mark the 
line of division between these suticenters. This division is so marked 
that it may yet necessitate a complete se\ering of the tranaeontin- 
«ntal conifer belt, (cf, Murray '70, pp. y2-3X) 

a. The Eastern Canadinn Conifer Fauna. 

This includes Canada east of the Rocky ?tIountains, north to the tree 
limit and the interior of Alaska: northeaslem United States; and 
southward on the Appalachians, 

Characterized by forms of extensive subarctic range, very few 
«ndemic elements or those peculiarly American; individuals abundant, 
variety moderate; beetles feeding on conifers, birches, and as{>ens. I>e- 
■Conte '50, '59. '7Sb. There is a posRibilily that this fauna has North 



100 MICHIGAN SURVEY. I9D3. 

£nropeaD nfBnities stronger than Asiatic ones; the i-ererse from the 
western conifer center. 

Local Hats including this fanna; Hubbard and Schwarz '78; Har- 
rington '84; Wickham '97; Hamilton "d^, 94a, '95, Klages '01. 

b. The Western Canadian Conifer Fauna. 

This ioeludes the western mountains northward and upward to the 
tree limit, eastward to the Great Plains and the eastern Conifers. 
North of the United States this area has been extensively glaciated but 
within the United States the glaciers wfre local. 

On account of this moderate glaciation in the mountains of the 
United States, this old land surface has been a center of origin, prener- 
ration and dispersal. It contains more endemic elements than the 
eastern conifer center, and more local faunae. Beetles feeding upon 
conifers and aspens are fairl.v characteristic. The Asiatic affinities of 
the fanna. are much more pronounced than those of the eastern con- 
ifer center. Part of the apparently European influence may more truly 
be considered Asiastlc — both the Western and European — having 
been derived from Asia. Elements of this fauna probably 8urvi*-ed the 
Ice Age on the Pacific Coast north of the United States, but tlie- 
humid interior has been invaded from the south, or is endemic. I^e- 
Conte '76; Wickham '96; Cockerel! '93; Hamilton '94, '94a; Fall and 
Cockerell '07; Keen '95. 

5. Alpitie and Arctic. This fauna occupies the area north of the 
tree limit, and above the tree limit on the mountains. This fauna i:* 
very imperfectly known and is limited in, variety and in the number of 
individuals. Many species are of circumpolar range in the Arctic re- 
gions. Composed of \-ery diverse elements and of diverse origin. The 
nnglaciated arctic areas are probably centers of origin as well as of dis- 
persal, as also slightly or moderately unglaciated alpine areas. The 
glaciated portions have been repopulated and show incipient endemisni 
but are mainly characterized by the extensive dispersal of species, tut i» 
apparently true of unglaciated Asiatic Siberia. Both of these centeriii 
(Alpine and arctic) have been much confused with regard to whethei- 
<bey are centers of origin or of dispersal. These types are curivntly 
stated as of boreal origin, but this is very improbable for perhaps 
the majority of the population. They may be of alpine origin on the 
western mountains with an extensive post-Glacial dispersal favored bv 
climatic conditions, and the low topographic relief of the northern land, 
areas. Schwarz '90 ; Murray '70, pp. 32-33. 

Eastern Alpine, Scudder '74 ; Bowditcb '96. 

Western Alpine. Carpenter '75 ; I^Conte '78, '79 ; Schwarz '90 ; Cock- 
erell (including Horn) '93; Wickham '03; Fall and Cockerell '07. 

TI. The Oenerat Characteristics and Affinities of the Isle Royale Fauna. 

1. Faunal Characteristics. The accompanying list of beetles col- 
lected in 1905 includes 89 species. The only previous list is that by 
Hubbard and Schwarz ('78) in which they list 123 species. A surprisinft 
feature of our 1905 collections is that of our 89 species, 66 are not 
listed by Hubbard and Schwarz, Such species are indicated by the 



ECOLOGT OF ISLE ROTALE. 191 

letter A. following the seientiflc name. • On the basis of these two lists, 
206 species are now recorded from the island.- It is not improbable 
that other species have been recorded in tbe scattered literature, but 
no effort has been made to search for them. Undoubtedly only a fair 
start has been made in the study of the beetle fauna. Careful detailed 
(.■ollecting, covering several years, would probably increase the number 
(ibout five times, or bring it up to about 1,000 or 1,100 species; that in, 
judging from other northern localities. Pcttit has recorded from 
Grimsby, Ontario 1,143 species and Harrington ('84) from Ottawa 
1.00.3 species. On the other hand it is not improbable that the present 
known 206 species give a fair sample of the dominant features of the 
l)eetle fauna. Wickham's ("J7) Bayfleld, WisconNin list contains Oftl 
species (six weeks collecting by an expert). Such statistics mean but 
little, beyond showing tbe reduction in variety toward the north when 
compared with southern localities. The two best local southern lists — 
tbe best in America — are those by Ulke for Washington, D. C, with 
2,975 species, and by Dury for the r^on about Cincinnati with 2,2£H> 
species. Two important intermediate locality lists between these north- 
em and southern ones are from the vicinity of Allegheny and Pittsburg 
by Hamilton, in which 2,153 species are listed or 2,500 as given by 
Klages ; and at Buffalo, where about 1,424 species are listed by Reinecke 
and Zescli. The variety in tieetle life is thus seen to drop off about Vi 
or more in passing from the latitude of Washington and Cincinnati to 
that of Lake Superior and the St. Lawrence valley. 

2. MisceUaneous XoUa on the FatuM, In the present list there are 
included 6 species which in the Hubbard and Scliwarz list are indicated 
as "Species found by Dr. LeConte, mostly catal<^pied in Agassis' Lake 
Superior, p. 203-239, which have not since occurred." These species 
are as follows: Carabus seratus, Calthus gngariut, Blechrua nigrinua 
(linearis Lee), Barpalua ruficoUis. Pachijta Uturata, Dtmacia proxima. 
All these and other rare species turned up in our collection. 

TjcConte and Horn describe the following three new species from 
Isle Boyale specimens in the Hubbard and Schwarz paper: Sabroceras 
magnua Lee, p. 598; Phymatodea mactilicollig Lee, p. 614 (from one 
specimen) ; Orcheatea vantia Horn, p. 020. None of these species were 
found in our collection. LeConte ('IS, p. 463) described Magdaiia 
alutacca {armicollia Say) from Colorado and Isle Boyale specimens. 

As numbered in the accompanying list of species collected during 
1005, the following are not to be found in tbe Bayfield list by Wickham ; 
Xo's. 2. 6, 7, 10, 12, 15, 16, 17, 18, 20. 21, 22, 25, 27, 30, 34, 36, 41, 
50, 52, 53, 55, 56, 59, 60, 62, 64, 73, 76, 77, 79, 81, 85, 86,-35 species. 



DigtizedbyCOOgle 



^IICHIQAN 8URYBY. 1908. 



VII. LISTS OF ISLE ROYALE BEETLES. 

1. LiaT OP SPECIES COLLBCTED IS 1905. 

Cichidelidac. 

1. Oidndela longilabria Say. A, One fipeciiiieD of the dark form 
\vaB taken from the clearing about Neiitson's resort (IV, S) on Julv 21 
(Q. 121). 

Geographic Range. Newfoundland; Ottawa, Canada: Hudeon Bay; 
Nova Scotia; Quebec; Mt. Washington (summit), N. H.; Michigan: Wis- 
consin; Nebraska; New Mexico; Colorado (10,000-12,000 ft.); Utah; 
Idalio; Montana; Albei-ta; California; Oregon; Alaska. 

Carabidm: 

2. Carahus aerratim Say. A. A single specimen was found crawling 
over and through the tults of Cladonia in the rock opening near camp 
on Siskowit Bay (V, 3) on August ."> {(3. 208). 

Geographic Range. Saskatchewan Itasin, Canada: Mt. Washington. 
N. H. ; W. Penna. ; Michigan; Indiana (A. B. Wolcott) : Chicago, III. 
(Wolcott); Kansas; Colorado; New Mexico. Hamilton '94ii, p. 35i. 

3. Caloaoma frigidum Kb.v. A. A single siiecimeu was found on July 
7 among the drift on the beach (I, 11 near Tonkin Bay (A. 7). , 

Geographic Range. Prummond's Island, Ottawa. Canada; Mt. Wash- 
ington (summit) N, H.; New York; Chicago, Illinois (Wolcott); W. 
Penna.; Michigan; Indiana; Wisconsin; New Mexico; Texas. 

4. liemiidium caritiula. Chand. A. "Very abundant July 8 on the 
sandy beach at the head of Conglomerate Bay (I, 1). Running rapidly 
over the sand and fine gravel just back of the wet strip along the shore." 
(G. 30), Oleason. 

Geographic Range, New Hampshire; Mass.; Adirondack Mts., New 
Y'ork; Port Arthur, Ontario; Saskatchewan Basin, Canada; Georgin; 
Ohio; Michigan; Indiana (Wolcott) ; Illinois; Wisconsin; Arkansas; 
Colo. (8,0(t0 ft.) ; Oregon; Brit. Columbia. Hayward, '97. p. 4G. 

5. Bembidiiim transcersale I>ej. Two specimens were taken about 
the camp at the Light-house (I, 7) on July 11 (G. 4SK 

Geographic Range. Canada; Gulf of St. Lawi-ence; Lake Superior 
region; Mich.; Wisconsin; Ne'braska; Kansas; Colo.; New Mexico; Ari- 
zona; Wyoming; Utah; Pacific Coast from So. Calif, to Alaska. 

(!. Urtmbidium grapii GyW.^nitenn Ijec. A, "On a low bare rock on 
the shore near the Lighthouse at Rock Harbor (I, 11. On July 11, early 
in the morning, with air temperature of 51° F. and surface temperature 
about the same, no specimens were seen ; but as the surface grew warmer, 
up to 95° F„ the beetles liecanie abundant. They jirobably conceHl them- 
selves in crevices in the rock when the temperature is low." (0. 46. i 
Oleason. 

Geographic Range. Greeulaud ; Hudson Bay region, Saskatchewan and 
Mackenzie Basins; Isle Royale, Michigan: White Mts., N. H. ; New York; 
southward on the mountains of the wwl to Colorado, New Mracico and 



ECOLOQY OP I8]J! HOYALE. 193 

Nevada; Alaeha; Siberia; Northern Europe. Hamilton, '94, p. 8; '94a, 
p. 351. 

7. BembUUum rariegatum Say.=patniele l^ej. "In debris cast up on 
the beach at the head of Tonkin Kay {I, 1) with B. versicolor and 
Plati/ntis." (G. 21i. Glenson. 

Geographic Rantjc. Nova Scotia; New England States; New York; 
New Jeniey; Peniia.; Maryland; Distr. Oolnmbia; Texas; Ohio; Mich- 
igan; Lake Superior region; Wisconsin; Illinois; Iowa; Missonri; 
Nebraska; Saskatchewan Basin, Manitoba; Colorado; Nevada; Calif, 
to Brit. Columbia. 

8. Bembidiiim versicolor TjCC. A. "In debris at the head of Tonkin 
l!ay (I, 1) with B. varicgutum and Platynns 4-punctatus (G, 21)." 
* i leasoQ. 

(leogrophir Rinige, General distribution in Canada and United States; 
fi-oni Auti,costi. Quebec to Florida, Texas and California and north to 
(Colorado and Manitoba; Pine, Ind. (Wolcott). 

9. Ptcroatichus coracinus Neivni. A. A specimen of this species was 
taken in the Lighthouse clearing (I, 7) on Julv 11 (G. 49) and on July 
28 (G. 179). 

Geographic Range. Ottawa, Canada; Mt. Washington (summit) N. 
H. ; Vermont; New York; New Jei-sey; W. Penoa. ; Maryland; Virginia; 
Itist. Columbia: Teun.; Ohio; Mich.; Northern IltinoiB; Iowa; Wyoming. 

10. PteroDticTms fcmoralis Kby. A. A specimen of this ground beetle 
was found under Cladotiia upon a sloping rock slioi-e (V, 2) just beyond 
the reach of the waves, on August 16 (A. 130). 

Geographic Range. Ottawa, Ontario; Saskatchewan Basin; Mass.; 
Mich.; W. Penn.; New York; Ohio (Dury) ; Colo.; New Mexico; No. 111. 
anil Ind. (Wolcott). 

il. Caiatkua grcgarius Say. A. A specimen was taken on or in leaf 
mould in a deeplv shaded balsam-spruce forest (I, 3) on July 24 (G. 140) , 
and (V, 4) on August 14 (G. 236). 

Geographic Range. Ottawa, Ontario; Quebec; Saskatchewan Basin; 
Vermont; New York; New Jersey to Florida and Texas; W. Penna.; 
Ohio; Mich.; No. Illinois (Wolcott); Wisconsin; Iowa; Kansas; Neb- 
raska; New Mexico, 

12. Cafaihus adtena Lee. A. "One was found crawling through soft 
decayed wood in the balsam-spruce forest (I, 3) on July 24 (G. 142)." 
Gleason. 

Geographic Range. Maine; Vermont; Mt. Washington, N. H. ; Mich- 
igan; Colorado; New Mexico; So. Alaska. Hamilton, '94, p. 11. 

13. Platynus 4 punctatua YieO. A. A single specimen of this 
species was found about camp at the Lighthouse (I, 7) on July 11 (Q. 
49), also in debris cast uji on the beach at the head of Tonkin Bay (I, 1) 
.where it was found alive (G, 21). 

Geographic Range. Ottawa, Ontario; Canada; Hudson Bay and 
Lake Superior regions; Mt. Washington, N. H. ; New York; W, Penna.; 
Mich,; Wisconsin; Idaho; Colorado; New Mexico; Montana; Alaska; 
Kamchatka; Siberia; Northern and Alpine Europe. Hamilton, '94, p. 11. 

14. Bl€chru8 nigrinug Mann.^/iwcuWs J^c. "In the debris under 
mats of bearherrv on the rock ridge north of the Lighthouse at Rock Har- 
bor (I, 3), (G. 64)." Gleason. 

Geographic Range. Saskatchewan Basiu, Canada; New York; New 

hyGoogIc 



194 MICHIGAN 8URVBT, 1808. 

Jersej ; Mich. ; Iowa ; Wisconsin ; Missouri ; Dakota ; Wyoming ; Colo- 
rado; New Mexico; Calif.; Brit. Oohimbia; possibly Siberia and No. 
Europe. Hamilton, '94o, p. 355. 

15. Harpalus megacephalua Lee. "In rock crevices and under debrin 
from bearberry on the jack pine ridge (1, 2) on July 13 (Q. 72)." 
Oleason. 

Geographic Range. Lake Superior; Isle Royale, MicUigan. 

HaHptidae. 

IB. Haliplua ruficollis DeG. A. "At the bottom of small pools in 
the partially drained sphagnum bog near Conglomerate Bay {I, 6) on 
July 18 (G. 116), and at the bottom of a. small stream flowing from 
a. tamarack swamp near Si^owit Bay (V, 5) on August 12 (O. 230). 
In each case the water wae shallow and the bottom composed of 
-sphagnum covered with dead leaves." Gleason. 

Geographic Range. Canada ; Hudson Bay region ; Mt. Washington ; 
Kew Hampshire; Vermont; New York; New Jersey; Mich.; W. Penna.; 
Ohio; 111. (Wolcott) ; Iowa; Colo.; New Mexico; Texas; .Wyoming; 
Kansas; Western Siberia; Europe; Tnrkestan. Hamilton, '94a, p. 355. 

I^fftiscida^. 

17. Hgdroporus triatta Pavk. A, "In the bottom of small streams 
draining a tamarack swamp (V, 5). (G. 237)." Gleason. 

Geographic Range. Ottawa, Ontario; Vermont; Mass.; Mich.; Lake 
Superior region; Hudson Bay; Colorado; British Columbia; Alaska; 
Arctic Miberia; Northern Europe to Finland. Hamilton, '94, pp. 13. 
'94a, 357. Sharp, '82, p. 472. 

18. Hydroporus modestua Aube. A. Taken at Benson Brook clear- 
ing (II, 1) on July 29 (A. 81). 

Geographic Range. Ottawa, Ontario ; Mt. Washington, N. H. ; Mass. ; 
W. Penna.; New Jersey; Dist. Columbia; "Carolina"; Wis,; Mich.; 
Ohio (I>ur>). Sharp, '82, p. 480. 

19. Ilyhiits pleuritictis Jjec. A. "In the water near the shore at 
camp on Slskowit Bay (V, 1) on August 7 (G. 213)." Gleason. 

Geographic Range. Penna.; New York; Isle Royaile, Mich.; Bayfield, 
Wis. ; Iowa ; Colorado. 

20. Agabus stridulator Sharp. A. Taken in a clearing (II, 1) on 
July 29 (A. 81). 

Geographic Range. Isle Rovale, Mich. ; Hudson Bay ; Canada. 
Sharp, '82, p. 509. 

21. Agabu8 congener Payk. A, "In the bottom of streamlets drain- 
ing a tamarack swamp (V, 5), (G. 237)." Gleason. 

Geographic Range. Greenland; Labrador; Hudson Bay; White 
Mountains N. H. ; Mass.; Penna.; Mich.; Missouri; Arctic and Western 
Siberia; Centnii and Northern Europe. Hamilton, '94a, p. 358. Sharp, 
'S3, p. 513. 

22. Scutoptcrtit hornii Cr. A. "In small pools in the tamarack and 
nrbor vitae swamp (1, 4) on July 28. These pools were under fallen 
logs and at the bases of trees; seldom more than 1.5 dm. in depth and 
with a bottom of sphagnum and vegetable debris (G. 181, 182)." 
<ilea9on. ,- , 

i.Cooglc ■ 



ECOI^OOY OF ISLB ROYALE. 19S 

Geographic Range, Canada ; lele Royale, Michigan. 

23. Rhanius iinotatus Horr. A. Two were found in rock pools on 
the beflch at the entrance to Tonkin Bay (I, 1) on Julj 13 (G. 73, 7i) 
nnd at Hcovill Point (IV, 1) on July 19 (G. 130). The beetles nsually 
remained on the bottom except when they came to the surface for air. 

Oeographic Range. Labrador; Ottawa, Canada; Hudson Bay region; 
Brit. Colambia; Mt. Washington, X. H.; New York; Xew Jersey; Mich.; 
"Wisconsin; Kansas; Nebraska; Colorado; Sew Mexico; So. Arizona', 
T^tah; Nevada; Calif.; Lower Calif.; Mexico; Guatemala. Sharp, '82, 
p. 614. 

Gprinidae. 

24. Ggrinm m'mutm Fab. A. "In sheltered coves of Siskowit 
T^ke (V, 6) on August 9, where the water was quiet. Most numeroas 
near the shore under the overhanging alders where they congregated in 
large flocks (G. 219)." Gleason. 

Geographic Range. Labrador; Canada; Hudson Bay region; Sad- 
katchewan basin (Hvans '03) ; Vermont; W. Penna.; Michigan; Wiscon- 
Hin; Washinffton; Or^on; Siberia; Central and Northern Europe. 
Hamilton, '94a, p. 360. 

25. G>/rinu8 pidpea Aube. A. In lai^e numbers near the shore of 
t^iskowit lake (V, 6) with the preceding species (fi. 219). 

Geographic Range. Labrador to Brit. Columbia; Vermont; Michi- 
gan; Idaho; Oregon; So. Alaska. Hamilton '94, p. 14, 

Siaphylinidae. 

26. Oyrophaena species. "Several specimens (G. 229) were taken 
from a ahelf fungus, Pleurotiia ogtreatua, on August 11 (V, 4)." Gleason. 

27. <^uediu8 fulgidus Fab. A. Two were taken from leaf mould or 
under dcfcared bark in the maple forest (III, '04) on August 21 (A. 
142). 

Geographic Range. Greenland to Alaska; south to No. Georgia and 
I^a. and Central' Calif. ; Peru; Mich.; West Siberia; Europe; Asia Minor; 
No. India; Java; Tasmania; Australia; New Zealand. Hamilton, '94, 
p. 18, '94a, p. 366. 

28. Philotithua politiia hiua.^aencui Kossi. A. Hamilton, '94a, p. 
19. One specimen was taken about camp at the Lighthouse (I, 7) on 
July 7 (G. 26). 

Geographic Range. Isle Royale, Mich.; Nova Scotia; Hndson Bay 
region; British Columbia; New York; Mass.; Penn.; New Jersey; La.; 
Ohio; Illinois (Wolcott) ; Wisconsin; Iowa; Kansas; Colorado; Neiv 
Mexico; Queen Charlotte Island; Alaska; Siberia; Amur region; 
Europe. 

29. Lathobium simplex I^ec. A. One specimen {A. 24) was taken 
July 17 on a jack pine ridge {I, 5). 

Geographic Range. Canada; Mass.; New York; Michigan; Wiscon- 
sin, Am. Knt. Soc, '80, p. 17G. 

30. Tachinua memnmiiua Grav. A. One beetle was found under the 
bark in the hardwoods along the Desor trail (III, '04) on August 24 
(A. 149). 



DigtizBdbyCOOgle 



196 MICHIGAN SURVEY, 1908. 

Qeoffraphic Range. Dist. of Columbia; W, Penna.; Ohio (Dury) ; 
WlBConsin; Michigan; III. (Wolcott). 

31. Bolctobius cincticollis 8aj. "In fresh plants of the bracket 
mnshrooiD Pleurotua sp. growing in the baisam-spruce forest (V, 1) on 
August 11." tileason. One speciineu (G. 229). 

Qeographic Range. Canada; New York; W. Feooa. ; New Jersey; 
Dist, of Columbia; Ohio; Wiacoasin; Iowa; Mich, to Brit. Columbia; 
Calif, and Arizona; cf. Hamilton, '94, p. 21, Alaska. 

CoccincUidae. 

32. Hippod^mia ISpunctata L. A. Taken about camp at Kocfc 
Harbor (1. 7) on July li (G, 98). 

Qeographic Range. "All America north of Mexico ;" WeBt Indies ; 
Alaska; throughout Europe and Central Asia; Siberia. Hamilton, '94a. 
p. 378. 

33. Anatis 15punctata 0\\y. ^=ocellata L. A. Found among drift 
cast up on the beach at the head of Tonkin Bay (1, 1) on July 6 (G. 21). 

Qeographic Range., Ottawa, Saskatchewan tKisin, Canada; Nova 
Scotia; New York; New Jersey; West Indies; W. Penna.; Ohio; 
Illinois; Mich.; Wisconsin; Iowa; Siberia; Europe. Hamilton, '94a, p. 
379. 

Erotylidae. 

34. Tritoma macro Lee. A. "One specimen (G. 229) found August 
11 in a shelf fnngus Pleurotua ostreatus (V, 4)." Oleason. 

Qeographic Range. Maine; Michigan; Illinois; W. Penn. 

35. Tritoma thoracica Say. A, From fre^ specimens of Pleurotua 
growing in the balsam-spruce forest (V, 4) on August 11 (G. 229). 

Geographic Range. Hudson Bay region; Saskatchewan basin; Ot- 
tawa, Canada; Vermont; New Y'ork; New Jersey; Va. ; Georgia; 
Florida; Texas; W. Penna.; Ohio; Ulinois; Mich.; Wis.; Iowa; Colo.; 
New Mexico; Washington. 

DascylUdae. 

36. Macropogon ruftpcs Horn. A. One specimen was found upon the 
beach of Lake Superior (I, 1) on July 12 {G. 60). 

Geographic Range. Illiuois; Isle Koyale, Mich.; White Mts., N. 11.; 
Horn, Amer. Ent. Soc, '80, p. 80. 

Elateridae. 

37. Adclocera brcvicornis Lee. A. One taken about camp at the 
Lighthouse (I, 7) on July 18 (G. 117). 

Geographic Range. Ottawa, Canada; Micb.; Wisconsin; Lake Su- 
Ijerior. 

38. Elater hepaticue Mels. A. Two taken about the camps both 
at the Lighthouse (I, 7) on July 13 (G, 86), and at Siskowit Bay iV, 
3) on Aug. 7 (G. 212). 

Geographic Range. Canada; Vermont; W. Penna.; New Jersey; 
Ohio; "Western States;" Wisconsin; Michigan. 

39. Elater apicatus Say. A. One taken at the camp on Siskowit 
Bay (V, 3) on .-Vugust 3 (G, 193). CoC^jIc 

' '' o 



ECOLOGY OF ISLE ROYALE. 197 

Qeofjraphic Raiiffe. Hankatrhewnn baoin ; Ottawa, Canada ; New 
Hampshire; Vermont; Xew York; Mich.; Wis.; Duluth, Minn. fWoI- 
cott) ; Colo.; Arizona; Kew Mexico; Idaho; Wafili.; Oregon; Calif.; 
"Xortbem U, 8. generally." 

40. Affriotes limosua Leo. Taken on flowera of the Cow Parsnip 
(Heraclcum latiatum) in the cleai-inR at the Lighthouse {I, 7} on 
.Inly 17 (G. 1031 and on July 23 (G. 1.161. Five »|>ecioiens. 

Qeoffntphic Range. Newfoundland; Mt, Washington (sammit), N. 
n.; Ottawa, Canada; Lake Superior; Sai^atchewaD basin; Michigan; 
Wisconsin. 

41. MelanotuH para^oxvfi )leli>h. A. One taken about the ramp 
at the Lightboutie (I, 7) on July 11 (G. 49). and near Lake Desor 
{VII, '04) on August 21 (A. 139). 

Geographic Range. Isle Royale, Mich. ; Colorado ; New Mexico 
{Snow). 

42. Corpmhites medianug Germ. One taken on the beach south of 
Tonkin Bay (I, 1) on July 10 (G. 41), "crawling over the sand in a 
shaded place near a rock cliff." Gleason. 

Geographic Range. Ottawa, Canada: Mt. Washington (summit), N. 
H. ; New York; W. Penn.; Michigan; Wisconsin. 

43. Corymiitcs acripcnnis Kby. One taken at Scovill Point {IV, 
1) on July 19 (G. 130). 

Geographic Range. Ottawa. Canada; Nova Scotia; Maine; Mt. 
Washington, N, H.; New York; Mich.; Wis.; Colo.; New Mexico; 
Idaho; Oregon; Brit. Columbia. 

44. Corymhites aratus I*c. On July 19 one was taken at Tobin 
Harbor (A. 29). 

Geographic Range. Canada; Lake Superior; Michigan; No. Wis- 
consin. 

Bupreatidac. 

45. Dicera prolongata Ijcc. A. Two were taken alMmt caniji at 
the Light-house (I, 7) on July 10 (G. 45) and on July 15 (G. 86). 

Geographic Range. Saskatchewan basin ; Ottawa, Canada ; New 
Hampshire; Mass.; New Jersey; Mich.; Wisconsin; Nebraska; Kansas; 
Colo.; New Mexico; Idaho. 

46. Dicera tenebroaa Kby. Taken about the campn at the Light- 
house (I, 7) on July 25 (G. l.">3), and at SisOtonit Bay {V, 3) on 
August 7 (G. 212), and 15 (G. 239). 

Geographic Range. Ottawa, Canada; Lake Sujwrior; Mt. Washing- 
ton, N. H.; Mass.; Mich.; Duluth, Minn. {Wolcott) ; Wisconsin; 
Colorado, 

47. Buprestis macuUventris Say. A. This was the most abundant 
species of the family, and was very abundant about the camp on ^is- 
kowit Bay (V, 3} during August; others were taken at the Light-bouse 
clearing (I, 7) during July (G. 86, 117, 179, 195, 212, 222, 231), (A. 
152). 

Food plants. Beetles have been found on balsam and spruce, and 
emerging from pine timber, (Felt, 1906, p. 674.) 

Geographic Range. Newfoundland; Ottawa, Canada; Lake Superior 
region; Mt. Washington {summit), N. II.; Yenaont; Mass.; New York; 



198 MICHIGAN SURVBT, ISDS. 

Petina.; Mich.; Wieconfiin; Xebraeku; Kaneas; Colo.; Xew Mexico; 
Utah. (WaBfaington; Oregon, cf. Bethune, '76, p. 65). 

48. Buprestia faaciata Fab. (aad vArietiee). Like ttie preceding 
species, this was also taken m large numbers; at the Light-house (1, 7) 
during July and at camp on Siskowit Bay (V, 3) during August. (G. 
117, 133. 153, 166, 195, 212, 231). Thi» is a large metallic green species 
which shows considerable variation in the amount of the light-colored 
spots on the elytra. In some Isle Boyale specimens the spots are well 
developed, in others completely tacking. The var. langii is credited to 
Isle Royale in the Hubbard and Schwarz list (78). This is a western 
and northwestern variety, Alaska, Brit. Columbia and western moun- 
tains. 

Food plants. Found on poplars, and the larva-bores in maple. (Felt, 
'06, p. 459.) 

Geographic Range. Ottawa, Canada; Nova Scotia; Xortheastem U. 
S. generally; W. Penna.; Ohio (Dury) ; Michigan; Wisconsin; Colo- 
rado. 

49. Bupreatis striata Fab. A. Taken on the open rock ridge north 
of the Light-house (I, 2) on July 13 (G. 68). One specimen. 

Food plants. Occurs on pine and spruce, the buds of which the 
beetles are said to eat; may also feed upon dead wood. (Felt, '06, p. 
655). 

Geographic Range, Ottawa, Canada ; New York ; Mass. ; New Jersey ; 
Penna.; Ohio (Dury) ; Michigan: Wisconsin. 

50. Melanophila aceuminata TteG.-longipeg Bay. A. Two speci- 
mens were taken at the Light-bouse (1, 7) on July 7 (G. 26) and on Jolv 
11 (G. 49). 

Geographic Range. Canada; Hudson Bay south to Virginia, and 
Kentucky ; W. Penna. ; Mich. ; Wisconsin ; Colo. ; New Mexico ; So, 
Calif,; Brit. Columbia; Alaska; Knrope; China. 

51. Mclanophila driinimoitdi Khy.^guttulata Gebl. A. Taken about 
the camps at the Light-house (I, 7) during July (G. 98), and on Siskowit 
Bay (V, 3) during August (G. 212, 231, 239). Five specimens. 

Food plant. Found on spruce logs. ( Blanchni-d, Ent. Ainer., 5, p, 30). 

Geographic Range. Maine to Alaska (Yukon) ; Mt. Washington 
(summit), N. H.; Mich.; Wisconsin; Idaho; Colo.; New Mexico; Utah; 
Calif.; Washington; Oregon; Alaska; Siberia, Hamilton, '94, p. 29. 
'94a, 391. 

52. Clirysobothris trtticrvia Kby. Found at the Lighthouse (I, 7) 
during Julv (G. 106) and verv abundant at the camp on Siskowit Bav 
(V, 3) during August (G. 212, 222,231, 239). 

Food plant. Found on spruce l(^, Blanchard, Ent, Amer., 5, p. 31. 

Geographic Range. Ijike WinHi|»eg; Alberta; Hudson Bay region; 
Ottawa, Ontario; N. H.; W. Penna.; North Cai"olina; Mich.; Colo.; New 
Mexico; Washington; Oregon. Hamilton, '94, p. 29, 

53. Agrihts aciitipf-nniit Mann. A. One specimen from the clearing 
at the Light-house (I. 7) on July 26 (G. 166). 

Food plant. Found on t)ak. lllanchard. Ent. Amer., 5, p, 33. 

Geographic Range. ''Mass. to Kansas, Floi'ida and Texas" ; W. Penna, ; 
Ohio (Durv); Mich.; Glendon I'ark, III. (Wolcott). Horn, Trans. Ani. 
Ent. Soc., 18, p. 309. 



,dbyGOO<^IC 



ECOLOGY OF ISLB ROYALE. 199 

Lampyridae. 

54. Podahrius diadema Fab. A. Found about the camp at tbe Light- 
honse (I, 7) on July 23 ((!. 1X1), and among beach drift at the head of 
Tonkin Bay (I, 1) on July 7 (A. 7). Two Hpeciniens. 

<;ei>f/raphic Banffe. Ottawa, Canada ; Mt. Washington, X. .H ; Ver- 
mont; Xew York; New Jersey; W. Pennn. ; Mich.; Wisconsin; Iowa. 

.55. Podahnts tumentomm Sav. A. Taken at the camp on Siskowit 
Bay (V, .3) on August 4 (<J. 201). 

ISeoijraphh Range. W. I'enna. ; Mich.; IllinoiH (Wolcotf) ; Colorado. 

.50. ilaHhodca n'lger I*c. Fonnd in a small rock ytool on the Lake 
shore (I, 1) on July 12 i^G. 75). 

Geographic Hange. Isie Boyale, Marquette, Mich.; fiake Superior 
region; Mt. Washington, N. H. 

i^carahaeidue. 

57. Oeotnipefi blackburnii Fab. A. Two of these beetles were taken 
about home dung on the Deiwr trail (IIT, 'U4) on August 21 (A. 143). 

Olographic Range. Ottawa, Canada; New York; Xew Jersey; Dist. 
Columbia; Ohio; Mich.; Wis. 

58. Hcriva vcitpertina Gyll. A. One specimen found on the grarelly 
lieach near the Ijigbt-houne (I, 1) on July 10 (G. 43). 

Geographic Range. Ottawa, Canada ; Nova Scotia : Saskatchewan 
basin; Vermont; New York; N. J.; I>ist. Columbia; Fla. ; W. Penna. ; 
Ohio; Mich.; No. Illinois and Indiana (Woicott) ; Wisconsin; Iowa; 
Nebraska; Kansas: Colorado; New Mexico. 

59. Diplotaxia Uberta Germ. A. A single dead specimen {Q. 102) was 
foand under a flat rock on a jack pine ridge (I, 5). 

Geographic Range. Isle Bovale, Mich; W. Penna.; New Jweer; Dist, 
of Col. 

(JO. /.arhnoatema arctiata Smith, A. Taken at the Lighthouse camp 
(I, 7), on July 26 (G. 166), 

Geographic Range. W, Penna.; Dist. Columbia; Michigan; Elliot and 
Carbondale, 111. (Woicott). 

61. Trichius a^nis Gory. Very abundant in the flowers of the Cow 
Parsnip (Seraclcum lanatum) in the clearing at the Lighthouse. (I, 7) 
during July (G. 26, 45, 49, 105, 133, 136, 137). 

Geoyru])hi<- Range. Saskatchewan basin: Ottawa, Canada; Nova 
Scotia; N. H.; New York; New Jersey; Virginia; W. Penna.: Ohio. 
" "■ " Wisconsin; Iowa; Colorado; New 

mbgcidae. 

\. Two specimens were taken on Sis- 
kowit Bay {V, 3) on August 15 (G. 239) and August 16 (A. 153). 

Food plants. I^rva feeds on the inner bark of dead and dying oaks 
and hickory. Probably has other food plant as hickory was not found 
on the island and oak is of very rare occurrence. (Fett,''06, p. 433.) 

Geographic Range. Mass. to .\labama ; W. Penna. ; Ohio { l>ury ) ; New 
Y'ork; Mich.; Wisconsin; Kansas; Colorado; Arizona. Hamilton, 'y7a, 
p. 395. IVobably introduced. 

63. Xylotrtvhiia undulatua Say. This active beetle was- exceedingly 



Mich.; No. 
Meiico. 


Illinois 


(Woicott) ; 










Cera, 


62. 


Phymatodi'H n 


inaJiilh 


iFnb. 



200 MICHIGAN SURVEY, 1908. 

abundant about the cnnips at the Ligfat-hoxise (I, 7) during July and 
on SiBkowit Bay <V. 3) during August. There is couBiderable variation 
in the tcUow elytral markings in the series secured. (G. 86, 212, 222, 
231, 239), (A. 5, 152). 

Food plants. Has been found on hemlock and spruce, but as hemlock 
is not found on the island, spruce is probably the food plant. (Felt, '06, 
p. 671). 

Qeoffraphic Range. Ottawa, Canada ; Lake Superior ; New Hampshire ; 
"New York; New Jersey; W. Penna.; Mich,; Wisconsin; Iowa; Nebr. ; 
Kansas; New Mexico; Northwest Terr.; Colorado; British Columbia. 

U4. Pachyta Uturata Kby. A. The one specimen is from the camp 
at Rock narbor (1, 7) on July 31 (G. 191). 

. Qeographie Range. Vermont; Hudson Bay region; Mich.; Colo.; 
New Mexico (Psyche 9, p. 303) ; Washington; Idaho; British Col. Ham- 
ilton, '94, p. 31. 

65- Acmaeops pi-oteua Kby. One taken at the Siskowit camp (V, 3) 
on August 15 (G. 239). 

Geographic Range. Labrador; Ottawa, Ontario; "common through 
Canada;" Hudson Bay; Saskatchewan basin; Mt. Washington (sum- 
mit), N. H. ; Mass.; New York; Mich.; Wisconsin; Kansas; New 
Mexico; Montana; Colo,; Oregon; Brit. Columbia. 

66. BelJamifa scalaris Say. A. A single specimen of this slender 
beetle came from the Light-house camp (1, 7) on July 22 (G. 133). 

Food plants. Beetle and larva have been found under the bark of the 
Yellow Birch. {B. lutoa) and has been found ovipositing on maple. 
(Beutenmuller, '96, p. 77.) 

Oeogmphio Range. Saskatchewan basin; Ottawa, Canada; N. II.; 
New York; W. Penna.; New Jersey; Maryland; Va.; La,; Ohio (Dury) ; 
Mich.; Wisconsin. 

67. Lcptura suhargenta-ta Kby. One specimen from the Light- 
house camp (1, 7) on July 11 (G. 49). 

QeograplUc Range. Canada ; Hudson Bay and Lake Superior region ; 
N. H.; Mass.; New York; Dist. Columbia; Georgia; Mich.; Ohio 
(Dory) ; Wisconsin; Montana; Colo.; New Mexico; Nevada; Utah; 
Calif.; Washington; Brit. Columbia; Alaska. 

68. Leptura nigrella Say A. A single specimen was taken at the 
Siskowit camp (V, 3) on August 7 (G. 212). 

Geographic Range. Ottawa, Canada; Hudson Bay region; Maine; 
Georgia; W. Penna.; Mich,; No. Illinois (Wolcott) ; Wisconsin; Colo.; 
New Mexico ; Nevada ; Washington. 

69. Lcptura sexmaculata L. A. Taken on the flowers of the Gow 
Parsnip in the clearing at the lighthouse (1, 7) during July {G. 105). 

Geographic Range. Hudson Bay to Lake Superior; Ottawa, Canada; 
Quebec; Mt. Washington (summit), N. H.; Mich.; Wisconsin; Colo.; 
Brit. Columbia; eastern and western Siberia; Alps and Europe. Ham- 
ilton, '94, p. 396. 

70. Leptura canadensis Fab. A. Only two specimens of this i-eH 
shouldered beetle were taken, one from the camp on Siskowit (V. 3> 
on August 7 (G. 212), and the other on August 13 (G. 232) from near 
the head of Siskowit Bay {VIII, '04). 



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BCOLOGY OP ISLB ROTALB. 201 

Food plants. Larva barrows in spruce and hemlock. (Beutenmnller, 
'96, p. 78). 

Oeographic Range. Ottawa. Can.; Nova Scotia; N. H.; Vermont; 
Mass.; New^ York; Penna.; ViiTiinia; Qa,; Mich.; WiBconBin; Mo.; Colo.; 
Hew Mexico ; No. ArizOQa ; No. Idnho ; Brit. Colnmbia ; eastern an4 
■western Siberia; Japan; Russia; Germany. Hamiltou, '94a, p. 396. 

71. Leptura chri/socoma Kby. This bright yellow beetle was the 
7i]0st abundant Oerambycid, occurring in great numbers in the flowers 
of the Cow Parsnip in the clearing at the Light-house (1, T) ; also 
found in the flowers of the Wild Rose on the beach (I. 1) ; and on the 
flowers of Opniaatcr opuUfoUiig. at the month of Benson brook (II. 1> 
during Julv. Also taken at the Siskowit camp (V, 3) on August u. 
((.). :17, 45. 49, 105, 133. 137, 148, 191.) 

Oeoffniphir Range. Ottawa. Can.; Hudson Bay region; Nova Scotia; 
Haine; N. 11.; New York; Mich.; Wisconsin; Colo.; New Mexico; No. 
Arizona; Priest's Ijike, Idaho. (T\'olrott) ; Utah; Nevada; Calif,; Brit. 
Columbia. 

72". Leptura proxitna Say. A. Two specimens were found on the 
flowers of the Cow Parsnip {I. 7) in July (G. 105. 179), and another 
specimen at the camp on Siskowit Bay (V, 3) on August 3 (O. 195). 

Pood plant. Reared from nmple. (Wickham. Can. Ent., 29. p. 192.) 

Oeographic Range. Ottawa. Can.; Vermont; N. H.; Mass.; New 
York; W. Penna. ; Virginia; fia.; Pist. of Columbia; Oliio; Mich.; No. 
111. (Wolcott) ; Wisconsin; Iowa; Missonri, 

73. Leptura tibialis I^ec. A. The one specimen is from the camp 
on Siskowit Bay (V, 3) on Angust 16 (A. 152). 

Oeographic Range. Mt. Washington, New Hampsiiire; Michigan; 
Oi'^on. 

74. Leptura mutabilig Newm. Four specimens were taken at the 
Light-house {I, 7) during July (G. 49, 105, 137, 16C). Some of these 
were taken on the flowei-s of the Cow Parsnip, 

geographic Range. Saskatchewan basin; Ottawa, Can.; Mt. Wash- 
ington (summit), N. H.; New York; Dist. Columbia; Mass.: New Jersey; 
W. Penna.; Ohio (Dury); Mich.; Wisconsin; New Mexico. 

75. Monohammua acutellatus Say, A. Six specimens of these large 
beetles were taken: one at the Liglit-house (I. 7) on July 24 (G. 152), 
and the others on Angust 7. 12 and 16 at the Siskowit camp (V, 3). 
(G. 212, 2S1; A. 152). 

Food plant. Taken on white and hard pine: beetle girdles branched 
and the larva bores in spruce trunk. (Felt, 'Oti, p. 364.) 

Geographic Range. Ottawa, Can. ; Hudson Bay i-egion ; Saskatchewan 
basin; W. Penna.; St. Joseph (Wolcott), Isle Boyaie, Mich.: Wisconsin; 
Dulutb, Minn. (Wolcott) ; Colo.; New Mexico; Brit. Columbia; Alaska; 
extensive N. American range in "'pine regions," Itistrict of Columbia. 

ChryKomclidac' 

76. Donaeia protrima Kby. A. "In the water-lily zone of Snmner 
Lake (III, 5) on July 27 (G. 171). The beetles fly' low, dragging the 
tip of the abdomen in the water, and apparently alight only on leaves 
of the waterlily." Gleason. Also taken July 29 (A. iS4). 

*cl. Chliu-iKlen CSS) tor rood htiblls or ililn family. 



202 MICHIGAN SURVET, 1908. 

Geographic Range. Ottawa, Can.; Lake Superior; N. H.; Mass.; New 
York; Penna.; Mich.; Wie. ; Hudson Bay Terr.; Idaho; Calif. Leng. 
Trans. Am. Ent Soc., 18, p. 167. 

77. Donccia cincticomia Jiewm, A. "Three apecimenB were taken on 
July 27 and 28 at Snniner Lake (III, 5), associated with the preceding 
Blieciee and with the same habit," Gleason. (G, 171, 1751. 

Geographic Range. Canada; Vermont; New HampshiFe; MaBS.; New 
York; Michigan; No. Illinois; Texas. 

78. Orsodachna atra Alir. var.=cfti7drrn»' Kbv. Two specimens were 
taken at the Liftht-hoiise (I, 7) on July 11 (G. 49). Horn. Tr. Am. Ent. 
Soc.. '92, pp. 6-7. Ent. Amer., I, p. 9. 

Geographic R«uge. Saskatchewan basin; Ottawa, Canada; New Eng- 
land and south on the mountains to N. Carolina; W. Penna. ; Mich.; 
Wisconsin; No. III. (Wolcott); Iowa; Alberta; Colorado; New Mexir^; 
Arizona; California. Psyche, 9, p. 303; Brit. Columbia. 

79. GalcruccUa ngmphaea L. A. These leaf beetles were taken in a 
small bayou (IV, 3) connected with ^Tobin Harbor on July 21 (A. 42). 
Larvae, pupae, freshly emerged and fully covered adults were all rep- 
resented in very lat^e nunibei«. The lily leaves were riddled by the 
innumerable lai-vae. Cf. Chittenden, '05, p. 58 and Mac Glllivcay, '03, |>. 
3^5 for the life history of this species. 

Geographic Range. In Canada westward to the Mackenzie Basin and 
into Alaska; New York; Va.; Ohio (Dury): W. Penna.; Mich.; Colo- 
rado; Texjis; Oregon; Calif.; Siberia into Europe. Hamilton, '94a, p. 
398. 

TcnchrioHidac. 

80. Vpi" rcramhoiflci* L. A. A single specimen was taken at the 
Lighthouse (1, 7) on July 23 (G. 153). 

Geographic Range. Ottawa, Can,; Hudson Bay; Saskatchewan basin; 
Lake Superior; Nova Scotia; Maine; Mt Washington, K, H.; Vermont; 
New York; New Jei-sey; W. Penna.; Mich.; Wisconsin; Estherville, 
Cass Co., Minn. (Wolcott); Colo.; Montana; Manitoba; No. Asia; 
Siberia; No, EnrojMj: Germany. Hamilton, '94a, p. 400. 

CixtcUdac. 

81. CistcJa ^erirca Sav. .\. Found under loo^e stones on the jack 
pine ridge (I. 5) on July'l4 (G. 81). 

Pood plants. Has been found on pine, oak and basewood. (Felt, '06, 
p. 518.) 

Geographic Range. Michigan; W. Penna.; New Jersey; New Mexico. 

Jlciandrgidae. 

82. fferropalpii* harhaliiit Schall. A. One specimen was taken at 
Tobin Harlwr on July 19 (G. 129). 

Food plant. I^irva bores in sap and heart wood of balsnm and sprnce. 
(Pelt. '01!. p. 071). 

Geographic Range. Canada; I>ake Superior and Hndson Bay regioD8<; 
Maine; Vermont; New York; W. Penna.; West Virginia: Colorado; 
Rocky Mrs. south to New Mexico; Manitoba; Oregon; Brit. Columbia; 
Alaska ; Kil>eriti ; I'urope. 



ECOLOGY OP ISLE ROYALE. 203 

MordcUidar. 

83. An^i^pis riifa Say. A. Many Rpwiineng of this fi[>e(-ief< were takeu 
about the camp at the"l-ighthouBe (1. 7) ou July 28 (G. 179). 

Geographic Range. Ottawa, Can.; Mt. AVaahington (summit), N. H.; 
Vermont; New York; New Jersey; Dist. Col.; Florida; Oliio; Michigan; 
"WiBooDBin ; Wyoming; Colo,; Utah; Lower Calif.; New Mexico; Mexico; 
'W'ashinfrton ; Brit. Columbia; Alaelia. 

84. MordelUstcna biplagiatn Helm. A. One specimen was taken on 
flowers in the clearing at the Lighthouse (I. 7) on July 11 (G. 4ft). 

Qeograpbic Range. New York; Dist. of Colnmbia; Ohio; Mich.; Illi- 
nois; Wis. 

85. Mordellistctia arapiilarin Hav. A. Two specimens were taken at 
the Lighthouse (1, 7) on July 28 (G. 179). 

Geographic Range. Dist. of Columbia ; "Middle and l^'estern States" ; 
lUich. (Isle Royale) ; Ottawa, Canada. 

Ciirculionidae. 

86. HyJobivs paleg Hbst. A. .\ single specimen was taken at the 
I>igbthouse (I, 7) on July 13 (O. 86). 

Food plant. Larvae lire in bark of white pines. (Felt, '06, p. 664). 
Geographic Range. Ottawa. Canada ; Maine to Florida ; Michigan ; 
\V. Penna.; Dnlnth, Minn. (Wolcott). 

87. HypomolyT pineti Fab. A. This lai^ snout beetle (G. 179) was 
taken July 28 in the Lighthouse clearing (I, 7>. 

Geographic Range. Canada ; Hudson Bay region ; Saskatchewan 
basin; Mich.; Wisconsin; Siberia; Enrnpe. 

88. Magdalig. "Apparently new," Wickham.; Taken at the Light- 
house camp {I, 7) on Julv 23 (G. 130), at Siekowit (V, 3) on August !"► 
(G. 239). 

Calandridac. 

89. Comoniin subarcata^ Boh. A. Taken at the Siskowit Camp (V, 3> 
on August 7 (G. 212). 

Geographic Range. Mt. Washington. N. H. ; Michigan; Wisconsin: 
OlendoQ Park, III. (Wolcott) ; Iowa; Kansas; Nebraska; Colorado; New 
Mexico; "Middle States." 



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MICHIGAN SURVEY, 1908. 



2. mPPLEMENTARY LIST OF ISLE ROYALE BEETLES. 

BY A. B. WOLCOTT, 

Field Museum of Natural History, Chicago. 

Thin Rupplementary lint of speciee records from Isle Royale all the 
speries taken by Hubbard and Schwarz ('78, pp. 627-043) but not foond 
in the 1905 collectiono. These two lists make a complete cataloff of the 
species so far found on this island, exocptinff those species which are 
scattered in the literature and have thus been overlooked. The general 
geographic range of each speoies is giren. 

Carabidac. 

1. Bp.mhidittm concoUir Kby. Xew York; Maine; Canada; Michigan 
( M ichipicoton River) ; Wyoming; Maine to the Pacific coast. 

2. Bembidium planatiim Lev. Michigan (Isle Royale) ; Colorado; 
Wyoming; Nevada; Oregon; Washington to British Columbia. 

\i. fntrobiua lonfficornis Say. Sew Jersey; Vermont; New York; 
I>i»t. Columbia; Ohio; Pennsylvania; Canada; Michigan (Escanaba) ; 
Wisconsin; Illinois; Indiana; Iowa: Colorado; Texas; New Mexico. 

4. PteroKtichus punctatisshiiua Rand. Massachusetts; New Hamp- 
shire; Vermont; Maine; Canada; Hudson Bay region; Michigan (Mich- 
ipicoton Island) ; Arctic Sibera; the Amur; Dnuria. 

5, Pierostichus mandibularia Kby, var. New Hampshire; Vermont; 
Massachnsetts ; Canada; Wisconsin; Michigan (Marquette. Michipicoton 
River); Hudson Bay region; Alaska; Arctic ^il>eria. 

0, Amara latior Kby. New Jersey; New Hampshire; Canada; Mich- 
igan (Escanaba. Ann Arbor) ; Wisconsin; Illinois; Nebraska; Colo- 
rado; Idaho; New Mexico; .\ri7,ona; Vancouver Island. 

7. Amara impuncticoUis Say. Hist. Columbia; Ohio; Michigan (De- 
troit) ; Wisconsin; Canada; Montana; Colorado; New Mexico. 

8. Calaihns advcna var. mollis Mots. Vermont; Maine; ^lichigau 
(Michipicoton Kiver, Michipicoton Island); Alaska. 

!t. Platyntis aemginoms l>ej. Dist. Columbia ; Indiana (Pine) ; 
Illinois (Chicago); Michigan (Escanaba, l>etroit) ; Wisconsin. 

10, Dromiua picctis r>ej. New Jersey; New York: Dist, Columbia: 
Massachusetts; Ohio; Michigan (Manjuette, Detroit); Wisconsin; Can- 
ada: Iowa; California. 

11, HarpaUta fulvilahris Maun. Michigan (Marquette, Michipi- 
coton River). 

12, Harpalus rufimatim I>ec. Jlichigan (Escanaba, Marquette) ; Wis- 
consin; Canada; British Columbia. 

i:i. Harpalus laticepa T-ec, New Hampshire (iSnmmit Mt, Washing- 
ton); Michigan (Escanaba, Mai-quctte, Lake Hui-on) ; Wisconsin; Can- 
ada (Ottawa); Colorado, 

14. Bradj/cflhin cordicollis Ta-c. New Ilaniiishii-e (Mt. Washington) ; 
Michigan (Marquette). CoOqIc 



ECOLOGY OF ISLB ROTALE. 205 

Hydrophilidae. 

15. Crenophilua {Hydrohiiis) diffeatus 1*0. Miohigan (Marquette, De- 
troit). 

fill phi dar. 

16. Necrophorua ^ re-ipilloidei* Hbst. New Jernev; New Hampshire 
(Mt. Washington);' Michigan (EHonnaba. Mii-hipicoton Island) ;'^^'i(*- 
consin; HtidsoD Bar Territor,v; Nova Bcotia; Ontario: Manitoba; Brit- 
ish Columbia ; Alaska ; Wa«bington ; Oregon ; East Siberia ; Kamt- 
schatka; Amurland; Europe; China. 

17. Choleia baniUaria Sa.v. New Jersey; New IIanii>shire (Mt. Wash- 
ington) ; Ohio; Michigan (Bault de Kte. Marie, Detroit); Wisconsin; 
Nebraska; Kansas; Canada: Hudson Bay Territory; British Columbia; 
Alaska; Nevada to Colorado; California. 

18. Choleva {Catopjt) terminann Lee. Virginia; New Jersey; Mass- 
achusetts; Dist, Columbia; Ohio; Illinois; Michigan (Bachewauung Bay, 
Michipicotou Island); Wisconsin; Canada (Ottawa). 

19. Anistoma aaaiinilis Lee. Dist. Columbia; New Hampshire (Buni- 
mit Mt. Washington) ; Michigan (Marquette, Michipicoton River) ; Wis- 
consin; Canada; Colorado; Vancpuver Island. 

20. Liodea globma Lee. New Hampshire (Mt. Washington) ; Michigan 
(Marquette) ; Canada (Ottawa) ; Colorado; New Mexico. 

21. Agathidium revolrens Lee. Canada (Ottawa) ; British Colum- 
bia; New Mexico. 

22. Clambus gibbulus Lee. Florida; Dist. Columbia; Michigan (Mar- 
quette, Detroit); Colorado; S. Arizona. 

Paelaphidae. 

23. Tychus longipalpua Lee. Florida ; Dist. Columbia ; Michigan 
(Marquette); Canada (Ottawa). 

24. Reichenbackia (Bryaxis] propinqua Lee. Canada (Ottawa) ; 
Michigan (Marquette, Point aux Pins); Colorado (si»ecies doubtfully 
identical). 

8taphyUnida€. 

25. Qvediiis lticrigatiif< Gyll, Geot^ia; New HampHliire (summit Mt. 
Washington); Massachusetts; Pennsylvania; Ohio; Illinois; Michigan 
(Marquette, Bachewanung Bay, Detroit); Canada; British Columbia; 
Alaska; Or^on; Nevada; Colorado; Kansas; New Mexico; California; 
eastern Siberia; northern and Alpine Europe. 

26. Stenus aemicolon Lee. Dist. Colu^iibia; Michigan (EscanabOr 
Marquette, Basbewanung Bay, Micbipicoton River), 

27. Lathrobium tcrmtnatum Grav. (piinc(M/a(«m Lee.). Florida; 
Georgia; Dist. Columbia; New Jersey; W. Pennsylvania; Ohio; "East- 
em States"; Massachusetts; Michigan (Escanaba, Marqnette, Detroit); 
Wisconsin; Iowa; Canada; Kansas; Colorado; Europe and Siberia. 

28. Tachinua fumipetinis ,Say. Florida; Dist. Columbia; Michigan 
(Marquette); Wisconsin: Colorado. 

29. Bolitobius cvngiilatus Mann. Virginia; New Jersey; New Hamp- 
shire (Mt, Washington); Pennsylvania; Michigan (Sault de Ste. Marie, 

27 



i06 MICHIGAN SURVEY. 1908. 

Bachewaniing Bay, Detroit) ; Wiflconsio; Canada; Oregon; Queen Char- 
lotte iBland; Britieh Columbia; Alaska; Caucasia; Europe. 

30. HobroceruS matfnus Lee. Michigan (Marquette). The type of 
this speoies came from Isle Royale. 

31. OliBthaerus mcgaoephalva Zett. Michigan (Michipicoton laland) ; 
Canada; Alaska; California; Siberia; Lapland; Sweden; Hungarj; 
Arctic and Eastern Siberia. 

32. Olisthacriis aub^triatua Payk. (nitidua JjCC.). Massachusetts; 
Michigan (Midiipicoton, Eagle Harbor) ; Wisconsin; Sweden; Germany; 
France; Arctic and Eastern Siberia. 

33. Ancyrophorua planua Lee. New Hampshire (Mt. Washington) ; 
Michigan (Isle Boyale). 

34. Anthophagua vertwalia Say. Michigan (Marquette, Detroit). 

35. Acidota creanta Fabr, {aeriata Lee). Massachusetts; Common 
on Islands and shores of Lake Superior; Michigan (Marquette, Mich- 
ipicoton River, Detroit) ; Canada; central and northern Europe; Siberia. 

36. Arpedium sp. Michigan (Marquette). 

Phalacridae. 

37. Phalacrua poUtua Melsh. Florida; Dist. Columbia; Ohio; Illinois; 
Michigan (Marquette, DetTOit) ; Canada (Ottawa); Iowa; Colorado. 

CoccineUidae. 

3*?. CoccineUa perplexa Muls. (trifasciata Linn.). New York; New 
HamjiRhire (Mt. Washington); Canada; Hudson Bay Territory; Mich- 
igan (Detroit. Marquette, Au Train Falls, St. Joseph); Wisconsin; Ill- 
inois (Chicago, taken by Wolcott) ; Alaska; Vancouver Island; Oregon; 
Washington to California; New Mexico; Kamtschatka through northern 
Siberia and Europe to Lapland. Circumpolar. 

39. Coccinetta tranaveraoguttata Fald, var, tranaveraalia Muls. The 
typical form or its varieties are known from New Hampshire (summit 
Mt. Washington) ; Greenland; Hudson Bay region; various places in 
Canada; British Columbia; Northwest Territory; Alaska; Illinois (Chi- 
cago, Wolcott coll.) ; Michigan (Bachewanung Bay, Chatham ; Wisconsin; 
Minnesota (l)uluth, Wolcott coll.); Nebraska; Nevada; Colorado; New 
Mexico; California; Rocky Mountains and Pacific r^ions to mountain- 
ous Mexico; eastern Siberia; Japan; northern China; Dauria; Lapland. 
Circumpolar. 

40. Cycloneda aanguinea Linn. Floiida; West Indies; "United States 
and Canada generally"; Michigan (Michipicoton River, Chatham) ; Wis- 
consin; Illinois; Indiana; Ohio; New Jersey to Colorado; New Mexico; 
N. Arizona ; Texas ; Baja California ; Europe. 

41. Cleia {Sarmonia) picta Band. Dist. Columbia; Pennsylvania; 
Canada to Colorado; New Hampshire (summit Mti Washington) ; Mich- 
igan (Escanaba, Marquette); Minnesota (Dulnth, Wolcott coll.); New 
Mexico. 

42. ficymnua lacustria Lee. Michigan (Escanaba, Marquette) ; Col- 
orado; Arizona. 



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ECOLOGY OP ISLB ROyALB. 



Endomychidae. 



43. Lycoperdina ferruyinea Lee. Dist, Culumbia; New Jersey west 
to Colorado; New Hampshire (Mt Washington) ; New York; "Middle 
and Southern States"; Ohio; Illinois (central and northern) ; Michigan 
(Bachewannng Bay, Detroit); Canada; Wisconsin; Iowa; Colorado; 
New Mexico, 

HUteridae. 

44. Biater haaalia 1*0. Ohio: Michigan (Marquette). 

45. Ptegaderua sayi Mara. "Middle States"; Michigan (Sault de Ste. 
Marie, Marquette); Canada; Wisconsin; Colorado; New Mexico. 



46. Omoaita diecoidea Fabr. Canada; Michigan (uorthem) ; Colo- 
rado; New Mexico; Europe and the Pa>ciflc States, east to Colorado. 

Lathridiidae. 

47. 8tepAo8t€thua (Laihridua) liratux Tjec. Dist Columbia; Ohio; 
Canada (Ottawa) ; Michigan (Detroit) ; Queen Charlotte Islands, Brit- 
ish Columbia. 

48. Lathridiua tninutua Linn. "Nearly all North America"; Dist. 
Columbia; Michigan (Detroit) ; Wisconsin; Colorado; "Alaska to Louis- 
iana and to Massachusetts and eastern Canada" ; all Europe and north- 
em Asia to Kamtschatka. 

49. Corticaria aerrtcolUs Lee Michigan (Michipicoton River, De- 
troit) ; British Columbia. 

Byrrhidae. 

50. Bffrrhua geminatua Lee. New Hampshire (summit Mt. Washing- 
ton) ; Michigan (Isle Royale only). 

Daacyllidac. 

51. Macropogon picena Lee. Michigan (Isle Royale only). 

52. Eurypogon niger. Michigan (Michipicoton River). 

53. Eascinettta terminalia Lee. New Jersey west to Colorado; New 
York; Vermont; Ohio; Illinois; Michigan (Bscanaba, Marquette, De- 
troit) ; Canada. 

Elaterid^Ie. 

54. Cryptohypnua bicolor Jlach. This species is believed to be merely 
a variety of nocturttim Esch. which is recorded with the variety from 
the following localities; — Labrador; Hudson Bay regions; New Hamp- 
shire (summit and alpine regions Mt. Washington) ; Canada; Michigan 
(Marquette, Sault de Hte. Marie) ; Dakota; Wisconsin; Utah; Colorado; 
Montana; Idaho; New Mexico; Or^on; British Columbia; Alaska; 
Kamtschatka; eastern Siberia. 

55. Crytohypnua tumcscena Lee. Michigan (Sault de Ste. Marie); 
Colorado; New Mexico. 

56. Elater nigrinus Pajk. var.? Elatcr nigrinua occurs in Vermont; 
Canada (Ottawa); Michigan (Escanaba, Marquette, Detroit); Alaska: 



208 MIGHIGAN aURVET. 1908. 

Vancouver Inland and Queen Charlotte Island; British Columbia; Kew 
Mexico; northern and central Europe; west Siberia; Amurlaud. 

57. Slater mtxtm Hbst. Dist. Columbia ; New Hampehire (sunmut Mt. 
Washiugton); Canada (Ottawa); Michigan (Marquette, Micbipicoton 
Island) ; Colorado. 

58. lietarmon bigeminatus Rand. Dist. Columbia; Canada (Ottawa) ; 
Michigan ( Marquette) . 

59. Melanotus Leonardr Lee. Michigan (Marquette, Detroit). 

60. AfeUmotus castanipea Payk. {acrobieotlia IjCc). "Middle States to 
Canada"; New York; Vermont; New Hampshire (summit Mt. Washing- 
ton) ; Dist. Columbia; Ohio; Pennsylvania; Canada; Michigan <Esca- 
naba, Marquette, Detroit) ; Wisconsin; Colorado; Europe; West Siberia; 
Amurland. 

61. Limon^a aeger Lee. New Jersey; New Hampshire (Mt. Washing- 
ton) ; Canada (Ottawa); Michigan (Marquette); Wisconsin. 

62. Campylus denticornis Kirby. New Hampshire (summit Mt. 
Washington); Maine; Peunsylvania ; Canada (Ottawa); Ohio; Mich- 
igan; (Marquette, Port Huron); Wisconsin. 

63. Paratuymvs costalis Payk. New Hampshire (summit Mt. Wash- 
ington) ; "The northern shore of Lake Superior"; Labrador; Europe 
(Sweden, Finland, Lapland) ; Amurland. 

64. Sericoaomua incongruua Lee, Canada (Ottawa) ; Michigan (Mar- 
quette) ; New Hampshire (Mt. Washington). 

65. Corymhitcs resplendcns Escb. Newfoundland ; Maine ; Lake 
Superior region northward to 56°; Vermont; Canada (Ottawa); Mich- 
igan ( Michipicoton Island, Marquette) ; New Hampshire (summit Mt. 
Wtibhington) ; Wisconsin; Queen Charlotte Island; Dritish Columbia; 
Alaska. 

66. Corymbitea apinoaua 1*0. New Hampshire (summit Mt. Wash- 
ington) ; Canada (Ottawa) ; Michigan (Escanaba, Marquette) ; Wis- 
consin; Iowa. 

67. Corymbitea mendax Lee, Michigan (Eagle Harbor). 

68. Cori/miitea inaidiosua Ijec. New Hampshire (Mt. Washington) ; 
Michigan (Marquette) . 

69. Corymbitea falalficua I^c. New Hampshire (summit Mt, Wash- 
ington) ; Canada; Michigan (Marquette): Wisconsin, 

70. Corymbitea triutidtilatus Rand. New Hampshire (summit Mt. 
Washington) ; Maine; Vermont; Michigan (Marquette) ; Canada 
(Ottawa); Wisconsin; Colorado. 

71. Corynibites propnia I^ec. New York; Vermont; New Hampshire 
(summit Mt. Washington); Canada; Michigan (Michipicoton River, 
Marquette) ; British Colmnbia. 

72. Corymbitea nigricolHa Bland. Michigan (Marquette); Colorado. 

73. Corymbitea aplendena Ziegl. Dist. Columbia ; Ohio ; Canada 
lUltawa) ; Michigan (Marquette). 

74. CorymAitea nigricomia Panz. New Jersey; New Hampshire 
(summit Mt, Washington); Massachusetts; Illinois (Ft, Sheridan, Wol- 
cott) ; Michigan (Marquette, Detroit) ; Canada; Iowa; WiBCOUsin; Col- 
orado; central and boreal Euro^ie and Siberia. 



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BCOLOOT or ISLE ROTALE. 



Buprestidae. 

76. ,Melanophila fulvoguttata Harr. New Hampshire (summit Mt. 
Washington); Canada (Ottawa); Michigan (Escanaba, Marquette, Port 
Huron) ; Eanaas. 

Lampyridae, 

76. Plateros (Eros) modcstus Say. Florida; Dist. Columbia; New 
Hampshire (summit Mt. Washington); Ohio; Michigan (Detroit, Mar- 
quette) ; Canada (Ottawa) ; Iowa (McGregor, Wolcott) ; New Mexico. 

77. EUychnia (Photintm) corrusca Linn. "Common in Canada and 
most of the United States Mst of the Rocky Mountains" ; Dist. Columbia ; 
Virginia; Georgia; New Jersey; New Hampshire (summit Mt. Washing- 
ton); New York;, Ohio; Indiana (Wolcott); Illinois; Michigan (Mich- 
ipicotoQ River, Detroit); Iowa; Wisconsin; Nebraska; Kansas; Col- 
orado; New Mexico; Arizona.; Canada (Ottawa); Nova Bcotia; North- 
west Territory. 

78. Podabrug modeatus Say. Georgia; New Jersey; New York; New 
Hampshire (Mt. Washington); Pennsylvania; Ohio; Michigan (Esca- 
naba, Mu-quette, Detroit); Canada (Ottawa); Iowa; Wisconsin; Colo- 
rado. 

79. Podabrus lacvtcoUia Kby. New Hampshire (Mt. Washington); 
Michigan (Marquette, Michipicoton River) ; Colorado. 

80. Telephorua CurtisU Kby. New Hampshire (summit Mt. Washing- 
ton) ; Michigan (Marquette, Michipicoton River); Wisconsin; Iowa; 
Hudson Bay region ; British Columbia. 

81. Malthodd$ laticolUg Lee. [traneversus T^ec.). Michigan (Isle 
Ko.vale only). 

82. Malthodes condavus Lee. Diet. Columbia; Michigan (Marquette, 
Detroit) ; Colorado. 

83. Malthodes fragilis I*c. Michigan (Detroit). 

Cleridae. 

84. Tha)iasimus (Clerus) undatulus Say. New York; Vermont; Maine; 
New Hampshire (summit Mt. Washington); Canada; Michigan (Mar- 
quette, Escanaba); Minnesota; Hudson Hay north (o lat. I>5°; Kansaa; 
Colorado; New Mexico; variety nubilii8 occurs in Northwest Territory 
and Alaska. 

Ptinidae. 

85. THnodervfi substriatns Payk, New Hampshire (summit Mt. 
Washington) ; "Northern States"; Canada; Pennsylvania; Michigan 
(Escanaba, Marquette, Bachewanung Bay) ; Alaska; eastern and western 
Siberia; Europe. 

Cioidae. 

86. Cis creberrimua Melli4. Florida; Dist. Columbia; Ohio; Michi- 
igaa (Marquette, Detroit). 



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210 HIOHIQAN SURVET, 1908. 

Cerambycidae. 

S7. Tetropium cinnam'opterum Kirby. New Jersey; New Hampshire 
(summit Mt Washington); Vermont; PennBylvania ; Canada; Michigan 
(Marquette); Wisconsin; Colorado; New Mexico; northern and mount 
ainous Arizona; California; Oregon; Washington; Northwest Terri- 
tory; British Columbia; Alaska; "north to 55°". 

SB. . Phymaiodes macitUcollis Tjec. New Hampshire (Mt. ^Vashing- 
ton) ; Michigan (Isle Royale-type locality) ; Colorado (7-9000 ft. el.). 

89. MicToclytus gazellula Hald. (Crytophorua gibbuJua Lee.). Dist 
Columbia; New Hampshire (Mt. Washington); Canada (Ottawa); 
Michigan (Detroit). 

90. Pachyta monticoJa Rand. New York; New Hampshire (summit 
Mt. Washington ) ; Vermont ; Maine ; Massachusetts ; Pennsylvania ; 
Michigan (Marquette) ; Wisconsin; Canada (Ottawa) ; Antlcosti Island; 
Alaska. 

91. Leptura rvfula HaM. Michigan (Isle Eoyale only). 

92. Pogonocherus mia^tus Hald. Dist, Columbia; New Jersey; New 
York; New Hampehii-e (summit Mt. Washington); Indiana (Clarke 
Junction, Dune Park, Wolcott coll.) ; Canada; Michigan (North Muske- 
gon, Marquette, Michipicoton River, Port Huron) ; Kansas; New Mexico; 
Colorado; Northern Arizona. 

Chryaomelidac. 

93. Zeugophora varians Cr. New Jersey; New Hampshire; Penn- 
sylvania; Indiana (Pine-Wolcott coll.); Illinois (Glen Ellyn, Wolcott 
coll.); Canada; Michigan (Detroit); Wisconsin; Kansas; Washington, 

94. Syneta fcrnigirwa Cerm. Dist. Columbia; Maryland; New Jer- 
sey; New York; New Hampshire (Mt, Washington); Vermont; Mass- 
achusetts; Ohio; Illinois (central and northern) ; Michigan (Mar- 
quette) ; Canada (Ottawa) ; Wisconsin; Nebraska; Colorado; New- 
foundland. 

95. Basmre.tin mamiiiifn- Newm. var. nrlUttus Suffr. {Cryptocephalu* 
Kcllafus Suffr.). DJst. Columbia; New Jersey; "Middle and Western 
States"; Ohio; Indiana (Clarke, Hessville, Wolcott coll.); Michigan 
(Escanaba, Mai-qnette, Detroit, North Muskegon, Holland); Wisconsin; 
Iowa ; Canada ; Colorado. 

96. PackybrachyH sp. Michigan (Sault de Sfe. Marie, Marquette). 

97. fionioctvna pallida Linn. New Hampshire (summit Mt. Washinft- 
ton) ; Michigan (Marquette, Bacbewanung) ; Minnesota; Wisconsin ; 
('olorado; Hudson Bay region generally; Europe and Siberia. 

9K Phylhdecta viilf/rithsiina Lii\n. \^i^inia; New Jersejj; New 
Hampshire (summit Mt. Washington) ; New York; Pennsylvania; Ohio; 
Illinois (central) ; Michigan (Detroit) ; Wisconsin; Iowa; Canada 
(Ottawa) ; Iceland; Siberia; China; Turkestan; Canaries. Perhaps also 
in Alaska. 

Cistelidae. 

99. Hymenorus nigcr Melsh, Florida ; Texas ; Dist. Columbia ; 
New York; New Hampshire (Mt. Washington); Pennsylvania; Ohio; 
Canada (Ottawa); Michigan (Escanaba, Marquette, Detroit); Wiscon- 
sin; Colorado. ^ ,, CoOgIc 



BCOLOGT OP ISLE ROTALB. 211 

Melandryidae, 

100. Emmem connect-;n8 Newm, New Hampshire (summit Mt. Waeh- 
ington) ; Michigan (Uarqnette). 

101. Scotochroa ftfwn/w Lee. Canada (Ottawa) ; Michigan (Esca- 
naba, Marqoette) ; Colorado. 

Pythidac. 

102. Lecontia (Crpmodes) disicoIUs Lee. New Hampshire (summit 
Mt. Washington); Alichigan (Marquette); Manitoba; Canada; (Ot- 
tawa) ; Idaho; Colorado; New Mexico. 

103. Boroa vnicotor Say. Piat. Columbia; Michigan (Marquette); 
Canada (Ottawa). 

104. Rhinoaimua viridiaencua Rand, (nitens Lee). Dist. Columbia; 
Michigan (Detroit, Marquette). 

Ciirculionidae. 

105. Pis»odvs duViut Hand. New Hampshire {Mt. Washington); 
Canada; Michigan (Maniuette) ; Wisconsin. 

100. Dori/tomiis brevimllia Lee. DiKt. Columbia; New Jersey; New 
York; New Hampshire (Mt. Washington); Ohio; Michigan (Marquette, 
Detroit); Colorado; New Mexico; Canada: Vancouver Island. 

107. Trichalophiia aUnnatua Say. Michigan (Michipicoton River) ; 
Wyoming (Laramie) ; Colorado. ' 

108. Apion sp. Michigan (Marquette). 

109. Maffdalia hiapoides Lee. Djst. Columbia; Michigan (Marquette, 
Port Huron); Colorado; British Columbia. 

110. Magdalia gentilia I^c, Michigan (Marquette); Colorado; Cali- 
fornia. 

lU, Magdalis armicolUa Sav (Magdalis alvtacca Lee. Bui. U. S. Geo!, 
and Geogr. Surv. Terr., 4 p. '4(i3, 1878). J*Conte described alutacea 
from Isle Royale, Lake KiiiHriof (Mr. K. A. Hchwara) and I/cavenworth 
Valley, above Georgetown, Ci>lunulo, sfiecimens; the species is not given 
in Hubbard and Schwarz's list. It has since t»een found at various 
places in the mountains in Colorado; Ohio; Canada (Ottawa); New 
Ilamiwhire (Mt. Washington) ; and a species doubtfully referred here 
occurs in New Mexico. 

112. Antkonomua cornilua Lw. Dist. Columbia; Ohio; Illinois 
(Itiver Foreflt, Bowmanvi lie, Wolcofl coll.); Miohigjin (Marquette, De- 
troit). 

Iia. Paeudanthonomun (Anthoiiomus) rralacffi Walsh. Florida; 
Disf. Columbia; Ohio; llliuuis (cenfrul and northern) ; Michigan (De 
iroit, Marquette). 

114. Orcheatca pallwiriiin Say, "Nova Kcolia to Texas, and to Puget 
Sound" (LeConte) ; Dist. Columbia; New Hampshire (Mt. Washington) ; 
Ohio; Michigan (Escanaba, Marquette, Detroit). 

115. Orchestcs cantia Horn. Ohio; Tyjie locality given thus; "Speci- 
mens are before me from Isle Koyale and Kscauaba. Michigan, and from 
San Juan, Colorado" Horn, Also known fi-oni Marquette, Michigan. 

116. Ctvemogonus epilobH Payk. Michigan (Marquette) ; British Col- 
umbia; Great Slave Lake, Northwest Territory; Colorado; northern and 
central Europe. 



MICHIGAN SURVEY, 1908. 



117. Dendroctonus rufipennia Kbj. Alaska; "Vancouver to Anti- 
costi. New BmnBirick and sonthweBt to Florida and New Mexico"; West- 
ern PennsylTania ; Michigan (Marqnette). 

. RGPERKNCBS. 

Refhiine, C. J. 8. (compiled by). 
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[From Kirby's Fauna Boreali-Americana; Insecta.] 
Beutenmnller, W. 
1893. On the Food-Habits of North American Rhyncopbora. Jonr. 

N. Y. Ent Soc, I. pp. 36-43, 80-88. 
1896, Food-Habits of North American Cerambycidae. Jour. N. Y. 
Ent. Soc., 4, pp. 73-81. 
Blanchard, F. 

1889. List of Buprestidae of New England. Ent. Am., 5, p. 29-32. 
Bowditch, F. 0. 

1896. List of Mt. Washington Coleoptera. Psvche, 7, Bnpp. pp. 1-11. 
Caudell, A. N. 

1903. Some Insects from the Summit of Pike's Peak, Found on Snow. 
Pro. Ent. Soc. Wash.. 5, pp. 74-82. 
Carpenter, W. L. 

1875. Report on tlie Alpine Insect-Fauna of Colorado. Ann, Rep. U. 
S. Oeol. Surv-ev Terr, for 1H73. (Havden), pp. 539-S43. 
Chittenden. F. H. 

1889. Notes ou the Habits of Buprestidae. Ent. Am., 5, pp. 217-220. 
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1900. Insects and the Weather: Otwen-ations Daring the SeasOD of 

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1901. Insects ami the Weather During the Benson of 1900. Bull. .30. 

U. K. Div. Entomolog.v, U. S. Dept. Agr.. pp. 63-75. 

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1893. The Entomology of the Mid-Alpine Zone of Custer County, 
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1906. Interglacial Periods in Canada, Inter- Oeol. Congress, Mexico, 

1906, reprint, pp. 1-26. 
Dury. C. 

1902. A Revised List of the Coleoptera Observed near Cincinnati, 

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Evans, J. D. 

1903. List of Canadian Coleoptera. Can. Ent., 35, pp. 239-243, 288- 

292, 317-320. 



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BCOLOOY OF ISLB ROTALB. 213 

Pall, H. C. 

1902. Some Insects of the Hudsonian Zone in New Mexico— Coleop- 
tera. Psyche 9. p. 303. 
Fall, H. C, and Cockerell, T. D. A. 

1907. Tlie Coleoptera of New Mexico. Trans, Am. Ent. Soc., 33, 
pp. 145-272. 
Fauvel, A. 

1889. Liste des Coleopteres Comiuana a I'Enrope et a TAmerique dn 
Nord. Rer. D'Rntomologie, 8, pp. 92-174. 
Feit, E. r. 

1905-OC. Insects affecting Park and Woodland Trees. Mem. 8, N. Y. 
St. Mns., Vol. 1, 1905; Vol. 2, 190C. 
Hamilton, J., and Hennbaw, S, 

1891-92. A List of Some of the Catalogues and Local Lists of North 
American Coleoptera. Psyche, 7, I (A.-G), pp. 160-162; 
II (H.-P), pp. 188-193; III (R.-Z.), pp. 205-209. (Con- 
tains an index of localities.) 
Hamilton, J. 

1894. Catalogue of the Coleoptera of Alaska, with the Synonymy and 

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1894a, Catalogue of the Coleoptera Common to 'North America, 
Northern Asia and Enropc, with Distribution and Biblio- 
graphv. Second edition. Trans. Am. Ent. Soc, 21, pp. 
345-416'. (First edition I. c, '89, 16, pp. 88-162.) 

1895. Catalogue of the Coleoptera of Southwestern Pennsylvajiia, 

with Notes and Descriptions. Trans. Am. Ent. Soc., 22, pp. 
317-381. 
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1884. List of Ottawa Coleoptera. Trans. Ottawa Field Natur. Club, 

No. 5, pp. 67-85. 
Hayward, R. 

1897. On the Species of Bembidium of America, North of Mexico, 
Trans. Am. Ent. Soc., 24, pp. 32-143. 
Henshaw, S. i 

1885. IJst of the Coleoptera of Ameri^-a, North of Mexico. Amer. 

Ent, Soc, 
1895. Third Supplement to the List of Coleoptera of America, North 
of Mexico. Amer, Ent, Soc. 
Horn, G. H. 

1871. Revision of the Tenebrionidae of America. Trans, Amer. Pbil. 

Soc, N. S. 14, pp. 'lotiiOi. 

1872. Coleoptera- Fifth Ann, Rep. U. 6. Geo!. Surv, Montana and Por- 

tions of Adjacent Territories (Hayden), pp. 382-392. 
Hubbard, H. G., and Schwan:, E. A. 

1878, Coleoptera of Michigan. Pro. Amer, Phil. Soc., 17, pp. 593-669. 
Keen, J. H. 

1895. List of Coleopterai collected at Massett, Queen Charlotte 
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Klages, H. 0. 

1901. Supplement to Dr. John Hamilton's List of the Coleoptera of 

Southwestern Pennsvlvania, .\nn. Carnegie Museum, I, uii„ 

265-294. "^ 

LeConte, J. L. 

1^0, General Remarks upon the Coleoptera of Lake Superior, pp. 



214 MICHIGAN SURVEY. 190«. 

1851. On tbe (ieojp-utihical DiHtribution of AtiimaU in Californiii. 
Pi-o. Ain. Assoc. Adv. Si-i., 6, pp. 248-254. 

1859. TTie Coleoptera of Kansas and Eastern New Mexico. 8mitb. 

Cont. Kuowl., 11, pp. 1-58. 

1860. Report upon Insects Collected on the Survey. Rep. Ezpl. and 

Snrv. from Miss. River to the Pacific Ocean, 12, Pt. 3, pp. 

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18ti2. Notes on the Coleopterous Fauna of Lower California, Pro. 

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187(i. Presidential Address. Pro. Am, Assoc. Adv. Sci., 1875, pp. 

1-18. (Also Pop. Sci. Mo., 8, pp. 285-299; Amer. Nat., !), 

pp. 481-498.) 

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Pro. Am. Phil. Soc, 17, pp. 470-471, 
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1879. The Coleoptera of the Alpine Rocky Mountain Regions — Part 

II. Bull. U. S. Geol. Sui-vey, Terr., 5, pp. 499-520. 
MacGillivray, A, D. 

1903. Aquatic Chrvsonielidac and a Table of the Families of Coleop- 
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1870, On the Geographical Relations of the Chief Coleopterous 
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Riley, C. V. 

1880. Food Habits of the T»ngicorn Beetles or Wood Borers. Amer. 

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1878. The Coleoptera of Florida. Pro. Am. Phil. Soc, 17, pp. 353- 
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1890a. On a Collection of Coleoptera from St. Augustine, Florida. 
Pro. Ent. Soc. Wash.. 1, pp. 169-171, 

1901. Semi-ti-opical Texas. Pro. Ent. Soc. Wash., 4, pp. 1-3. 
Scudder, S. H, 

1874. The Distribution of Insects in New Hampshire. Final Report 
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1894. The Effects of Glaciatlon on the North American Fauna. Am. 

Jour, Sci,, 48. p. 180. 

1895. TTie Coleoptera Hitherto Found Fossil in Canada. Cont, to 
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1900. Additions to the ('oleopterouR Fauna of the Infer^Iacial Clays 
of the Toronto District, With an Appendix by A. D. Hopkins 



ECOLOGY OF ISLE HOYALE. 215 

OD the Scolytid Borings from tbe Same Deposits. Coot. 

to Can. Paleon. II, Pt. II, pp. 67-92. 
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1882. On the Aqnatic CarnivoroiiK Col«>ptera or Dytiwidac. Soi. 

Trans. Gov. Dublin Noc, (2>, 2, pp. 17fi-10<Kt. 
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1907. Prelimiunrv Note on the Distribntiou of the Tiger Beetle (Cin- 

cindela) and its Belation to Plant eucceneion. Biol. Bull. 

14. pp. 9-14. 
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1900. Insects of New Jersev. Snpp. Twentv-seveuth Ann. Rep. N. 

J. St. Bd. Agr., 1899. pp. 107-S67. 
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1874. Catalf^ie of the Coleoptei-a from the Region of Lake Pontehar- 
train, La. Bui!. Bnffalo Soc. Nat. Sci-, 5, ,pp. 78-09. 
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1906. An Investigation of Evolution in the Chrysomelid Beetles of 
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1902. A Ust of the Beetles of the District of Columbia. Pro. U. S. 

Nat. Mus., 25,' pp. 1-57. No. 1275. 
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1901. Observations upon the Faunal Relations of California from the 

Standpoint of a Coleopterist. Jonr. N- Y. Ent. Soc, 9, 

pp. 197-199. 
Wickham, H, F. 

1893. Report on an Entomological Rcconnaisanoe of Southern 

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perior, With Remarks on Geographical Distribution, Pro. 

Davenport Acad. Sci., (i. [jp. 125169, 
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, New Mexico and Arizona. Bull. Lab. Nat. His, Univ. Iowa. 

3, pp. 15;!-171. 
1897a. The Coleoptera of the Lower Rio Grande Vailey. Bull. Lab. 

Nat. His. Univ. Iowa, 4, pp. 96-115, 

1898. Tile Beetles of Southern Arizona, Bull. Lab. Nat. His., Univ. 

Iowa, 4, pp. 295-312. 

1902. A Catalogue of the Coleoptera of Colorado. Bull. Sci. I^b. 

Nat, His, Univ. of Iowa, .^, pp. 217-310. 

1904. The Influence of the Mutations of the Pleistocene I^kes upon 

the Present Distribution of Cicindela. .Vmer. Nat., 38. pp. 
643-654, 

1905. Insect Distribution in the Great Basin Considered in tbe I^ight 

of its <;eological Historv. Thirtv-flfth .\nn. Rep. Ent. Soc. 
Ont. pp. 42-46. 



jiGoogle 



BCOLOOT OP ISLE ROYALB. 



NOTES ON THE VEGETATION OF ISLE ROYALE, HICHIGAN. 

W. p. HOLT, CENTRAL. HIGH SCHOOL, TOLBIK), OHIO. 

/. Qeneral Oltservations on the Plant Societies. 

Situated in tlie northern part of Lake Superior, in ni^ht of the 
Canada shore, and bisected by the parallel of 43° N., ThIp Royale offers 
a most attractiire field for summer work. That its Horn is strikingly 
northern may be inferred from its proximity to the Cana^hn shore, as 
well as by the many species of northern plants include*! in the anno- 
tated list of plants. 

The island, 45 miles long and containing about 210 square miles, has 
had almost no permanent population since the "copper days;" and only 
a few localities along the water's edge are frequented by summer visitors, 
thus leaving the island largely free fromi man's occupancy. With the 
exception of the building and mining improvements of the Wendigo 
Mining Company, at the bead of Washington Harbor, little remains on 
the island to mark the vanished population, except the burnings and 
clearings, which are easily recognized by tbeir characteristic floras. 

Large parts of the island, however, have remained quite free from 
man's invasion. That the present natural conditions are not likely to 
remain long undisturbed, and that the past Bummer's observations and 
records were made none too soon, is shown by the fact that contemporane- 
ous with the work of the Museum party there were at least three differ- 
ent parties of timber estimators working over large parts of the island . 
looking toward the cutting off of the forests. 

The general plan of the summer's work on the biota of the island was 
to select the most typical and representative parts as general stations, 
wolfing these through sub-stations as carefully as time would allow, 
and comparing other similar localities with these. 

The principal plant societies of Isle Boyale may be considered under 
four heads, viz.: B(^ societies, shore societies, forests, and bumingSi. 

I. Bog Societies. It is doubtful if there could be found anywhere 
in an area of the same size a more interesting and more complete series 
of bogs than occurs ou Isle Royale. On the geologic^ map of the 
island, by Lane and Stoekly, there are shown over 100 smaller bog 
areas, exclusive of the extensive bog region in the southwest part of the 
island, to the west of Siskowit Bay. Add to this the various stages 
of partly filled takes, and there is shown almost every conceivable stage 
in the life history of bogs, from the open tarns, or lakes, to the climatic 
bog forest. 

It is unnecessary to call attention to the very interesting manner in 
which the bog floras respond to the various stages of physical changes; 
such responses of vegetation in any physi<^aphic series are too well 
known to need comment. For any student who desires to work out in 
detail these stages of successions as carefully and minutely as Cowles 
has done for the dunes of the Chicago region, Isle Royale presents all 



218 MICHIGAN SURVEY. 1908. 

that conid be desired; and one important advantage here is that most 
of the bog areas are comparatively small, and all in a reasonably limited 
area, thus offering an unusaal opportunity for their comparative study. 

There is, on the other hand, one difficulty that should be mentioned 
in connection with any kind of fleld work on Isle Hoyale which takes 
one far back from the shore, and that is the difficulty of penetrating 
the dense tangles of the forest. The absence of roads, the limited num- 
ber even of old blazed trails, the unusually dense underbrush, includ- 
ing a very rank growth of Tawu-a minor (Ground Hemlock) and Dumer- 
oua windfalls, together with the necessity of carrying by pack one's 
supplies — all render the penetration of many parts of the island a mat- 
ter of such difficulty that it has been remarked by all who have at- 
tempted it. 

In the limited time at our disposal during the summer It was im- 
possible to visit all of the 100 or more bc^ areas on the island : our 
attention was therefore confined to^a limited number of those which 
are typical of a certain stage of development, or to those having in- 
dividual points of special interest. 

Three general stages of the lake-bog series will be briefly touched upon, 
(1) the open lake withovt marginal vegetation, (2) the partly open 
lake icith marginal vegetation of varying width, (3) the wholly carpeted 
b<^ area ; the vegetative carpet in some cases being recent enough to give 
beneath the feet, in other cases old and solid enough to be more or less 
forested. 

The first, or open stage, includes only a few of the lai^est lakes of 
Isle Royale, such as Lake Siskowlt and Sargent Lake. Of these Lake 
Siskowit is hy far the largest, being at least a mile and one-half broad 
in places. The principal reasons for the absence of vegetation in the 
lakes of the first class seems to be that their size and openings renders 
the sweep of the wind and the resulting wave action so vigorous thsft 
even annuals cannot get a foothold along their shores. 

Wave action in a few places is clearly marked by a narrow but well- 
defined beach, as along the north shore of Lake Biskowit. 

Another factor that has to be reckoned with in the larger lakes of 
the first class is the work of ice. Ice destroys shore vegetation in two 
ways, first — hy pushing, due to expansion by freezing (and this total 
expansion in a lake as large as Lake Hiskowit is considerable) ; seconil 
— the open expanse allows large ice floes to blow ashore during the 
spring breakup. Tlie most interesting example of ice pushing noted 
was along tht north shore of Lake Siskowit, where there is an irre- 
gular ridge, varying in height up to 15 feet and composed of bowlders 
and various fragiiiental materials. Along this ridge there were, in 
places, even overturned trees of considerable size, pointing away from 
the lake, back 20 to 25 feet from the present shore. This ridge seems 
certainly to be the work of ice as in the case of the so-called "bowlder 
rim" lakes of the western United States, or the ice floe ridges at Put- 
in-ltay in Lake Erie. 

In drawing a line between the lakes that will long continue to remain 
free from the encroachment of vegetation and those which are being 
gradually captured by vegetation, the size and openness seem to be the 
most important factors, affecting the vigor of wave action as well as 
the work of ice in one or both of the ways suggested. 



ECOLOGY OP ISLE ROYALE. 219 

In the smaller lakes, espeoiallr those nestled in depressions, the 
surrounding forests protect their surfaces from vigorous wind action 
BO that there are practically no waves at all to check the encroachment 
of regetRtion along the lake margins. 

That the slope of the shore, in the ense of protected lakes, has much 
to do with retarding or aBsisting the encroachments of plants is self 
evident. The Isle Boyale lakes of the protected class show numerous 
examples where the plant zone is much farther advanced on the gently 
sloping side than on the opposite one with a more abrupt slope. 

A typical example of a lake midway in the process of capture is 
Sumner Lake. This lake, which is roughly one-half mile long and one- 
third as wide, with its long axis nearly east and west, has already 
been captured at its east and west ends. Had its north and south 
borders been less steep the entire lake would doubtless have been 
covered ere'this. This lake has an outlet into Conglomerate Bay, bat 
at its west end it receives a small creek. The west end is covered by 
a bog carpet still so young and elastic as to render the crossing of it 
diflBcult. Along the more abrupt sides, and connecting the bog carpet 
at the ends, was a narrow, irregular zone of CaJla palaatris and Iria 
versicolor, with the Mcnyanthes (lEuckbean) and Comarv/m palustrc 
(Marsh cinquefoil) mixed in places. Parts of this zone, where the shore 
is less steep, were closely backed up by willows, Comns stoUmifera, and 
Alitu« inmna. thus giving to the marginal zone the aspert of a swamp 
rather than of a bog. Growing on the wet bog carpet at the ends we're 
the Sarracenta purpurea, Droaera rotundi folia. Drosera intermedia. Men- 
yanthea trifoliata, Comarum palvstre, Droacra linearis (the latter two 
in wetter places generally than the former), Oxycoccus oxycoccva, 
Hahcnaria pai/codrs, Hahcnaria diJatnta, Pogonia ophioglossoidea, Vtri- 
cularia minor (wetter parts), Campauula oparinoidea, Scutellaria galer- 
icuJa, Cicvta bulbifcra, Triadenum- t-irginicum, Pamassta pahistris, 
Sotidago neglecta, etc. 

A word in passing in regard to the "false bottom" of Sumner Lake, 
for in no place on the island was this better shown. In paddling aronnd 
the open part of this Iflke on a raft it appeared in places that the water 
was only 6-10 feet deep. This was a matter of surprise since even a 
raft's length from the shore we could not touch bottom with our 15 foot 
pole. Further investigation showed a "false bottom" in various parts 
of this lake. This was composed of the fine, disintegrated remains of 
leaves and other light organic material. In places there were great 
breaks in this "false bottom," doubtless due to the escape of gases 
which has lifted this flue, ooze-like material from a greater depth; and 
through these breaks one could look down several feet through the . 
brownish colored water. While this "false bottom" was so tenuous 
that a pole could be thrust through it almost as easily aa through the 
clear water, it seemed to play an important part in the distribution 
of patches of CastaHa odorata (White Pond Lily) so abundant on the 
surface of the lake, and also served to call attention to the manner in 
which this material assists in lake filling. 

An area illustrating the final stage of bog covering was examined 
at the end of the cabin trait from our Kiskowit Ray camp. In this 
sphagnum hog (V, 5). containing 80-100 acres, all lias been covered 



220 MICHIGAN SURVEY, 190S. 

except an area of open water about 60 feet long and 'balf as wide 
surrounded by an exceedingly wet, unstable margin. 

A few years hence and even this will be covered. The main part of 
the bog was covered with sphagnum hummocks, upon which were grow- 
ing Ledum ffroenlandicum, Chamacdaphne calyculata and Andromeda 
poUfolia in dense patches. Young Tamaracks and Black Spruces were 
pushing out from the older parts of the margin, with Balsam Firs close 
behind. 

Along the south margin of this bog, in the tension zone between the 
bog and the adjacent forest, there was being waged one of the most 
intense and most interesting struggles for plant supremacy that we 
have ever seen. Working up the gentle slope from the bog margin the 
sphaguum invasion (after the manner of a large snowdrift) was push- 
ing out its lobate fingers, over the forest carpet of leaves; and during 
a single season by its rapid growth had surrounded sucb plants of 
the forest as Aralia nudieaitlis, Trientalis americana, Clintonia borealis. 
Lycopodium lucidulum, all of which were completely helpless in the 
path of the sphagnum invasion. Even lai^e, fallen trees were able 
to check its advance only temporarily, for instances were noted where 
entire fallen trunks were covered, only the upward projecting branches 
being out of reach of the Sphaguum. In a dry carpet of forest leaves 
the clean-cut forward margin of the sphagnum was so wet that water 
could be wrung from it at a distance of 15-20 feet from the original 
bog margin, thus showing how readily water is transferred through the 
sphagnum patches, even up a slope. 

While the sphagnum invasion was eminently successful against all 
the scattered plants of the woods there was at "least one species of 
moss {Polytrichum cmnmtine) growing in dense formations which was 
succe-fuful in holding the sphagnum in check. The moss colonies were 
so dense that the sphagnum could not penetrate them; on the other 
hand the moss was actually invading the moist sphagnum and growing 
over it. 

Before leaving the semi-enclosed lake bogs a few questions surest 
themselves regarding the trembling bog carpet adjacent to the water's 
edge. What is the thickness of this elastic, quaking water cover which 
is, at the same time, strong enough to enable one to walk out to with- 
in a single step (in some cases) of the water's edge? Also of what is 
it composed? In all the measurementfl taken it was found that this 
vegetative cover, within two to three feet of the water's edge, had a 
thickness varying from 22-24 inches. Back from this younger and more 
unstable margin the bog cover becomes thicker and firmer. In one bog, 
back about 100 yards from the water's edge, where the surface was 
firm and unyielding, the boring pole broke through into open water at 
a depth of 5 feet 6 inches. In another instance, at the west end of 
Wninner Lake, at a distance of over 100 yards back from the water's 
edge, I found the bog cover still so thin and trembling that I broke 
through in one place in attempting to walk across it, and anticipated 
that the same might happen in several other places. These and other 
instances all go to show that no definite statement can be made as to 
the exact distance from the water's edge at which the bog cover be- 
comes thick enough to support one. ,This may vary with the depth 
of water underneath, as well as the distance from the original shore 



ECOLOGY OF ISLE ROYALE:. 221 

It ia to be regretted tbat more borings and measurements coald not 
bave been taken in the limited time at our disposal. A summer spent 
with suitable borinj; tools in malting an extended series of borings over 
various parts of several of the Isle Royale bogs and boi;-lake margins 
would doubtless bring to light some very interesting data. 

Now as to what gives strength to the bc^ cover. Kince the Rphagnum 
is so predominaut on bog areas, covering large parts of the surface, 
and often extending out uluioKt to the water's edge, oue is apt to think 
only of the sphagnum surface and fail to consider the important net- 
work below that gives such strength to the trembling bog carpet ont 
almoKt to the very water's edge. It iw scarcely necessarj' to add that 
the delicate sphagnum alone i.i not sufficient to make a strong bog 
cover. The weakness of the individual sphagnum plants to resist strain, 
the lack of interlacing parts, or of even ''felting" properties are clearly 
shown in that one can reach down a foot or more into the loose, soft, 
sphagnum and pull out a handful^ of it without seriously disturbing 
the adjacent plants. Moreover, the Nphagnuni does not grow along the 
water's edge in advance of its supporting mat — at least we failed to 
find a single instance of this on Isle Boyale->-wbiIe in many cases it 
did not extend out to within several feet of the water edge of the 
supporting mat. 

0^ pulling up large masses of the floating mat at the water's edge 
it was found to consist of a dense tangle, or network, of tough fibrona 
roots and rhizomes of KcdgeM, Mi-nyanth<:n trifoliata and Comarum pal- 
ustre, all so tightly interlaced that it was very dilBcult to separate any 
part of the tangle from the rest. Such tou^ parts are in striking 
contrast to the delicate sphagnum, as is also the manner of growth;. 
and furnish the platform on which the sphagnum works out toward the 
lake margin. 

In the last, or wholly covered, division of bogs a wet and a drier stage 
may be reci^nized; the former may be cbaractwized by the Sarracenia 
purpuria, Menj/anthes trifoliata, Comarum palwstre, and one or more 
species of Droaera. Rpbagnum hummocks may occur in both of these 
covered stages, or the surface may be comparatively smooth. These hum- 
mocks, of varying size up to 4 feet in height, seem to be due in most 
cases to the sphagnum growing up around tree trunks, shrubs, or other 
objects. Instances were noted of where the rapidly growing sphagnnm 
had so nearly covered the Ledum groenlandicum that only the ends of 
the upper branches were to be seen. It is possible, however, that some 
of the hummocks may be formed in other ways, e. g. one large hummock 
was noted that was inhabited by ants. This suggested that possibly the 
sphagnum had overgrown a large ant mound, although it is also possible 
that the ants may have inhabited the mound only after its formation in 
some other way. 

The pioneer trees to apjMiar in Isle Royale bogs are the /rfin> laririmt 
jTamarack),and Picea mariana (Black Sprnce) , which appear simultan- 
eously, and seem equally well adapted to bog conditions. Owing to the 
advance of the bog cover from the margin toward the centre, one nat- 
urally expects to And the youngest trees farthest in, and this is utrik. 
ingly well illustrated in many of the bogs. , 

By counting the rings of trees cut in the bogs, and comparing with 
the same species just outside, it was found tbat the growth of those in 



222 MICHIGAN SURVEY. 1908. 

the bogs was strikingly slower; The rings of the bog species were in 
many cases so close together as to render a band lens desirable for 
cooating them, while the annnal rings of the same species in the adja- 
cent forest were widely separated. 

2. Shore Vegetation. The work done on shore forms was confined 
entirely to the south shore, including the group of small islands near 
the abandoned Light-hoase at Buck Harbor. The northern shore is steep 
and cliff-like, the southern shore gently sloping. While the northern 
shore is strikingly different from the southern, and might have hronght 
to light many interesting things (especially in the way of lichen forma- 
tions), it seemed best to confine the limited time at our disposal to work 
on the south shore. 

Of the special shore forms, the crevice plants are both interesting 
and attractive. The crevices in most cases are due to fissuring, altbongh 
some long, narrow grooves were made by the differential weathering of 
the softer vein rock. The bed-rock of the shore is often amygdaloidal, 
and many small depressions in this, due to the more rapid weathering, 
afford a foothold for the hardy plants of the rock shore. In their narrow 
rock crevices and confines, with little soil, and on dark-colored rock which 
in summer becomes highly heated, at all times exposed to the strong 
lake winds, and in winter often washed by the powerful storm waves of 
I^ke Superior, their struggle for existence is certainly a most strenuous 
one. On the whole their size and appearance is strikingly alpine, as 
is also their coloring in many cases. 

A partial list of the crevice plants is as follows: Campanula rotun- 
difolia, Potentilla trideatata, PotentiUa littoralis, Saxifraga tricunpi- 
data, Saxifraga aizoon, Saxifraga nivalis, Artemisia canadensis, Senecio 
dalsamitae, Primula miatasainica, Solidago virgaurea ( ?) , Sagina sag- 
inoidea, Achillea millefolium, Aster ptarmacoides, Sisifmbrium humile. 
Lobelia kalmii, Nabahia racemosa, etc. The insectivorous Pinguicuia 
vulgaris (Butterwort) occurs in rock pools and on wet rocks along the 
rocky shores. 

The most common crevice shrubs were the Juniperua nana, Junipcrua 
procumbena. Arctostaphylos uva^irai (Bear-berry) , Shepherdia cana- 
densis (Shepherdin), Opulaater opuUfoliua fNinebark). On the exposed 
rocks at Scovill Point and at the eastern end of the island, the Empctrvm 
nigrum (Crowberrj) was also found, forming a part of the heath mat. 
Of all these the ./. procumbena is easily of the greatest importance in 
preparing the way for other larger forms of plants. Certainly no shrub 
of Isle Royale precedes it or has better claims for pioneer distinction. 
Its hardiness, prostrate manner of - growth, and its thick, sheltering 
branches are all of great importajice in making it an excellent pioneer. 

A study of the small rock islands' near the Bock Harbor light-house 
was very interesting not only on account of the striking differences in 
the individual flora of each, but also for a comparative study of the plant 
successions upon them. All stages of successions were noted from an 
unusually rich mesophytic flora, growing on a humus soil 3-10 inches 
in depth, down to islands almost bare except for a few crevice plantn. 
In general ail the islands showed a less development of the flora on the 
side exposed to the open lake than on the more protected land side: in 
some cases the vegetation of the two sides was strikingly different. 

The advantages of crevices in enabling vegetation to get a start upon 



BCOLOOT OF ISLE ROYALB. 223 

bare islands was well illnBtrated in the caee ot one of the email ielands 
of this group. Ita smooth, sloping surface was bare except for a few 
small patches of crustaceous licbens and a single large procumbent 
juniper. The juniper -was growing in a crevice along which it had 
reached for several feet in either direction, occasionally rooting along 
the crevice which held it more securely in place. The spaces between its 
dense sheltering branches were filled with a vigorous growth of moss 
which doubtless started on the wind-blown material that had lodged 
there. So solidly had the moss filled the spaces between the branches 
where it was growing that in breaking off a portion of the juniper every- 
thing was stripped oS down to the bed-rock. As such a juniper patch 
spreads, and the humus made by the moss increases, other plants come 
to grow on the juniper patch, and an ever-increasing heath mat is formed. 
Other similar crevices may, in like manner, spread to join this, and in a 
comparatively short time the entire surface is carpeted with vegetation. 
On other small islands crevice trees and shrubs have contributed shade 
and partial protection from the wind, and the process has gone on even 
more rapidly. Had there been ito crevices to enable these higher pioneer 
plants to secure a foothold the process of vegetative capture would have 
gone on infinitely slower. What the possible steps are in such a case 
may best be considered in connection with the rock shore-heath-forest 
series to be referred to presently. 

To suggest the severe and varied conditions of the exposed rock shore 
the following is cited. On a bare, gently sloping {10°-12'') portion of 
the rock shore near Rock Harbor, there were, in an area approximately 
40 fteet square, over 100 fresh scars where the thin (1-6 to 1-8 inch^ 
patches of rock had recently been broken off. These patches varied in 
size from 12 inches in diameter down: some were covered in part by 
licbens (principally Parmelias), others were entirely bare. The intense 
daily heating and expansion to which the immediate surface of the dark 
colored, exposed rocks is subjected, together with the rapid cooling and 
the resulting contraction at night, doubtless has much to do with weak- 
ening the immediate surface, and starting the chipping. The freezing 
of moisture in the rock surface may have been responsible for the final 
breaking away and lifting. 

For a brief survey of the vegetation from the water's edge back through 
the heath zone to the forest at the top of the gently sloping rock shore 
area, V 2, (designated as "the heath zone and beach" of Siakowit Bay) 
will be selected as a typical locality, and supplemented by additional 
observations on similar places elsewhere along the southern shore. The 
IKtrtion of rock shore to be considered has a rather uniform slope of 
about 10° and a width of 200-250 feet from the water's edge back to the 
forest at its summit. 

The first zone of no vegetation extends back about 20-25 feet from the 
water's edge, although the winter waves doubtless reach far beyond this. 
Back of this occur, in turn, the crustaceous and foliaceous licheu zones, 
which meet in a somewhat irregular tension zone that can, nevertheless, 
be distinguished by looking up or down the shore. The lichens of these 
zones are included in the annotated list, and will not be enumerated here. 

Numerous crevice plants (as already listed under shore forms) make 
their appearance in the crustaceous and foliaceous lichen zones, also 
Thuja occidentalia and Picea oatiaderma, the former being the hardier 



224 MICHIGAN SUBVEY, 1908. 

pioDeer of the two. In parts of the upper, or third, lichen zone there 
are unnanally denne and luxuriant formations of CladoniaB, often 50-60 
feet across. Kcattered among the Cladonias were Junipertts nana, J. 
procumbcm, Arctostaphylon uea-ursi, and -Vaccinium pennsglvanicum. 
The upper, op back iiortion, of this zone will be designated as the 
Cladonia-beatb zone, for it is here that the real shore heath begins. The 
baelt of the heath zone contains numerous young Balsam Firs and White 
Birches which have worked in irrt^ularl.v from the adjacent forest. 

\ similar sloping rock shore near our Siskowit cabin camp (V, 4) 
showed some interesting later stages. The shore was here better pro- 
tected from waves and wind by the flat neighboring wooded islands; and 
the forest development had gone on more rapidly, having extended irreg- 
ularly from the higher shore down to the very water's edge sug- 
gesting the ultimate condition elsewhere along the less "protected rock 
shore. There were still open places, NUggesfing the irregular manner 
in which the trees had pushed out to take possession of the lower shore; 
but the forest was here far better established (seemingly on account of 
the better protection) than elsewhere along the beach where exposed 
directly to the lake winds and waves. 

As the trees increase in number, and afford better conditions of shade 
and moisture, vigorous mosses and wood plants begin to invade the 
<'ladonia patches still occnpying the more open places. A series of 
jrhotographs was taken showing various stages of this invasion of 
uuder-growth wood plants, from a pure formation of Cladonias to a 
climax of a dense society of wood plants with not a vestige of Cladonias- 
remaining. These later back shore formations were equally well shown 
along the heath forest tension zone at Kock Harbor. 

If carefully worked out the i-ock shore series, from the water's edge 
Imck to the neighboring foi-est, might be made to rival in interest the 
liike-bog-forest series, so deserving of more cai-eful study on Isle Royale. 

.'t. ForeHta. The forests of Isle Royale include about 21 species of 
ti-ees, 13 of which ar^ deciduous, the remainder evergreen conifers. The 
paucity of species has been more than offset by a generous distribntion 
and abundance, for the island as a whole is heavily forested. The largest 
and dominant trees of Ihe i)resent forest are Abies balaamea (Balsam 
Fir), lictuia papyrifern fWhite, op Canoe Birch), and Plcea vatmden»it 
(White Hpruce), with the exception of the western end of Greenstone 
ridge where Acer sacvharum (Hard Maple), Bctula lutea (Yellow 
Binh), and Bctula htita (Black, or Chepry Birch) are dominant. 

Between the end of Washington Harbor and Lake Desor there are 
places where almost pure stands of Hard Maple and birches obtain. 
The scarcity of Abii-^ balnniin'a hei-e. which is so abundant ovep almost 
all other parts of the island, is an interesting matter of speculatiou. 
Young Balsam Fire wei'e noted growing in the shade and shelter of 
the maple groves, and they appeared to be vigorous and thriving, yet 
scarcely a large fir could be found associated with the maples in this 
jmrt of the (Si-eenstone ridge. An examination of the soil here showed 
that it is only 4-6 inches deep. This, together with the laterally 
limited root system exhibited by the larger overturned firs, seemed to 
suggest that, after attaining a certain height and rigidity, they be- 
came BufBciently exposed to be overtamed by the powerful winds that 
sweep that exposed part of the Greenstone ridge. r- [ 



ECOLOGY OP ISLE ROTALE. 225 

The forested bog areas are characterized by the domiDance of Tam- 
arack, Black Spruce and White Cedar. As a rale, where the Tamarack 
is more abundant the White Cedar is less abundant, and tbe opposite. 
Where the White Cedar is dominant (ae it is in many bog areas, tbe 
largest trees being 2 — 314 f«et in diameter), the few Tamaracks pres- 
ent are lai^ and appear as pioneer relicts. The White Cedar, more- 
over, does not appear with the Black Spruce and the Tamarack in the 
earliest, wetter stages; but seems to come in only when a drier con- 
dition has been reached. 

It may also be added that none of these characteristic bog trees are 
here eo closely confined to their bog habitats as to the south of here; 
but they have a much more general distribution. The Black Spruce, 
for example, one of the earliest pioneers of the bogs, occnrs sparingly 
dietribated in the original forests along with the White Cedar and 
Balsam Fir; and I have also noted it growing on dry exposed rocks 
where very little soil was present. The Tamaracks also get out of the 
bogs and occur sparingly distributed in the npland forests — sometimes 
in most unexpected places. 

Of all the island conifers the Abies bahamea is easily the most com- 
mon, and seems to be superseding the sprures and tamaracks. Tbe 
young seedlinf^ of it grow in dense shade, as well as in more open 
places. Seedlings of the Balsam Fir come np abundantly under the 
White Spruces in place of the seedlings of that species which do not 
seem to be able to endure the shade of the dense forest. It will doubt- 
less form an important part of the climatic forest of the island. 

The Picec eana4ensis is fairly common along tbe margins of forests, 
and in the more open parts — even in the deeper parts of tbe forest — 
when it has come in as a pioneer with firs and other, conifers of the 
present generation; but the White Spruces do not seem likely to suc- 
ceed themselves and become a considerable part of the dominant forest, 
on account of the inability of their seedlings to withstand deep sbade. 

4. Burnings. The burnings and old clearings are everywhere char- 
acterized by an abundance of PopuJus trcmuloidea and Betula papyri- 
fera. while the undergrowth consists largely of Diervilla diercUJa. Aster 
macrophylla, Cha^naenerion anguatifoUum, Hubus parviftorum, Comu» 
canadensis, and in places an abundance of TaxuK minor. Burnings of 
different periods were suggested by uniform stands of Quaking Aspens 
and W'hite Birches which were of difFerent heights. 

Tbe Pennsylvania Cherry {Primus pennaylvanicus) occurs in burned 
areas and elsewhere where there is little soil, sometimes growing out 
of the crevices of exposed rocks where the conditions of growth were 
strikingly unfavorable. Perhaps no other tree on Isle Boyale can 
withstand more xerophytic inland conditions, with the possible excep- 
tion of the Jack Pine {Pinns divaricata) which was occasionally found 
associated with it on high exposed ledges. In one locality where the 
two were growing in close company — a high rocky ledge near Conglom- 
erate Bay (III, 5) — there was an almost total absence of soil, dne 
to its removal to lower altitudes by wind and rain; there was a strik- 
ing range of SO^-TO" F. in the daily temperature, and a complete ex- 
posure to the powerful Lake Superior winds which overturn so many 
trees when they had developed enough heart wood to become rigid and 



226 MICHIGAN SURVBT, 1908. 

resisting. There might be added to the unfavorable conditions of 
growth on such exposed ridgea the work of Hares, for the Northern 
Varying Hare often resorts to the expoaed heights — as in the case just 
noted — for its winter feeding ground, since there is probably less snow 
left there by the sweeping winter winds than at lesser elevations, and 
the Hares can get about more easily. The principal damage done to 
the trees by Hares consists of the cutting off of the yonng branches, 
and gnawing the bark, and this in some cases amounts to considerable. 

Scattered individuals of White Pine {Pinu* atrohus's occur along 
the ridges and on the north side of Siskowlt Lake, hut it is nowhere 
abundant. Only a few Norway Pines {Pimus resinoaa) were noted — 
these occurring principally on ridges and in exposed places, as on the 
ridge north of Sumner Lake (III, 5). 

The Qrecn, or MonntainAlder (Alnua alnobctula) was widely dis- 
tributed on higher ground, and can seemingly stand as much shade as 
any broad leaved teee on the island. The Speckled Alder {AUtua 
incuna) was common near the water's edge and on low ground ; and is 
places had worked back some distance from the water. 

The Sorbus americana, found on many parts of the island, was in 
moat cases solitary in its distribution and nowhere abundant.^ It ap- 
pears to be most abundant along the water's edge. 

The successions of the burnings and clearings due to the attempts of 
the early copper prospectors to clear the land, as well as the results 
of later forest flres, present an interesting problem; also the peculiar 
distribution of the Hard Maple and White Pine on the island. Students 
of fleshy fungi may also find a most fascinating field for later summer 
work at the west end of the island, especially along the forest road 
from the Washington Club grounds to Lake Desor. Never have we 
seen a more inviting field for mycologists; and in a region as yet un- 
touched as to its fungi. 

In conclusion, we beg to call attention to the fact that, owing to 
time limitations, no attempt was made to work out in detail any of the 
Lirge and interesting problems that presented themselves; our object 
being rather to make a general reconnaisance of the plant life on as 
many different parts of the island as possible. It is to be regretted 
that time did not permit the party to investigate several habitats on 
the north side of the island in addition to the work done on the south 
side, for the physiographic conditions there are different from those 
on the south shore, and a comparison of the environmental conditiona 
of the two localities would doubtless throw additional I^bt upon the 
teries of shore societies. 

To give a more detailed account of the plants noted and collected on 
the island than could be attempted in this resume of conditions the 
following annotated list is herewith presented. 

The -writer is indebted to Dr. C. A. Davis of the Michigan Biological 
Survey for the determination of the sedges and certain flowering 
plants; to Prof. Bruce Fink of Miami College, Oxford, Ohio, for the 
d4 termination of the lichens; for the determination of the momes to 
Prof. J. M. Holzinger of Winona Normal School, Winona, Minn., and 
Dr. J. Boll, Germany. • 

The nomenclature is that of Britton and Brown's Illustrated Floro of 

iX.ooglc 



ECOLOGY OF ISLE ROTALE. ■ 227 

tbe Northern States and Canada, 189S. The report of the expedition 
for 1904 followed the nomenclature of Britton'a Manual of the Flora 
ot Northern United Btatea and Canada, IdOl. 

II. Annotated List of Plants. 

lAchens. 

Dy lichen zone No. 1 in meant to inclnde the crustaceovs lichens; 
liiese forming Bmall patches on the rocke as in Placodium elegans, in 
which the thnllue is principally imbedded in the rock bo closely that 
I he rock must be broken away to secure them. Zone No. 2 (Foliose 
zone) includes the flat thallus species which are attached by rhizoids, 
as ParincUa. These can he scraped or pnlled off. Zone Xo. 3 (Fruiti- 
cose zone) includes the upright lichens like Cladonia. 

1. Ramalhia calicaria farinacea (C) Pr. Vertical rock cliffs at 
water's elge. Principal branching lichen on vertical cliffs. Common 
8ta. I, Sub. 1. 

2. Cctraria larunosa .\ch. On nearly bare surface of rocks Sis- 
bowit Bay, (V. 2). Occasional. 

3. Evemia prKrmatria (C) Arch. A light green, branching form 
growing in Cladonia zone (I, 1). 

4. ilsnea barbata'cerotina (Ach.) Schaer. Hanging from trees along 
Siskowit cabin trail (V, 4), also on trees at Rock Harbor (I, 3). Com- 
(tion. 

5. Usnea longissima Ach. A pendulous form 12-15 inches or more 
in length. Occurs on conifers. T^sb common than preceding species. 

B, Parmdia perlata (Jacq.) Ach. Top of cliff at Bock Harbor ([, 
H). Sot widely distributed. 

7. Parmelia saxttalis sulcata Taiyl. A gray foliose form growing on 
very thin black humus on sloping tops of cliffs. Noted to be abundant 
top of rock cliff at Bock Harbor (I, 1). 

8. Pmynelia caperata (C) Ach. On rocks of foliose lichen zone at 
Bock Harbor, I, 1; V, 2. 

9. ParmeUa conspersa Ehr. One of the most common lichens on 
the island, and the principal form in the second (foliose) lichen zone of 
the sloping rock shore. Also fairly abundant on rock surfaces back 
fi-om the shore. I. 1; V, 2; V, 3; I, 2. 

10. Phpscia pulverulenta (Scrieb.) Nyl. Natural openings on bed 
rock, growing on very thin hard humus. V. 3. 

11. Oyrophora hyperborca Ach. On almost bare rock in the foliose 
lichen zone (No. 2). Scarce. V, 2. 

12. Qyrophora (Vmbilicaria) vellea (C) Ach. Occurs in patches 
on vertical rock faces. I, 1. Fairly common. 

13. Bticta pulmonaria (C) Schaer. Cmnmon along Siskowit cabin 
trail, on trunks of fallen trees. Fairly common in similar places else- 
where in forest. V, 4 ; II, 

14. Peltidea (Peltigera) apkthosa (L) Ach. In forest along cabin 
trail to bog at Siskowit Bay. Fairly abundant (V, 4). 

15. Peltigera canina (C) Hoffm. On moist moss patches in woods, 
and in shady places. Common. V, 4. 

16. Placodittm elegans (Link) DC. On exposed rock surfaces along 



228 MICHIGAN SURVEY, 1908. 

shore. Very abundant. Also on conglomerate. Gives a striking darfc 
orange color to the cliffs along the main shore and on surfaces of small 
rock leefs and islands. I, 1; V, 2. The most striking ticben of the 
crustaceons lichen zone. 

17. Lecanora rttbina (Vill.) Ach. In foliose lichen zone. Not com- 
mon. V, 2. 

18. Lecanora mwalis Schrieb. In low rocky reefs Scarcely above- 
tlif action of nunimer waves. I, 1. Sot abundant. 

19. Lecanora fmstulosa (Diclis.) Ach. Rocky shores and cliff facea^ 
also in patches among parmelias. I, 1 ; V, 2. 

20. Lecanora atihfusoa allophana Ach. Rocky shores and exposed 
rocks. Fairly common, I, 1 ; V, 2. 

21. Lecanora cinerea gibhosa (Ach.) iJfyl. Back 20-25 feet fronn 
water's edge in second or foliose lichen zone, in lower edge of sanie. 
V, 2. 

22. Stereocaulon corallotdes Fr. In foliose lichen zone. V, 2. 

23. Stereocaulon paachale (C) Ach. Rocky openings near Sidcowit 
cabin. V, 3. Not abundant. 

24. Cladonia rangiferina (C) Web. Very common in cladonia zone- 
on all parts of shore where ciadonias occur. Probably most abundant 
of all ciadonias noted. 

25. Cladonia ejflvatica (C) Hoffm. Very common in cladonia zones- 
Lighter and smaller than preceding. Common in V, 2; I, 1. 

26. Cladonia alpestris (L). One of the principal form» in cladonia 
zone in the shore-heath swiee. Common. I, 1; V, 2. 

27. Cladonia cocoifera (C) Willd. Thin earth on exposed rocks. 
Fairly common. II, 3; V, 7 (in burned area). 

28. Cladonia dcfornus (C) Hoffm. On partly decayed bark' and wood 
of fallen trees. Not common. V, 4. 

29. Cladonia cristatella Tuck. On old wood. V, 4. 

30. Cladonia crispata (Ach.) Fib. Forest trail Siskowit. Also found 
growing into moss patches in woods. V, 4. 

31. Cladonia amoatirocraea (Flk.) Schaer. A cladonia in heath- 
cladonia zone at Rock Harbor, I, 1. Patches of this are being invaded 
by moss. , 

32. Cladonia furcata pi nnata Flk. Growing along cabin trail oa 
debris, and on fallen conifers. V, 4, Not abundant. 

33. Cladonia turgida (Ehrh.) Hoffm. Growing on thin humus ac- 
cumulation on open rocky places, V, 4. Also along portage to Siskowit 
Lake (V, 9). 

34. Cladonia gracilis dilatata (Hoffm.) Wain. On thin humus cover- 
ing of bed rock. Natural forest "openings." V, 4, 3. 

:!5. Cladonia rerticiHata Hoffm. Woods and rock clearing at Sis- 
kowit Bay station, V, 4, 5. 

36. Cladonia pyxidata (C) Hoffm. Rock shore in foliose and fmiti- 
cose lichen zones. V, 2. Not abundant. 

37. Cladonia finibriata simplex (Weis) Wainio. On decaying bark: 
of fallen trees in forest. Forest trail from Washington Harbor to Lake 
Desor. 

38. Cladonia fimhriata coniocraca (Flk.) Wainio. Bark of falleik 
trees in forest along Siskowit cabin trail. V, 4. Not abundant. 



ECOLOGY OF ISLS ROTALB. 229 

39. Leoidea lactea (Flk.). In rather amall patches an exposed slop- 
lug aod vertical rocky cliffs. GommoD on the water Bide of crustaceous 
or liiwer Hchen zone. A white lichen with black dots. I, 1; V, 2. 

40. Endocarrpon miniatum (0) Ach. Bock sorfaces with little soil, 
Along Siakowit cabin trail, V, 4. Also in lower crustaceous lichen 
zone. V, 2; I, 1. Not abundant. 

41. Endocarpon miniatum aaauaticum. In lower crnstaceous lichen 
zone within reach of winter wafes. Not abundant. I. 1; V, 2. 

42. lohmodophila aeruginosa. On decaying bark of fallen trees along 
fiiekowit cabin trail. Scarce. V, 4. 

43. Rhieocarpon (BuelHa) geographioum <C) DC. A small green 
lichen occnrring in small patches on rocky shores in crustaceous lichen 
zone, often near tbe water. Can only be removed by chipping away 
the i-ock on which it grows. Fairly common. I, 2; V, 1. 

if 08868. 

44. Sphagrwm teres Aug., var. tenelhtm Bl., bicolor. 

45. Sphagnum robustum Bl., var. gracile Rt., palleus. 

46. Sphagnum girgensohnii Ruas., var. molle Crev., palleua. 

47. Sphagnum platyphyllum Bull., var. subsimphx Cdbg., glaucum. 
The above sphagna were abundant in all the bog areas of the island, 

and were of occasional occurrence on the low ground along creeks and 
elsewhere on low wet ground. I, 4; I. 6; II, 2; II, 5; III, 5; IV", 10; 
V, 5; V, 8; V, 11. 

48. (Seorgia peXludda {Tetraphis pelliicida). Woods along Biskowit 
«abia trail. V, 4. 

49. Polytrichum commune C. A moat vigorous moss growing in 
-dense colonies; in places along the forest-bog tension zone successfully 
Invading the sphagnum masses. The only plant of the woods that 
could hold its own against the invasion of the Sphagnum into the forest. 
Confined to moist or wet places. V, 4, 

aO. Polytrichum atrictum Banks. Rather bare exposed places along 
<Jreenstone Ridge, also "natural openings" along the Biskowit cabin 
trail. V, 2; y, 4. 

51. Dicranum gchreberi. Near sphagnum bog at end of cabin trail. 
V, 5. 

n2. Dicranum fuaceacvns Turn. Occurs in small beads or clumps on 
dead wood. In woods. V, i. 

53. Dicranum longifoUum Hedw. Woods along Siskowit cabin trail. 
V, 4. 

54. Dicranum acoparium (C) Hedw. Woods along Siskowit cabin 
trail. V, 4. 

55. Dicranum unduiatum Volt. Woods along Siskowit cabin trail. 
V, 4. 

56. Qrimmia unicolor Hook. Confined entirely to small crevices and 
cavities in the bed rock of the gently sloping shore. It occurs nearer 
to the water's edge than any other form of vegetation observed on the 
island. No other mosses approach it in nearness to the water's edge. 
and it surpasses even the hardiest lichens of the crustaceous zone in 
this respect. Very hardy, and at times highly xerophytic. I, 1; V, 2. 
It is of a very dark greenish brown color. OoC^jlc 



230 MICHIGAN SURVET. l»Og. 

57. Lmeohryum glaucum (L) Schimp. Grows in beade of varying^ 
size, principally in woods. I, 3 ; V, 4. 

58. Tortella tortuosa (h) Limpr. Bock ridges, and other rocky 
plHces. firo^fi in dense rounded fufts. II, 3. 

59. Vlota americana (Beauv.) Liitdb. Growing on gently 8lopiD|r 
rock shore, sometimeB covering crustaceous and foliose lichen patcbes. 
I, 1 ; V. 2. 

60. Bartramia pomiformis (L) Hedw. Shady, moist niches and 
crevicefl in rock cliffs, A beautiful moss having the appearance of green 
wool. I. 1; V, 4. 

61. BryuM palleus Swartz. Gn dead wood, and on thinly covered 
rock surfaces In woods, V, 4, 

62. Aulocomium palustre (L) Schwaegr. Near b<^ at end of cabin 
trail ei8k<>wit Bay. V, 5, 

63. Mnium punctatum Hedw. Moist woods along Benson Brook; 
also in moist places along giskowit cabin trail through woods. II, 1; 
V, 4. 

64. Leakea nermaa (Schwaeg.). Myr. Closely associated with Vlota 
americana on the sloping rock shore where it sometimes coverB patcbes 
of cpustaceous and foliose lichens. I, 1 ; Y, 2. 

65. Thvidium abietinum (L) B. & 8. Growing on fine material that 
has accumulated among the close branches of the low Procumbeat 
Juniper, It was noted on one of the small rock islands ia Bock Harbor 
which had little if any vegetation besides the crevices plants. It here 
plays an important pert in the early formation of a hamus soil by 
solidly filling in the spacer between the Jnnii^er branches. I, 1. 

66. Hypmim crUta-castrensia L. On decayed wood in coo! moist 
woods near peat bog, V, 4, 5. 

67. Eppntim ackreberi Willd. Rich, moist woods along forest road 
Washington Harbor; also noted growing in Cladonia patches in woods 
along Siskowit cabin trail. It seems to be replacing and Bucoeeding the 
Cladonias in places. V, 4; I, 3. 

68. Bypmun scorpoides L. B(^ margin of Forbes Lake. II, 5. 

69. Hiypnum inemicoswm Lindb. Bog beyond Malone's fishing camp. 
Back from V, 2. 

70. Hypnum polarc Lindb. Protected rock crevices, Bock Harbor. 
I, 1. 

71. Bypnum, ftuitans L. Bock pools Scovill Point. IV, 1. 

7-. Bypnum «tramincum Dicks. Bog beyond Malone's fishing camp. 
Back from V, 2, 

73. Hypnum aduncam Scb. Bog at end of Siakowit Bay cabin trail. 
A", 5. 

74. Hi/ptuiiii, (uUiiicum inttrmcdiitm Seh. Growing in water in mar- 
gin of a brook emptying into Forbes Lake, II, 5. 

75. BypHviii uncinatum Hedw. formaplumoaa Sch. Moist woods 
along trail to Monument Bock. iV, 4. 

76. Byloconiium triquetiim (L) B. & S. Woods along Siskowit cabin 
trail. V, 4. Fairly common. 

77. Bkylocomiiim aplendcns. Woods along forest road from Wdbhing- 
toii Harbor to Lake Desor, III, '04. 

78. Distichiiim capillaceinii. From a partly protected vertical rock 
crevice 6-8 feet above water. Rock Harbor. I, 1. i, CoOqIc 



■ ECOLOGY OF ISLB ROYALB. 231 

79. Neckera oligocarpa B. &■ S. Forest road, Washington Harbw: 
woods. 

80. Dickelyma uncinatum Mitl. (?) Growing in a pool on small 
island at upper end of Rock Harbor. IjDaBnally lai^. Ill, 1. 

81. Palndella squarrosa (L) Brid. Bog margin of Forbea Lake. II, 
6. 

Pteridophytes. 

Ophioffloasaceae — Adder's Tongue Family. 

82. Botrychium lunaria (L) Sw. Moonwort. Bare. Partially shaded 
rocky ground near Rock Harbor Ugbt-honse. I, 1. 

83. Botrychium virginicum (L) Sw. Yii^nia Grape Fern. Spar- 
ingly distributed in rich woods. I, 3; III, 4; V, 4. 

Oamundoceae. 

84. Osmunda regalia L. Royal Fern. One locality. Rich low ground 
near small creek emptying into Forbes Lake, II, 5. 

85. Ogmunda cinnamomea L. Cinnamon Fern. Moist thickets and 
low ground. II, 6. 

86. 0$munda claytoniana L. Interrupted Fern. Mesophytic woods. 
!Not common. 

Polypodiaceae — Fern Family. 

87. Onoclea teneibilis L. Bensitive Fern. Fairly abundant. 

88. Onoclea strutkiopteris (L) Hoifm. Ostrich fern. Few localities : 
not abundant. 

89. Woodaia ilvenaia (L) R. Br. Rusty Woodaia. Several small, 
dense patches on rock surfaces, and along rock crevices. Island upper 
end of Rock Harbor and I, 1 ; V, 2. 

90. Cystopteris hulbifera (L) Bernh. Bulblet Cystopteris. Thinly 
scattered on moist, shaded cliff faces. Cliff near Bock Harbor ligbt- 
bonae. 

91. Cystopterig fragilis (C) Bernh. Brittle Fern. Shaded, moist 
places. Not abundant. 

92. Dryopteria the^teria (L) A, Gray, iMarsh Fern. Wet mar- 
gins of .bogs, and other low, wet places. Common in such places. II, 
5; III, 5. 

93. Dryopteria fragrana (L) Schott. Fragrant Shield Fern. Com- 
mon in patches on cliffs and rocks along shore, I, 1. 

94. Dryopteria filiwmas (L) Schott. Male Fern. Fairly abundant 
in rich, moist woods. Especially abundant near Benson Brook. II, 1. 

95. Dryopteria apinuloaa (Retz.) Kuntze. Spinnloee Fern. Rich, 
moist woods. Fairly common. Unusnally large and vigorous on Malone'a 
Island in Siskowit Bay. Ill, 4; V, 4. 

96. Phegopteris phegopteria (L) Underw. Long Beech Fern. Moist 
woods (IV, 4). Less common than P. dryopteria. 

97. PJiegopteria dryopteria (L) Fee, Oak Fern. Bich, moist woods. 
Fairly common. IV, 4; III, 4. 

98. Aaplenium trtchomatu^a L. Maiden-hair Spleenwort. On thinly 
soil-covered rocks. Rare. Rock cliflf along Siskowit cabin trail. V, 4. 



232 UICHIOAN SURVBT, 1908. 

&9. Adiantum ped^tum L. Maidenhair Fern. Sparingly distri 
bnted in the mesophytic forest. Ill, 4, and at Washington Club (forest). 

100. Pteria aquilina h. Brake. Abundant in open, drier places, 
especially in bnmed areaH. 

101. Cryptogramma acrostiekoidca B. Br. American Rock Brake. 
In denfie jKitches on exposed bed-rock where thinly soil-covered. Upper 
end of Rock Harbor and I, 1. 

102. Pokypodium milgare L. Common Polypody. Tops and exposed 
edges of cliffs. Common. I, 1. 

Eguisetoceae — Horsetail Family. 

103.. Eqtii»etum arccnse L. Between forest and bog margin Forbes 
Lake, II, 5. 

104. Equieetum sylvaticum L. Wood Horse-tail. Moist woods. II. 
1. Noted in one locality only. 

105. Eguiaetum palustre L. Marsh Horse-tail. Wet, back-margin of 
bog. 

106. Equisetum fluviatUe L. Swamp Horse-tail. In water apper 
end of Rock Harbor. Ill, 3. 

107. Equisetum ncirpoides !tlichx. Depression in Arbor-vitae swamp 
along Siskowit I^ake portage, (V, 9). 

Lycopodiaceae — Club-Moss Family. 

108. Lycopodium selago L. Fir Club Moss. Rare. . Exposed rocks 
at Scovill Point, IV, 1. 

109. Lycopodium lucidulum Michx. Shining Clnb Moss. Edge of 
rock pools Scovill Point, IV. 1, and in moist woods. III, 4. 

110. Lycopodium invndatum L. Bog Clnb Moss. Wet bog margin, 
Sumner Lake. Ill, 5. 

111. Lycopodium obgcurnm L. Ground Pine, Sparingly distributed 
in moist woods. I, 3. 

112. Lycopodium clavntum L. Running Pine. Common in dry to 
moist woods. Ill, 4; V, 4; I, 3. 

113. Lycopodium complanalum L, Fairly common in woods and 
shady places. V. 4; III, 4. 

114. Lycopodium annotinum h. Stiff Club Moss. Cool, dry woods. 
V, 4. 

Selaginellaceae — Selaginella Family. 

115. Selaginella rupestria (L) Spring. On thinly soil-covered rocks 
along Siskowit Lake portage (V, 9). Sparingly distribnted. 

Isoetttceae — Quillwort Family. 

116. lioetcB sp? In shallow water at upper end of Bock Harbor, 
III, 3. 

Spermatopkytcs. (Seed Plants). 

Xaiad4iceae — Pondweed Family. 

117. Potamogelon natans L. Fairlv common on mai^n of Sumner 

'-'«■ ™'^- ' ,„ Google 



ECOLOGY OF ISLE ROYALB. 233 

118. Potamogetott perfoliatus L. Claeping leaved Pond-weed. Mar- 
^n of Snmner Lake. 

119. Potamogeton heterophyllaa. fichreb. Washington Creek. 

120. potamogeton hillii {?) Hill's Potamogeton. Uargia of Butnoer 
■Lake. 

121. Potamogeton. pectinatus L. Margin of Sumner Lake. 

122. Tfaiag ^ctili* WHld. Slender Naias. Shallow water at head of 
Bock Harbor. 

Scheuchzeriaceae — Arrow-Grass Family. 

12.t. Triglochin maritima L. B(^ margin of Sumner Lake. Not 
Abundant. Ill, 5. 

Vallianeriaceae — Tape-Orass Family. 

124. Valliineria spiralis L. Tape-GrsBs, Eel-Orass. Shallow water 
-at bead of Bock Haii>or. 

Graminae — Grass Family. 

125. Panicum ranthophysum A. Gray. Dry rocky ridges, and rocks 
"with little soil. 

126. Agroatia hyemaUe (Walt.) B. S. P. Rather dry ground. 

127. Calamagrostia canadensis (Micbz.) Beauv. Bock pool margins. 
■Scovill Point, IV, 1. Creek margin upper end of Bock Harbor, III, 3. 
Wet places generally. 

128. Trisetum subsptcatum (L) Beauv. Common in rock crevicee 
and dry places. Bock shore where little soil is present, where it occurs 
■as the picmeer grass. I, 1; V, 2. 

129. Pkragmites phragmHes (L) Karst. Washington Creek. 

130. Poa pratenaia L. A dry ground form. Island in Bock Harbor, 
III, 1. 

131. Panicularia canadensis (Michx.) Kuntze. Upper end of hog at 
«Dd of Siskowit cabin trail. V, Q. 

132. Panicularia elongata (Torr.) Kuutze. Mar^n of Siskowit cabin 
trail bog. V, 5. 

133. Feaeuta ovina L. Bock crevices and on thinly soil-corered rocks. 
I, 2; V, 2. 

Cyperaceae — Sedge Family. 

134. Eleocharig paUistria (L) B. & S. In shallow water at upper 
«nd of Siskowit cabin trail. Y, 5, 

135. Eleocharia patuatris glauceacens (Willd.) A. Gray. (?) Wet 
part of island in Tobin Harbor. 

136. Scirpus caespitosug L. Margin of Forbes I^ake, II, 5. Bock 
poors, Scovill Point, IV, 1. 

137. Scirpua cjfperinua (L) Knnth. Tx>w ground along "Island^ 
mine" road, head of Siskowit Bay. 

138 Eripphorum alpinum L. Alpine Cotton-Grass. Slost common 
'"cotton-grass" on the island. Common in all the bogs. II, 2; III. 5. 

139. Eriophorum vaqinatum L. Sheathed Cotton-Grase. Spht^nnm 
tKigs. V, 11; III, 4; V, 5. 



234 MICHIGAN SURVEY. 1808. 

140. Eriophorum gracile L. Bog margin of Samner Lake, II, 5. 

141. Rynchospora alba (L) Yahl. White Beaked Bu^. Common in 
vet bog margiuB. V, 11; II, 5. 

142. Carex paucifiora Lightf. Few-flon*ered Bedge. Margin of 
Siskowit cabin trail bog, V, 5. 

143. Carex folliculata L. Long Sedge. Associated with preceding- 
species. 

144. Carex monile Tnckerm. Necklace Bedge. Wet creek margin 
of bog near Malone's fishing camp, (V, 11). 

145. Carex tuchermani Bewev. Along Washington Creek. 

146. Carex retrorsa Schwein. Betrorse Sedge. Along WashingtoD 
Creek. 

147. Carex riparia Curtis. Biver-bank Sedge. Creek margin heaif 
of Bock Harbor, III, 3. 

148. Carex fiUformis L. Blender Sedge. Common in 1k^ mat;gin»> 
IL 5; IIP, 5. 

149. Carex atricta Lam. Tussock Bedge. Bog margins. Y. 11; 

II, 5. 

150. Carex a^uatilis Wahl. Water Sedge. Bog margins. V, 11 r 

III, 5. 

161. Carex Umoaa L. Mud Sedge. Bock pools, Scovill Point, IV^ 
I. Siskowit cabin trail bog. V, 6. 

152. Carex crinita Lam. Along road to "Island Mine" head of Sis- 
kowit Bay. 

153. Carex arctata Boott. Drooping Wood Sedge. Dry woods,^ 
Washington Harbor. 

154. Carex viridula Michx. Edge of rock pools, and on moister 
parts of rock beach. I, 1; V, 1, 2. 

156. Carex chordorhiza L. Creeping Sedge. Bog margins. V, lip 

III, 5; II. 5. 

156. Carex teneUa Schk. Soft-leaved Sedge. Arbor-ritae depree- 
sioD, Siskowit Lake portage, (V, 9). 

157. Carex aterilis Willd. Bock pools, Scovill Point, TV, 1, and 
Siskowit cabin trail bog. Y, 4. 

158. Carex hruanesoena (Pers.) Poir. Bock pools, Scovill Point. 

IV, 1. 

169. Carex triaperma Dewey. Three-fmited Sedge. B(^ margins, 
n, 5; III, 5. 

160. Carex acoparia Schk. Pointed Broom Bedge. Washington 
Club QroiiDds. I, '04. 

161. Carex fe8tucacea> Willd. Fescue Sedge. Dry rocky places;, 
rock ridges. II, 3; V, 3. 

NOTE. — For more convenient reference the principal trees of the 
island will be grouped together instead of being placed under their- 
respective orders and genera. 

Pinaceae — Pine Family. 

162. Ptnus strobM L. White Pine. I<ai^, isolated indiyidual» 
occur along the Greenstone Bidge, and on other ridges; bat is no- 
where abundant. It is confined almost entirely to higher ground, and 



ECOLOGY OF ISLE HOYALE. 236 

to Open, sunny places. Large, cbarred trunks 3-4 feet in diameter are 
still fairly abundant along the Greenstone and other ridges. Very 
few young trees of this species were noted; and there are do indica- 
tions, at present, to suggest that it will again become abnndant on tlie 
island. I, 3; II, 3; III. '04; VII, '04. 

163. Pinus resinoaa Ait. Red, or Norway Pine. Not abundant; 
noted in two localities only. Occurs on high, exposed ground. Ill, 4, 

164. Pinus divaricata (Ait.) Budw. Labrador, or Gray Pine. Fairly 
common on exposed, dry rock ridges, and on a few of the rock islands. 
Several in heath-forest tension zone near Bock Harbor light-honse 
Able to withstand highly zerophytic conditions. 

165. Larix Iwicina (DuBoi) Koch. Tamarack, or American Liirch. 
Principally in recently filled bogs or working in along mai^ns of 
partly filled ones. The tamaracks and Black Sprnces are the pioneer 
trees of the bogs. In the older bogs the few large tamaracks present 
are relicts, and few young ones appear to be coming on. Scattered in- 
diyidnals occur throughout the upland forests but are nowhere abund- 
ant outside the bogs. Largest individuals noted (V, 8) were over 3 
feet in diameter, I. 4; T, 6; II, 2; II, 4; II, 5; III, 5; IV, 4; IV, 8; 
V, 5; V, 7; V, 8; V, U; V, '04. 

166. Abies balaamca (L) Mill. Balsam Fir. The most character- 
istic and abundant e\'ergrcen of the upland forest. Abundant on all 
parts of the island except the Greenstone Ridge, and in the more re- 
cently filled b(^. Along the forest road from Washiagton Club to 
Lake Desor the absence of the larger firs was probably due to the 
shallowness of soil, exposure to the powerful winter gales (as soon 
as they overtop the maples and other trees among which they start to 
grow), and the reduced root system in proportion to the size of the 
tree. It reproduces i-eadily in dense shade as well as more open 
places, and is not only succeeding itself but other forest trees, as the 
White Spruce. It will certainly occupy a large and important place 
in the climatic forest. Up to 3 feet in diameter. I, 3; I, 4; III, 4; IV. 
4; IV, 8; IV, 9: V, 4; V. 7; ITT, '04. - 

167. Picea oanadensia (Mill.) B. 8. P. White Spruce. Older trees 
are fairly common where they have come in with Abies as pioneers. 
It does not appear to be succeeding itself except along the edge of 
clearings and in more open parts of the forest. Since the Fir seedlings 
ere common under the older trees instead of those from the present 
spruces it appears that the White Spruce will be replaced by the Fir 
in the climatic forest, the Fir seedlings being able to endure much 
deeper shade. I, 2; I. 3; III, 1. 

168. Piwa mariana (Mill.) B. 8. P. (Possibly Pioea brevifoHa 
Peck). Black Spnice. Confined principally to sphagnum bogs where 
it cou'es in with the tnniaratk as a pi»neer. The largest trees noted 
were '2'/j feet in dijimeter. AIho H)mringly disirihnted outside of bogs. 
In a few instances it Wiis found growing on the exp<!sed tops of cliffs 
(as at Roek Harbor) where there is only a thin covering of soil. I, R. 

16!). Thuya oifidrntalis L White ("cdar or Arlwrvitae. Occurs iit 
all bog ureax except those mont recently cnriieted ovr. It does not 
appear to ronie in as a pionM-r but follows closely Pirra iitariatm and 
Larij- lanritia. Ijargpsl n|>ecinien8 in old bog areas, V, 8, were 40 inctieR 



236 MICHIGAN SURVEY. 1908. 

■D diameter. Occasional in upland forest, in fact, fairly abnodant in 
placee; also one of tbe trees to occnr In crevices on the small rock 
islands and along the rock shore, in which cases they have a decidedly 
Rtanted appearance, and are often broader than high. I, 4; I, 6; II, 2; 

IV, 4; IV, 8; V, 6; Y, 7; V, 8; Y, 11; II, '04; Y, '04. 

170. Jumperus nana Willd. Low evergreen shrub common on the 
back heath zone and along the rock shore. It appears to follow 
rather than to precede Juniperus {procwnbens) sahitui with which it is 
80 commonly associated. A. common form on the rock islands and in 
the rock shore crevices. Also in the natural rock openings back from 
the shore. I, 1; I, 2; I, 5; III, 1. 

171. Jwniperus {procumhcns) sabiva L. Procumbent Juniper, A 
very important pioneer on the rock Islands and on the sloping rock 
shore, starting as a crevice plant and sending out its dense prostrate 
branches 6-10 feet- It offers a favornble place for wind blown material 
which there accumulates, and this is of great importance for the 
pioneer mosses which contribute so largely to the first humos soil. 
Rome most interesting examples of these pioneer stages were noted on 
one of the low, nearly bare rock islands near the Rock Harbor light- 
house, I, 1, and at V, 2, it was very abundant. 

Taxaceae — Yew Family. 

172. Taxtu canadonsia Marsh. Ground Hemlock, American Yew. 
Everywhere abundant in the upland forests of the island. On account 
of its low, spreading growth it forms one of tbe greatest impedimenta 
in penetrating the island forests. The rankest growth was noted in the 
lower forest region around Washington Harbor, where it attains a 
height of four to five feet. I, 6; lY, 4; lY, 8; IV, 9; V, 4; V, 5; V, 7; 

V, '04. 

iSflltcaccflc— Willow Family. 

173. Popuhia grandiientata Michz. Large-toothed Aspen. Princi- 
pally along tiie Qreenston« Bidge; not at all common as compared with 
P. tremuMdea. 

174. Popuhut tremuloides Michx. American Aspen. Very cwnmon 
on almost all parts of the islands where burnings and clearings have 
occorred. This and the Betula papyrifera are the pioneer decidaous 
trees is bnmed and cleared areas, where the two seem about equally 
abundant, colonies of both being intimately associated. Younger and 
older staods of this as noted along the Greenstone Ridge near Rock 
Harbor, suggest the vounger and older burnings bv the copper profi- 
pectoFB. I, 1; I, 2; I, 3; I, H; I, B; I, 7; II. 1; 11, 3: III, 4; IV, 5; IV, 
9; V, 8; V, 9; V, 3; Y, t; V, 5; V, 7; I, '04; III, '04. 

76. Populus balaamifera L. Balsam Poplar. One locality only ; 
head of Siskowit Bay. 

Betulaceae — Birch Family. 

176. Corylua roatrata Ait. Beaked Hazel. Rocky slopes and sum- 
mits of ridges. In thickets along the Qreenstone- 

177. Betula papyrifera Marsh. Paper, or Canoe Birch. Common 



KCOLOGY .OF ISLE ROYALE. 237 

everywhere In forested portions as well as burnings and clearings. 
This and the Balsam Fir seem to he the climax trees of the upland 
forest. I, 2; I, 3; I, 7; 11, 1; III, 4; IV, 8; IV, 9; V, 3; V, 4; V, 7; 
I, '04; III, '04. 

178. Betula lutca Michx. F. Yellow Birch. Noted only along the 
forest road from Washington Harbor to Lake Desor, where it was very 
common along the Oreeui^tone Uidge. (Specimens SU inches in diameter 
were noted. Ill, '04. 

179. Betula lenta L. Black or Cherry Birch. Associated with D. 
lutca as mentioned above. Also attaining great size. Ill, '04. 

180. Alnv8 alnobctuUi (Ehrh.) Koch. Green, or Mountain Alder. 
Fairly common in npland forest at Rock Harbor. Common shrub along 
with birches and nspens. 

181. Alnus incajia (L) Willd. Speckled Alder. Low ground, Irorders 
of streams and margins of lakes. Along water's edge at Bock Harbor, 
and sparingly associated with Alnu^ alnobctula in the forest back from 
■water. 

Note. — Thru an ovw-sight the 3 species of Salix observed wera 
omitted in preparing this list for the press. 

Fagaceac — Beech Family. i 

183. Quereiis ruhra L. Red Oak. .\ single specimen was noted along 
the forest road between Washington Club and Ijike Desor, (III, '04). 
The onl}/ oalc noted on the island. 

Pomaceae — A]>ple Family. 

183. SorbuS americana Marsh. American Mountain Ash. Fairly 
common along the forested margins of the principal inlets, as Rock 
Harttor, and sparingly distributed through the inland forest. Always 
more or less isolated, never in colonies. 

184. Aronia nigra (Willd.) Britton. A single specimen noted on 
north side of Rock Harbor. 

186. Amelanchier ainifolia Nutt. Nlorthwestem Juno-lrerry, A 
shrub 6 feet or less in height. Rock openings also rock ridge near 
Conglomerate Bay (I, 5), 

186. Amelanchier oUgocarpa (Michx.) Roen. Oblong-fruited June- 
berry. A shrub about the size of preceding, but occurring on lower 
ground. 

Dnipaceae — Plum Family. 

187. Prumia penaisylvanioa L. Wild Red Cherry, Pennsylvania 
Cherry. Characteristic of xerophytic places as rock openings, talus, 
slopes, and bnmings; and able to thrive in exposed rocky positions- 
where subject to great temperature extremes, and where there is very 
little soil. I, 5. 

188. Prtmtis mrginuina h. Choke Cherry. Woods: not common. 

Aceraceae — Maple Family. 

189. Acer aaccharum Marsh. Sugar or Hard Maple. One part of 
the island, on the summit of the Greenstone Ridge along the forest 



238 MICHIGAN SURVEY, 1908. 

road from Wasbington Harbor t^ I^ake Desor, it is very abundant. 
Reported to occur eparingl; along other parts of the Oreenstone, bnt 
8eemB to be coufloed entirely to the higher parts of the aummit ridge. 
Along this "forest road" it forms almost pure stands, iu other places 
there is B. lenta and B. luted mixed with it. Some of the trees are 2-3 
feet iu diameter. {Ill, '04.) 

190. Acer apicatum Lam. Mountain Maple. Generally distributed 
in the forest, but nowhere very abundant. Largest trees over 30 feet 
high. One of the lower growth forms to invade the forest roads. Oftea 
in rocky places. V, 7; III, '04. 

191. Acer pennaylvanicum. Striped Maple, Moosewood. Rare on 
island. 

Oomaccae — Dogwood Family. 

192. Comn8 atolonifera Michx. Common in low ground and back 
margins of bogs. A prominent member of the shrub zone Hnrronoding 
amall lakes. 

193. C<rrnvs circinata L'Her. Round-leaved Cornel. Sparingly dis- 
tributed in rich woods. 

Araceae — Arum Family. 

194. Calla palustris L. Water Arum. Lake margins, especially 
abundant at Sumner Lake, III, 5. 

195. Spathyema foetida (L) Baf. Skunk Cabbage. Common in low 
grounds in woods, and near logs. Ill, 5; II, 1, 2; II, o; IV, 4. 

Juncaceae — Rush Family. 

196. Juncus effusus L. Along old road to "Island mine," and in 
shallow water at upper end of Rock Harbor. Ill, 3, 

Meianthaceae — Bunch-Flower Family. 

197. Tofieldia paluatria Huds. Asphodel, Rocks at Scovill Point, 
TV, 1. 

189. Uvularia perfoliata L. Perfoliate Bellwort. Rich, moisi woods. 
Scattered. 

lAlwceae — Lily Family, 

199. TJUum phiUidelphicum h. Bed, or Wood Lily, Common in drier 
pnrts of woods ; even occurs as a rock crevice plant on the small islands 
in Rock Harbor, 

ConraHorifloMe—Li!y-of -the- Valley Family. 

300. Clintonia iorealia (Ait.) Raf. Yellow Clintonia. Common 
everywhere in moist, rich woods; verv abundant in places. I, 3; TV, 
4; V, 4; V, 5. 

201. Vagnera trifoUa (L) Morong. Three-leaved Solomon's Seal. 
Frequent in forest margins of bogs, and in cool, moist woods. I, 4: II, 
2; V, 5, 



D,3-ZBcbyCOO<^IC 



ECOLOGY OP ISLE BDYALE. 239 

202. VnifoUum eanadetme (Deef.) Greene. False Lily-o{-YaUey, Two- 
leaved Solomon's Seal. Rather open patches in rich, tnoiat woods. 1, 
4; 11, 2. 

203. Streptopas amplexicaiUia (L) DC. Claaping-Ieaved Twisted- 
stalk. Woods along porta^ toSiskowit Lake (V. 9). 

204. Trillium grandiflorwn iiiicb:!.) Salisb. Showy, White Triliitim. 
Flood plain of Washington Creek. 

Iridaceae — Iris Family. 

305. Iris versicolor L. Larger Blue Flag. Common in low wet 
places, as lake and bc^ margins. Y, 5; III, 5; II, 5. 

Orehidaceae — Orchid Family. 

206. Cypripedium reginae Walt. Showy Ladies-Slipper. Wet places 
in woods : not abundant. II, 1 ; near II, 5. 

207. Gypripedium hirmtutn Mill. Larger Yellow Ladies-Slipper. In 
drier parts of woods than preceding. lY, 4. 

208. Orchis rotundifolia Pursh, Small Bonnd-leared Orchild. Bare: 
tamarack forest. 

209. Habenaria orbiculata (Pursh) Torr. Large Round-leaved 
Orchid. Rich woods near lY, 2; few localities. 

210. Habenaria oMusata (Pursh) Richards. Small Northern Bog 
Orchid. Fairly common in forested hog margins, and bog forests. I, 
4; II, 2; Y, 5. 

211. Eabenaria h}/p^borea (L) R. Br. Tall Leafy Oteen Orchid. 
B<^ and wet woods: margin of Sumner Lake, II, 5. 

212. Habenaria dilatata (Pursh) Hoc^. Tall White Bog Orchid. 
Trembling hog margins of Forbes and Snmner Lakes, very abundant 
in latter place. II, 5; III, 5. 

213. Habenaria pstfoodes (L) Gray. Smaller Pnrple-fringed Orchid. 
Associated with H. dilatata as given above, and also abundant. Ill, 5; 
II, 5. 

214. Pogonia ophiogloasoidea ( L) Eier. Rose Pogonia. Common 
along wet bog margins. Ill, 5; II, 5. 

215. Arethusa bviboia L. Aretbusa. Wet bog margins. Xot so 
common as preceding species. 

216. Qyrostachya romancoffiana (Cham.) MacM. Wet margins of 
Sumner J^ke and Forbes Lake. 

217. lAstera cordata (L) R. Br. Heart-leaved Twayblade. Moist 
woods and ravines. 

218. Peramium repens (L) Salisb. Lesser Rattle-snake Plantain. 
Cabin trail woods, Y, 4, Siskowit. 

219. Peramium pubesoens (Willd.) MacM. Downy Rattle-anake 
Plantain. Rather dry woods. Y, 4, III, 4. 

220. Peramium. menzieaii (Lindl.) Morong. Menzies' Battle-snake 
Plantain. Rich woods. Y, 4; III, 4. 

221. Acroanthcs monophylla (L) Greene. (?) Sumner Lake mar- 
gin. III. 5. 

222. Leptorehis UUifoIia (L> Kuotze. Large Twayblade. Uoist 
woods and along bog margins. Woods of I. / - ^ i 



240 MICHIGAN SURVEY, 1908. 

223. Leptorchit lofselii (L) UacM. Loesel'B Twayblade. Wet thick- 
ets and flpring banks, 

224. Calypso iulbosa (L) Cakes. Calypso. Wet, cool woods and 
ravines. 

225. Corallorhiza corallorhiza (L) Karst. Early Coral-root. Rich, 
moist woods. Woods at upper end of Bock Harbor. 

226. Corallorhiza multi flora Nutt. Large Coral-root. Fairly com- 
mon in rich woods, I, 4; III, 4. 

Santaiaceac — Sandalwood Family. 

227. Comandi'a livida Richards. Northern Comandra. Tfaiu soil on 
rocks, and in open, xerophytic plac^. Pine ridge near Sumner Lake. 
Ill, i. Fairly common. 

Ariatolochiaceae — Birthwort Family. 

228. Aaarum canadense L. Wild Ginger. Flood plain, Washington 



il///»-icac(;ae— Bayberry Family. 

229. Myrica gale L, Sweet Qale. Margin of bayou off Tobin's Har- 
bor; wet rocks at Sco\iIl Point. Also V. 6. 

CaryophyllaGeae — Pink Family. 

230. Sagina saginoides (L) Britton. Arctic Pearl-wort. A hardy, 
low, rock crevice plant. I, 1. 

231. AUine longifoUa (Muhl.) Britton. Long-leaved Stitch-wort. 
Scattered ruderal. II, 1. 

232. Silene antirrhina L. Sleepy Catch-Fly. Side of Greenstone 
Itidge, and exposed xerophytic places. Not abundant. II, 3. 

Nymphaeaceae — Water-Llly Family. 

233. Brasenia purpurea (Michx) Casp. Water Shield. Open water 
in a few bogs; not abundant. Bog near Malone's fishing camp, V, II. 

234. Nymphaea advena Soland. Large Yellow Pond-Lily. Margins 
and shallower water in a few bogs. Ill, 5. 

235. Castalia odorata (Dryand.) W. & W. Sweet-scented White 
Pond-Lily. Abundant in Sumner Lake, III, 6, where it seems to grow 
in part on tbe uplifted "false bottom." 

Ranunculaceae — Crowfoot Family. 

236. Caltha palustria L. Marsh Marigold, Cowslip, Wet places in 
woods. Low woods at head of Bock Harbor, II, 1, 2; III, 5. 

237. Coptia trifolia (L) Salisb. Gold-thread. Hummocks in wet 
woods and filled bogs, and in wet bog margins. Common. 

288. Actaea rubra (Ait.) Willd. Bed Baneberry. Sparingly dis- 
tributed in woods. V, 4, 9; IV, 4. 

239. Aquilegia canadensis L. Wild Bed Columbine. Rocks near 
lieht-boQse at Bock Harbor. Not abundant. r^ ' i 

D= z.<:byL.OOg[e 



ECOLOGY OF ISLE ROTALB. 241 

240. Anemone multifida Poir. Red Wind-Flower. Bare. Bock shore 
or one island in Rock Hai-bor. 

2*1, Eepatica hepatica (L) Karet. Bound-lobed Hepatica. Woods: 
not abundant. 

242. Sanunculus abortivua L, Kidney-leaved Crowfoot. Scattered as 
a rnderal. II, 1, and on Waahington Club grounds. 

243. Ranunculus omlia Eaf. Thin soil on rock islands. Upper end 
of Rock Harbor, III, 1; also near Biskowit cabin (V, 1). 

244. Ranunculus maeounii Britton. Macoun's Buttercup. Bare, one 
locality, near Biskowit cabin (V, 1). 

245. Thalictrum purpuraacens L. Tall Purple Rue. Moist, rich 
woods near Benson Brook (II, 1), and along Washington Creek. 

Papaveraceae — Poppy Family. 

246. Capnoides sempervirena (L) Borck. Pink Corydalis. Rocky, ex- 
posed places along the Greenstone. II, 3. 

CruCT/erac— Mustard Family. 

347. Thlaspi arvense L. Field Penny Crew. Washington Club 
groonds. Only locality. Bnderal. 

248. Siaymbrium. altissium L. Tall SiBymbrinm. Washington Clulx 
grounds: waste places. I, '04. Rnderal, 

249. Arabis bracht/carpa (T. &G.) Britton. Pnrple Rock Cress. Ex- 
posed rocks. Greenstone Ridge along the McCargo Cove trail. II, 3. 

SarracfmMceae — Pitcher Plant Family. 

250. Sarracenia purpurea L. Pitcher Plant. Common in bog areas 
everywhere on island. I, 6; II, 2; II, 5; V, 6; V, 11. 

Droaeraceae — Bundew Family. 

261. Droaera rotundifolia L. Round-leaved Drosera. Common along 
wet bog margins, especially III, 5. 

252. Droaera intermedia Hayne. Bpatulate-Ieaved Bundew. Bc^ mar- 
gins, bat generally in wetter parts than the preceding; often elevated 
on a short stem extension. Ill, 5. 

253. Droaera liru:aria Goldia Slender-leaved Drosera. Bog margins; 
fairly abundant. Ill, 5. 

Saxifragaceae — Saxifrage Family. 

254. SaiBifraga tricuspidata Eetz. Three-toothed Saxifrage. Fairly 
common as a crevice plant along the low rock shore. L, 1. 

265. Saxifruga aizcKyn Jacq. Livelong Saxifrage. A rock shore crev- 
ice plant. Bare, V, 2. 

266. Sawifraga nivalis L. Clustered Alpine Saxifrage. Exposed rook 
shores, growing on scanty soil. I, 1 ; Y. 2. 

257. MitelUt nuda L. Naked, or Low Mitrewort Very common in 
moist woods. I, 3. Woods at end of Rock Harbor and on forested 
ialands. ^~. ■ 

32 lyCoogle 



242 MICHIGAN SURVEY. 1908. 

258. Panmssia paJu^itrig L. Northern Omss of FamaesaB. "Bog mar- 
gin of Sumner Lake. Sparitigl; distributed. 

OroxmtJariaceac — Gooseberry Family. 

259. Ribea setoaum Liadl. BriBtly GooBeberry. Shore of Siskovit 
Lake. Bare. 

260. Ribes proatratum L'Her. Fetid Currant. Rich, moist woo&; 
also one of rock islands at Hock Harbor. Fairly abandant. 

261. Ribea rulrum L. Bed. Currant. Growing wild in abundance in 
vicinity of Biskowit Lake portage. V, 9. 

■ Hoaaceae — Rose Family. 

262. Opulaster opuUfoUus (L) Kuntz. Kinebark. Occurs principally 
along the shores, often aa a crevice plnnt on the root islands, as well 
as on rocky shore of main land. I, 1; III, 1; V. 6, 

263. Rvbua parviflorus >'iitt. White-flowering Raspberry. A very com- 
mon and characteristic plant of clearings and burnings; also occurs in 
thickets and open parts of woods. VIII, '04. 

264. Ruhut arcticus L. Arctic Raspberry, or Bramble. Sparingly 
distributed in moist woods and filled bog areas. 

265. Rubua atrigoans Michx. Wild Red Raspberry. Found most 
abundant in the burned areas at head of Siskowit Bay. ' 

266. Riibua americanvs (Pers.) Britton Dwarf Raspberry. Occa- 
sional in woods; rather common in bog forests. 

267. Fragaria veaca L, Sparingly distributed. I, 1. 

268. Poteatilla arguta Pursh. Tall "WTiite Oinquefoil. Common 
around light-house clearing at Rock Harbor as a ruderal. I. 

269. Potentilla monspelienais L. Rough Oinquefoil. Exposed rocks 
having scanty soil. 

270. Potentilla tittoratia Rydberg. Coast Cinqnefoil, Fairly common 
as a rock crevice plant along main shore, and on small rook islands. I, 
1. 

271. Potentilla tridentata Soland. Three-toothed Oinquefoil. Very 
common as a rock crevice plant along shores and on small rock Islands. 
I, 1; V, 2. 

272. Potentilla fruttcoaa L. Shrubby Cinqnefoil. Rocks at Scovill 
Point, IV, 1. Occasional shore crevice plant. I, 1. 

273. Comarum paluatre L. Purple Marsh Cinqnefoil. One of the 
most common and most characteristic plants of all bog-lake margins, 
and contributing an important part toward the vegetative bog carpet. 

274. Waldsteinia fragariodeg (Miclix.) Tratt. Barren or Dry Straw- 
berry. Large patches on the side of tlie Greenstone range along McCargo 
trail. II, 3. 

275. Boaa adcularia Lindl. Prickly Bose. Only species of rose found 
on Island. Fairly common around lighthouse clearing and in open 
places. 

Oeran'taceac — Geranium Family. 

276. Qeranium JticknelUi Britton. Bicknell's Cranebill. Rock crevice 
plant: also on rocks with thin soil covering. Few localities only. 1, 
1. 



ECOLOQT OF ISLE ROYALE. 243 

Polygalaceae — Milkwort Family. 

3T7. PolygaUt pauoifolia Willd. Fringed Polygala. Fairly common Id 
rich, moist woods. I, 3; III, 4. 

Empetraoeae — Crowberry Family. 

278. Empetrum nigrum L. Crowberry, Heath-berry, On exposed, 
nearly bare rocka at Scovill Point. IV, 1. 

Anacardiaceae — Sumac Family. 

279. RhMS hirta (L) 8udw. Btaghorn Sumac. Sparingly distributed 
on higher parts of Greenstone. II, 3. 

Bypericacnae — St. John's-wort Family. 

280. Triadenum virginicum (L) Raf. Marsh St. John's-wort. B(^ 
margin of Sumner Lake (III, 5) ; also margin of Forbes Lake (II, S). 
Common. 

VMaceac — Violet Family. 

281. Viola rotundifoUa Michx. Round-leaved Violet. Fairly com- 
mon in ricb, moist woods, especially near I, 6. 

282. Viola lahradorica Schrank. American Dog Violet. Few Speci- 
mens in low moist ground near shore at Siskowit Bay cabin, V, 1. 

2^. Viola areneria DC. Sand Violet. Rocky shore near Siskowit 
cabin V, 1. 

Onagraccae — Evening I'rinirose Family. 

284. Ckamaenerion angttatifoUutn' (L) Scop. Fireweed. Very abund- 
ant everywhere in burnings and clearings. 

285. Epilobiiim Hneare 'iiahl. Narrow-leaved Willow-Herb. Bogmar- 
, g'la Sumner Lake, III, 5. 

286. Epiiobium adetuicaulon Haussk.' Northern Bog Willow-Herb. 
Wet soil near shore of Siskowit cabin, V, 1. 

287. Circaea Alpiiia, Li Smaller Enchanter's Nightshade. Cool 
moist woods at head of Rock Harbor. Not conunon, 

Pt/rolaceae — Wintei^reen Family, 

288. Pyrola chlorantha Sw. Greenish-flowered Winte^reen. Rich, 
moist woods. 

2S9. Pyrola asarifoUn MicUx. Liver-leaf Pyrola. Most common of the 
island pyrolas. Wooda. 

290. Pyrola scciniil<i 1.. One-sided Wintergreen. Ricb, moist woods. 
Not common. . 

291. Pyrola minor L. Jjeseei' I'vroln. Woods. Scarce. 

292. Manages iini flora (L) A. Omy. One-flowered Wintergreen. 
Rather widely distributed in rich, moist woodx, although nowhere abund- 
ant. I, 3, 4; III, 4; IV, 4. 

293. Chimaphila umbeVata (L) Nntt. Pipeissewa. Dry woods and 
fxitosed sunny places, as the pine ridge near Snmner Iiake./ 



"C.oogic 



244 MICHIGAN SURVEY, 1908. 

Monotropaccae — Indian Pipe Family, 

294. Monotropo unifiora L. Indian Pipe. Qnite abundant in ricli, 
dark, moiet woods. Unusnally large, vigorooB specimens in wet mar^n 
of woods beyond Cabin bog at Siskowit (V, 5) . 

295. Bypopityg hypOpitya (L) Bniall. Woods at Siskowit Bay, V, 4. 
Bare. 

Haioragidaceae — Water Milfoil Family. 

296. Hippuria vulgaris L. Mare's Tail. Head of Bock Harbor in 
shallow water. Ill, 2. 

Araliaceac — Ginseng Family. 

297. Aralia nud'waulis Ij. Wild Sarsaparilla. Abundant everywhci-e 
in rich moist woods, where it is one of the characteristic plants of the 
mesophytic forest. V, 4. 

298. Aralia hispida Vent. Bristly Sarsaparilla. One single colony 
on a burned-over island in Bock Harbor. I, 1. 

. VmbelUferae — Carrot, or Umbel Family. 

299. Heracleum lanatum Michx. Cow Parsnip. Light-house clear- 
ing at Bock Harbor; also an old mine clearing along Bock Harbor. 
Budera). I, 7; V, 3, 

300. Cicuta hulbif&ra L. Bulb-bearing Water Hemlock. Occasional 
in bog margins, as II, 5; III, 5. 

301. PaatimMca aativa L. ^Vild Parsnip. Clearing at banning of 
McCargo's trail. 

Cornaceac — Dogwood Family, 

302. Conttia oanadensia L. Low, or Dwarf Cornel. Bunchberry. 
Very abundant in filled bog areas and in moist woods. Also occurring 
abundantly in open places. One of most common herbaoeons plants od 
the island. 

Cornua stolorUfera {See tree and shrub list.) 
Comiis circinata (See tree and shrub list). 

Ericaceae — Heath Family. 

303. Ledum ffroeiilandioum OEder. Labrador Tea. One of the most 
characteristic bog shrubs. Common in b(^ everywhere. I, 6; II, 2; 
II, 5; V, 5; V, 11. 

304. Ealmia glauca Ait. Swamp Laurel. Fairly common in tx^, 
but nowhere so abundant as the preceding. 

305. Andromeda polifolia L. Wild Bosemary. Abundant in nearly 
all the recently filled bogs. 

306. Chamaednphne calyculata (L) Moench. Dwarf Cassandra. 
A very characteristic and common shi-ub of nearly all the ho^. 

307. Arctoataphylos uva^urai (L) Spreng. Bearberry. Very abund- 
ant as a heath plant along the rock shore, and on the thinly-covered 
"rock openings." I, 1, 5 and V, 2. ■ ("^qooI,. 



ECOLOGY OP ISLB ROYALB. 245 

Vacciniaceae — Huckleberry Family. 

308. Vaccinium uliglnostim L. Great Bilberrj-. Bocks at Scovill 
Point. IV, 1. 

309. Vaccinium pennayhanicum Lam. Low, or PennsylTania Huck- 
leberry. Abundant as a heatU plant along shores, and oo nearly bare 
.mountain sides. V, 2; 11, 3; IV, 8, and on some of the small islands at 
Kock Harbor. 

310. Chiogenet hMpiOitla (L) T, & G. Creeping Snowberry. Edge 
of bogs, and on sphagnum hummocks. I, 6; V, 5; II, 2. 

311. Oxycoccua outyroccua (L) MacM. Low Cranberry. Confined to 
wet, UDforested bogs: only fairly abundant. I, 6; II, 2; V, 5, and bog 
near Malone's flsliing camp, V, 11. 

Primulaccae — Primrose Family. 

312. Primula Jtiistammica Miclis. Dwarf Canadian Primrose. A 
crevice plant along the rock shore. Not abundant. I, i; I^', 1; V, 2. 

313. fji/simachia terrct^tris (L) B. 8, P. Bulb-bearing Ixiosestrife. 
Thinly soil-covered rock shore nenr Kiskowit cabin, V, 1. 

314. Naumbcrgia thyrsifolia (L) Duby. Tufted Loosestrife. Margin 
of Siskowit I^ake near head of Trout Creek, V, G. 

315. Trientalia amcricana Pursh. American Star-Flower. Moist, 
rich woodg. Sparingly distributed. I, i; IV, 4; V, 4. 

Gentianaccac — Gentian Family. 

31G. Oentiana andretvuii Griseb. Closed, or Bottled Gentian. A 
few specimens from the Siskowit cabin trail bc^, V, 5. Rare. 

317. Tetrag(mantku8 deftexus (J. E. Smith) Kuntze. Spurred 
Gentian. Moist woods, head of Bock Harbor. Few localities only. 

Jfenyanfftocp«e— Buck-bean Family. 

318. Menyanthes trifoUata L. Buckbean. Abundant in wetter parts 
of bogs : very important contribution in tbe formation oT the "bog car- 
pet." II, 5, III, 5, and bog near V, 2, 

Apocynaceae — Dc^bane Family. 

319. Apocynum andtvaaetnifolium L. Spreading Dogbane. Wash- 
ington Club grounds. 

Conporeu^accof— Morning-glory Family. 

320. Com?olvulus repetis, var. pvbeacens. Pubescent Bindweed. Xearly 
bare sides of the Greenstone along the McCargo Cove trail, II, 3. 

Hydrophyllaceae — Water-leaf Family. 

321. Phacelia franklinii (R. Br) A. Gray. Franklin's Phacelia. Few 
ftpecimens taken on a thinly soil-covered rock elevation near Rock Harbor 
lighthouse. 



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246 MICHIGAN SURVEY. 1908. 

Labiatae — Mint Family. 

323. S!cateUaria laterifoUa L. Mad-Dog Sknllcap. Along flood plain 
of Washington Creek. 

•323. Scutellaria galerhvlata L. Mflnth Skull-Cap. Wet bog mar- 
ginti, as of Bamner (III, 5) and B'nrbeR (II, 5) lafcee. 

324. Prunelia vulgarin L. ^If-heal. Clearings: occurs as a rnderal, 
Washington CInb grounds, II, 1, etc. 

326. Clinopodium vulgare L. Wild Basil. Woods on Oreenstone 
along McCargo trail. Not abundant. 

326. Lycopus americanus Muhl. Cnt-leaved Water Hoar-hound. Wet 
bog margin of Aumner Lake. Ill, 5. 

327. Mentha canadensis L. American Wild Mint. Near water's edge 
at Siskovit cabin, V, 1. Also on Washington Club grounds. 

Sorophulariaceae — Pigwort Family. 

328. Scrophularia leporella Bicfcnell. Hare Firwort. Along Wash- 
ington Creek. 

329. Veronica americana Schirein. American Brooklime. Along 
Washington Creek on low ground. 

330. Castitteja acuminata- (Pnrsh) Spreng. Lance-leaved PaJnted- 
Onp. Common around light-house at Bock Harbor. Fairly abundant 
in open, moist places. 

331. Melampyrum Utiearc Lam. Narrow-leaved Cow-wheat. Fairly 
common on dry. open, to partly shaded [riaces. Exposed Norway Pine 
ridge near Sumner Lake; also occasional in open woods. 

Lentibula riaceac — B I adderwort Family. 

332. Vtricularia mmor L. (?) Ijcsser Bladderwort. In shallow water 
on bog marginal carpet at Sumner Lake (III, 5) ; also occurs at Forbes 
Lake (II, 5). 

333. Pinguicula vulgaris L. Butterwort, B<y Violet. Rock pools or 
moist rocks near water's edge ; fairly common. IV, 1 ; I, 1 ; V, 2. 

Riibiaceac — Madder Family. 

334. Galium spurium L. I^esser Cleavers, Low ground along Wash- 
ington Creek, 

336. Oalium trifloram Michx. Sweet-scented Bed-straw. Woods 
along McCargo trail. 

336. Oalivm trifidum L. Small Marsh Bed-straw. Wet bog mai^ins 
of Snmner (III, 5) and Forbes (II, S) lakes. 

Caprifoliaceae — Honey-suckle Family. 

337. Sambuctfs puhetu 2tlichx. Red-Berried Eider. Along Biskowit 
portage. Fairly common. 

338. Sambucue canadenne L. American Elder. Light-house clearin;^ 
at Bock Harbor. 

339. Yiburnum acerifoUum L. Fairly common in woods. L, S; III, 



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BCOXXIQY OF ISLE ROTALE. 247 

. 340- Tiburtmm paudfoUum Pyl. Few-flowered Cranberry, Moist 
woods: abundant. I, 3. 

341. Linnaea bor€aU$ L. Tn-Ln-dower. Very common in woods and 
partial clearings. Even occurs at I, 1, as a rock crevice plant. Widely 
distributed. 

342. Lonicera dioica L. Glaucous Honey-suckle. Occasional in 
woods. I, 3. 

343. lAmic^a ciliata Muhl. American Fly Honey-suckle. Fairly com- 
mon in woods. I, 3; III, 4; IV, 4, S; V. 4. 

344. Lonicera hirauta Katon. Hairy Hooey-suckle. Rare: in woods. 

345. Lonicera involucrata (Richards) Banks. Involucred Honey- 
suckle. Border of lighthouse clearing, and in open parts of woods. 
Fairly common. I, 3. 

346. Diervilla dicrviUa L. Bush Honeysuckle. A very abundant and 
characteristic shrub of burnings, clearings, and natural openings in 
woods. 1, 2 (and on rock islands in Rock Harbor), II, 1, 3; III, 4; IV, 
4; V, 2, 3. 

Campanulaceac — Bell-Flower Family. 

347. Campanula rotitndifoUa L. Blue Hare-bell. Common rock 
crevice plant on. rock shore and small rock island. Also growing on thin 
soil along the shore. I, 1: V, 2. . 

348. Campanula aparinoides Pursh. Marsh Bell-flower. Common in 
wet bog margins of Sumner (III, 5) and Forbes (II, 5) lakes. 

349. Ij>})eUa khlmii L. Brook, or Ealm's Lobelia. A rocfe crevice 
plant, and on moist rocks near water's edge. Common. 1, 1 ; IV, 1 ; V, 2. 

Chicoriaceae — Chicory Family. 

350. Lactuea pulchella (Pursh) DC. Large-flowered Bine Lettuce. 
IS'oted in one locality only, rock clearing on side of Greenstone along the 
McCargo trail. II, 3. 

351. Hieracium umhcllatum L. Karrow-leaved Hawkweed. On rockn 
or in rock crevices. V, 2. 

352. NaAalus albua (L) Hook. White Battle-snake Root. Woods: 
not abundant. Forest along Riskowit portage, V. 9. 

353. Nabalui raeemosus (Michx.) I)C. Glaucous White Lettuce. 
Rocks and rock crevices along shore. IV, 1 ; V, 2. 

Componitoc — Composite Family. 

354. Etipatoriiim pupuiTiiin L. Joe-P>"e AVeed, or Purple Boneset, In 
moist land near ci'eek, upper end of Rock Harbor, III, .1. 

356, Holidago rirganrea L. var. (?) European Golden-rod. Rook 
crevices, and thin soil on rocks and rock islands. I, 1, 

356. Solidago neglecta T, & G, Bwamp Golden-rod. Fairly cotnnion 
in most of the wet bogs. I, fi; III, 5; V. 5. 

357. Solidago jimcfa .\it. On thinlv Koil-co\-ered i-ock surfaces. Rock 
Harbor, I, S, 

35fi, SuUdago nJiginow Nntt, Siskowit cabin trail Iwp. Y. ."i. 

.139. Aatcr maci-ophfilhiH L. I.aifre-lenved Aster. Very abundant and 
rbaracteristfc in nearly nil clearings, and in natural rock openiuf^ in 
woods. Often forms large colonies. >»jIc 



248 UICHIOAN SURTBT, 1908. 

360. Aster ptarmacoides (Nees) T. & G. Upland White Aster. . 
Fairly common on smal) rock Island, and as a rock shore crevice plant, 
the only crevice aster. I, 1; IV, 1; V, 2. 

361. Aster hirsuticauUs Lindl. Hairy-stemmed Aster. Woods along 
Washington Creek ; one locality only. 

362. Anaphalis margarttacea (L) B. & H. Large Pearly Everlasting. 
Dry soil, and rock clearings along Greenstone (McCat^ trail). II, 3, 
and on Washington Cluh grounds. 

363. Artemisia canadensis Michx. Canada Wormwood. Crevice 
plant along rock Hhore, and on small islands in Rock Harbor. I, 1 ; V, 2. 

364. 8enecio balsaniitae Mubl. Balsam Groundsel. Common rock 
crevice plant on Rock Harbor islands, and elsewhere along the rock 
whore. I, 1; III, 1; IV, 1; V, 2. 



SUMMARY. 

Species of lichens 43 

Species of mosses 38 

Species of Pteridophy tes 35 

Species of Spermatbpbytee 248 

Total 364 



Wheeler, W. A. 

1901. Notes on Some Plants of Isle Boyale. Minn. Bot. Studies, 2, 
pp. 619-620. 
Kuthven, A. G. 
1906. Notes on the Plants of the Porcupine Mountains and Isle Bov- 
ale, Michigan. Bep. Mich. Geol. Survey for 1905, pp. 86-92." 



D,gt,ZBdbyG06<^le 



ECOLOGY OF ISLE ROYALB. 



ANNOTATIONS ON CERTAIN ISLE ROYALE INVERTEBRATES. 

niL CEtAS. C. ADAMS. 

The follovJDg annotated list of Isle Royale invertebrates includes the 
;;roups whioh have not Ireeu made the basis for separate papers b,v 
fipecialists. For the determination of these I nm indebted to the follow- 
ing persons : Prof. N. A. Harvey, the Sponge ; Dr. T. H. Monl^oinery, 
Hair-worms; Prof. Fmnk Smith, Earthworms; Dr. J. Percy Moore, 
Leeches; Miss Ada Weckel, Amphipods; Dr. Harriet Richar^n, Sow- 
bug; Mr. Nathan Banks and Mr. J. H. Emerton, Spiders; Dr. J. W. 
Folsom, Spring-tail; Mr. E. B, Williamson, Dragonflies; Prof. Herbert 
Osbom and Mr. J. B. de la Torre Bueno, Hemiptera; and to Prof. A. J, 
Snyder and Dr. James Fletcher, Lepidoptera. 

In general, in addition to the field notes which include those made 
by Dr. H. A. Gleason and myself, the geographic range has been outlined 
and a selected series of references is given for the convenience of the 
i<tudent of the Michigan fauna. The insects were largely collected by 
Dr. Gleason, the writer, and B. F. Saverj who collected insects about 
tl»e camps, but all members of the party aided in the collection of the 
fipecimens. The field numbers are indicated in parentheses, preceded by 
O in the case of Dr. Gleason's numbers and A in my own. 

An examination'of these lists will reveal their incomplete character, 
as an effort was made to make representati^'e rather than complete col- 
lections of the groups. On account of the small amount of zoological 
information which we possess from Isle Royale it has been thought desir- 
able to make the determinations of the collecfions as complete as cir- 
cumstances would allow. 

FOSIPBKA. 

SpongilUdae. 

Spoitgitla lacuatrk (Linn.). Fresh Water Sponge. A small colony 
of this sponge was found on July 26 in shallow water near the head of 
Bock Harbor (III, 3) by Gleason. Prof. N. A. Harvey examined the 
specimen and makes the following comments: "Small encrusting sponge, 
with a tendency to branch. One branch cylindrical, full of gemmules. 
Skeleton spicules smooth, pointed, slightly curved, nomerous. Dermal 
spicules, few, half as long as skeleton spicules, densely spined, spines 
short, more numerous towards the ends, slightly curved. Gemmules 
with very thin membrane, destitute of spicules, foramen oval. 

"This sponge is evidently a weak form of Spon^Ua laciistris (Linu.), 
It is very close to the paupercula of Bowerbank, Theif can be no 
question about the identity of this sponge, although it is somewhat un- 
usual to find gemmules so well developed in July. It develoi>es its 
gemmules ordinarily very late in the fall. The spicules on the gemmule 
iil)pearto be wholly wanting, and the dermal spicules ni-e not numerous. 



250 MICHIGAN SURVEY, 1908. 

The speeimeo is too small to show the peculiar branching habit very 
distiiictlv." 

A targe (]oantity of this sponge was also secured during 190i in the 
Porcupine Motmtains. Qntonagon county, Mich., by N. F. Macdutf. The 
specimens came from Carp Ci-eek {Sta. VI) in Augnat. Concerning these 
specimens Prof. Harvey also remarks: "Skeleton spicules smooth, 
slightly curved, pointed at both ends. Dermal spicules half the length 
of skeleton spicules, or shorter, spinous spines numerous but short. 
Gemmules not well develoi)ed. TJie branches contaiuing few or none. 
The encrusting portion of the sponge manifesting some. Gemmiile 
spicules wanting. The geniiiiule crust very thin, or altogether wantinp. 
Its branching character is well marked, and the small size of the 
branches indicate the weak form which seldom shows many gemmuleH 
or in which the gemmule sjiicnles are seldom well developed. In con- 
sequence of the size and striking branched habit of this sponge it is the 
one that is usually first found by collectors. I am surprised to find any 
gemniules matured in these s]K)ngeH at the season when these were col- 
lected." 

NEMATODA. 

Gordh'dae. 

Gordius aqiiaticHS robiistiis (Ijeidy) Montg. Hair Worm. This spe<ifs 
of hair worm was fairly abundant; siwcimens were taken at the head 
of Tobin Harbor {IV, 7) among Potamogcton perfoliatum on July '2it: 
among sedgee at the head of a small island in the Harbor (IV. fi), an<l 
on the beach at onr camp on Siskowit Hay (V, 1) August 3 and 6. Both 
sexes are represented in our series, cf, Montgomery, '!>S, pp. 30-31, 

Ocoffraphic Range. Bay of Fundy; Maine; Massachusetts; Maryland; 
District of Columbia; New York; Pennsylvania; Montana; Michigtin ; 
Kansas. The typical form occurs in Europe. 

During August, 1903, Mr. A. G. Rnthvensecnred specimens of (^or'/iif^ 
lincatus Leidy, in a spring in l^e Porcupine Mountains (Station \l), 
Ontonagon county, Midi. cf. Montgomery, 'OS, p. 32. This species seems 
to frequent springs. It is recorded from New York, Pennsylvania and 
Maryland. 



1898. Montgomery, Jr., T, H. The Gordiacea of Certain American 
Collections with Particular Reference to the Korth American Fauna. 
Bull. Mus. Conip. Zool., 32, pp. 23-59. 

1899. Goi"diacea (Hair worms). Amer. Nat., 33, pp. 047-652. 

IIlRt'DlNEA. 

(Ilosaiphonidae. 

Glomphonia coiiiplannta (Linn.). One specimen of this leech was 
taken in a tamarack swamp (V, 5) on August 12. Moore {^01, p. 4!Kti 
states that it aliounds under stones in running water and "feeds chiefiy 
on small snails and annelids." 

Geographic Range. Connecticut; I^ke Ontario; Lake Erie; Ontario; 
Ohio; Slichigan; Illinois; also found in Europe. 

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ECOLOGY OF ISLE ROYALE. 251 

Tfirudinidae. 

MacrobdeUa devoiti (Sav) Verrill. This sixscies wjik quite abuodant 
fit a iiiarshY margin af ^iiiiiuer Lake (III. 5>, whore many specimeiiH 
Ttere tatfeu duriug Jnt.v. This is a larfce speriex and easily racogniKetl 
l)y its dark brown dorsal surfaoe and reddish colored ventral surface. 
They are very gracefa) and con»picuouA objects when seen swimining. 
Odg Bpecimeu was taken upon a frog. Moore (1901, p. 511> reports this 
i'jtecieH as a true blood-sucker and that it is frequently found goi^jed. 

Geographic Jiaiifjc. Maine; New York; Tonnecticut; Virginia; >liph- 
igan; Minnesota; Illinois; Kansas; Nebraska. 

Baemopnia grandii Verrill. One specimen was taken, .July 5, on the 
f.outh shore of Siskowit Luke (V, 6). This is a mud leech and at times 
leaves the water in seaiTh of earthworms (Moore, '01, p. 527), Also 
taken in 1004 l»y Rnthven at Lake Desor (VII, '04), Ruthven, 'Ofi, p. 51. 

Oeofft-aphic Hange. Conne«-ticut ; Lake Huron ; Afichigau ; Lake 
Superior (Ven-ill. '74, p. 6721: VeUowstone I'ark; Kansas; Alaska {E. 
vtarmoratis Mooi-e, "OS, p. 560); Michigan; Illinois; Nebraska. 

En-pohdeUidav. 

HcrpobdfUa lateralis (Verrill) Moore. On the south shore of Siskowit 
Lake (V. G) one »|)eciinen was taken August 5, and another specimen 
was taken Jnlv 14. in shallow water, at the head of Rock UarI)or 

(in. 3). 

The type of this si)ecies came from near the northern shore of Lake 
Superior. 

GeograiiHlc Range. Maine; Connecticut; Lake Huron ; I-ake Hui»erior; 
Colorado, (Verrill. '74, p.*67.">). 

Xephalopsis ohitrnra \'en'ill. Egg capsules of these leeches were taken 
July 27 at Sumner T^ke (III, 5), on a yellow water lily leaf ( .Vj/mpAflci 
advena) and apparently young were associated with them; and similar 
capsules were also found on Potamogeton leaves. \ capsule i^as also 
found in Bock Harbor at Neutson's Resort (IV, 5) also in a swampy 
bayou off Tobin'e Harbor (IV, 3) on July 21. Specimens of 
the leeches themselves were taken in Siskowit Bay on the beach 
at our camp (V, 1), in Siskowit Lake at a small island <m the south shoi-e 
(V, 6) and at the swamp nmrgin of a pond in the yellow water-lily and 
Potamogcton rone (V, 11). Also taken in 1904 by Rnthven at Tjake 
Desor (VII, '04), Ruthven, 'OC, p. Bl. 

Geographic Range. Wisconsin; Colorado (Verrill, "74, p. 074); Mich- 
igan. 

ILErCRENCES. 

1874, A'errill, A. E. Synopsis of the North American Fresh-water 

r^eeches. Rejmrt V. S. Fish Comm. I't. II, 1872-7^, i>p. 

666fi.SD. 
ISflS. Moore, .7. T. ■ The IxH>che8 of the V. S. National Museum- Proc, 

U. S. Nat. Mus. Vol. 21. No. 1160, pp, r)43-.=ili;!. 
1901. Moore, .1. r. The Hirundinea of Illinois. Hull. HI. State I,«b. 

Nat. His. Vol. 5. pp. 479-547. 



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252 MICHIGAN SURVEY, 190S. 

lOOC. HtmdiDea and Oligochaeta Collected in the Oi-eat Lakes 
Region. Bull. U. H. Bureau of Fiaheries, 25, pp. 153-171. 

1906. Rnthveu, A. G. An Ecological Survey in the Porcupine Mount- 
ains and Isle Royale, Michigan. Ann. Rep. 1905, Mich. Oeol. 
Survey, pp. 17-55. 

1902. Ward, H, B. Notes on the Ijceches of Nebraska. Report Neb. 

State Board of Agriculture for 1901. pp. 229-242. 

OLIGOCHAETA. 

Lumbricidae. 

Helodrilua {Allolobophtna) caliglnoaua (Savigny), neater to tffpicvtt 
than to H. {A.) C. trapesoiilea (.\nt. Dug.). These earthworms were 
taken along the beach near the lighthouse at Rock Harbor (I, 1) July 
9, and in a "rock clearing" (I, 2) on July 13, aud also at the outlet of 
Siskowit Lake (V, 9) on August 15, An undetermined Bpecies was 
taken in the shallow humus on the i-ock beach under Cladonia (Y, 2). 

OeograpJiic Rangr. Northern Europe and North America. (Mlch- 
aelsen, '00, p. 483). 

Lnmbricus terrestris Mull. A lai^ specimen was taken fi-om the 
mouth of a Garter Snake ( 7'/tanmo/)/i is sirtalis) found in a clearing which 
was formerly the location of the Ransom settlement (II, 1). The snakes 
were very abundant near the shore in the grass. It is not improbable 
that these earthworms were introduced at this locality. Other specimens, 
doubtfully referred to this species came from the balsam-spruce forest 
<I, 3), where there was also found an Enchytraid. (G. 140). 

Qeographic Range. Europe; New England; Illinois; M^co. (Mich- 
aelsen, '00, p. 512, '03, p. 144). 

REFEBEN'CES. 

1900. Michaelsen, W. Oligochaeta. Das Tierreich. 10 Lieferung. 

1903. Michaelsen, W. Die geographische Verbreitnng der Ollgochae- 

ten. Berlin. 



Gammaridac, 

Eucrangonym gracilis (Smith). Among the dark colored vegetablo 
debris on the north shore of Sumner Lake (III, 5) this species was found, 
July 29; also in a small si ream flowing from a tamarack-spruce swamp 
<V, 5) on August 8, in the Potamogrton and Nf/mphaea adtxna zone of 
a small pond, and also back from the pond in small footprint-like pool» 
of water in the buck-bean ( Maiyanthc» trifoUiita) and sedge zone. (A. 
126, 128, 77,97). 

ByaUlla knickcrhockcH (Bate). Taken at the west end of Rock Har- 
bor in the bulrushes about the mouth of a small stream (III, 3) on 
July 26, (G. 159). Other s|)ecimens were taken in 1904 at I.ake Desor 
('04, VII) on -Vuguat 30; and on water plants in Washington river ("04. 
II) on August 18. 

Gammarus Ihmiarus Smilh. Found in the same conditiopa as the 
above species (Til, 3) and in abundant'. i.,dOO»^?IC • 



ECOLOGY OP ISLE ROYALE. 



KEFEREXCE8. 



Smith. S. I. Tbe Crustacea of the Fresh Watei-s of the United 
States. Report U. 8. Fish Comm. 1872-73. Pt. II, pp. 63T-665. 

Weckel. A. L. The Freshwater Amphipoda of North America. 
Tro. U. S. Nat. Mus., 32, pp. 25-58. 



Oniscidae. 

Ctjlisticiia convexws (PeG,'). One specimen of this sow-bug was taken 
at camp at the Light-bonse (1,7) on July 15. (Q.99). Another specimen 
was taken July 2. at Mackinaw Island, Btraits of Mackinac, Michigan. 
cf. Richardson, '05, p. 609. 

Geographic Range. Jifapsachusetts ; New York; Washington, D. C, 
Ohio; Michigan; Illinois; New Mexico; also Norway; Sweden; Pen- 
marii; British Isles; Germany; Bohemia; Holland; Belgium; fiance 
Turkey. This Hi)ecie8 may have been carried to Isle Royale by man. 

REFERENCE. 

1905. Richardson, H. A Monograph of the Isopods of North Amer- 
ica. Bnll. No. 54, U. S. Nat. Museum. 



JiJtijiicolopkidae. 

RhjfticoJophiis simplex Bks. This mite was found in the thin soil and 
debris beneath the mats of bearben-y in a dry rock clearing (I, 2). 
Banks, '04. p. 30. 

Sydrachnidac. 

Limnochares (xtcndcns Say. This water mite was taken from the 
water in the sedge zone near the open water at Sumner Lake (III, 51. 

Geographic Range. "It occurs in northern states, from Maine to Mich- 
igan, perhaps farther west." (Banks, in letter). 

ARACHNIDA. 

Phalangiidac. 

Lacinitm ohio€ii8i.i Weed. Only three specimens of this Harvest Spider 
were taken, one was found under stones on a shallow soil among the 
Jack Pines (I, 5) in a very hot and dry locality; the second from the 
margin of the sedge none about a pond (V, i 1) , and the third from under 
Cladonia on a rocky beach with a ver>- shallow soil (V, 2). 

Geographic Range. Ohio, Illinois, Weed, '9.3, p. 559; Michigan. 

Dicti/itidac. 

Amaurohiiis beiitictti Blk, A few specimens were tnken under dry 
hark in the hardwoods on a ridge north of the Club House at Washington 
Harbor (V. '04) ; also from under the bnrk of decayed log in the maples 



254 MICHIGAN SURVBY, 190S. 

on tlie Deeor Trail (III, '04), In the moist vpftetnble mold in the baleam- 
sprfice forest (I, 3) and under tUe bark of dead trees near the rock 
clearing at the camp on Siskowit Bay (V, 3), One was found in the 
mouth of a Winter Wren phot by McCi-eary (il, 1). In the clearing at 
Benson Brook (II, 1), and in the "i-ock clearing" at camp on Siskowit 
Bay (V, 3). 

Geographic Range. Canada, Marx, '00, p. 510; Porcupine Mountains, 
Michigan. 

Drassidac. 

Dranmiit negleetuit Key8,=/>. taccatuR, Emerton. '02, p. 6. One speci- 
men was taken from under a stone, upon the jack pine ridge where the 
soil was very shallow and the heat intense during the middle of the day 
(I, 5). It was enclosed in a rather comi>ac-t close fitting web. Also taken 
from the margin of a pond among the vegetation (V. 11). One col- 
lected on the rock ridge north of the light-house (I, 2) was in a small 
pocket-like web about 2 by 2.5 cm. in a cavity under a flat stone, sur- 
rounded by moist soil, at a depth of about 6 cm. (Gleason). 

Geographic Range. New Hampshire, Slosson, '98, p. 247; Michigan; 
Hist. Columbia, Marx, '96a, p. 154. 

Gnaphoita hnimalis Th. The only specimen (G. 22) was taken in a 
small Cladonia clearing on the north side of Conglomerate Bay (I, 2) 
near the beach. 

Geographic Range. Labrador; Anticosti Island, Quebec; White Mts.. 
above tre^ limit, N, H, ; Ithaca, >'. Y.; Massachusetts; Colorado; 
I^ggan, Alberta, cf. Emerton, '94. p. 413; Banks. '9S, pp. 417, 421; 
Marx, '90, p. SOS. It is not unlikely that the New York and Massaehu- 
sets localities are from "boreal islands" — swamps or cool ravines, and 
are thus outliers from the principal range of this form to the north- 
ward. Alaska, Marx, '9([a, p. 189. 

A<p-iopi(iac. 

Linyphi^t phri/giana Koch, One specimen was taken at our camp at 
the Light-house (1, 7). Emerton, '02, p. 141. 

Geographic Range. Gaspe, Quebec; Mt. Washington, K. H.; Maine; 
Mass. ; New York ; Connecticut ; Colorado ; - Rocky Mts. of Canada ; 
Calif.; probably all over the United States and Northern Europe. 
Emerton, '94, p. 409, and '82, p. fi3; Banks, '95, p. 425. 

Tetragnatlut ejienaoi Linn. One specimen was taken on the window 
sill at the Lighthouse (I, 7). Emerton, '02, 201. 203. 

Geographic Range. Labrador; Mass.; New York, Marx, '90, p. 552 
Anticosti Island, Quebec; Saskatchewan River; \Vliite Mts., N. H.: 
Adirondack Mts., N. Y.; Connecticut; Dist. Columbia; Alaska; Siberia, 
Lapland; EuroiM.', Emerton. '04, p. 40C; Beaver Island, Mich., Pettitj 
'01. p. 39; Calif.. Collidge, Civn. Ent. 39, p. 376. Marx. '96a, p. 196. 

Epeira patagiata Clerck. Taken in the cassandra zone of a tamarack 
swamp (V, 5). Emerton, '02, p. 160. Comstock, '03, p. 38. 

Geographic Range. Lapland; Labrador; New Foundland; New Hamp- 
shire; New York; Pennsylvania; Maryland; District of Columbia; 
Vii^inia; Illinois; Alontreal. Anticosti Island. Quebec; Lake of tho 
Woods; Saskatchewan River; Colorado: British Columbia; Washing- 



EKIOLOGY OP ISLE ROYALK. 255 

ton; Oregon; Sitka, AiaBka; Europe. Enierton, '04, p. 404, and '84, 
p. SO.'i; SloBson. '08, p. 248; Banks, 'S5, pp. 417. 425. Marx. '96a, p. 
104. It eeems probable that the most southern localities of this species 
are confined to some restricted habitat. 

Thommdae. 

Ebo lalithorax Keys. One specimen was taken on the beach near the 
Lighthouse (I. 1). Emerfon. '02, p. 38. 

Geographic Uange.. Mass.; New York, Emerton, '92, p. 378; Virginia; 
District of Columbia; Maryland; Utah. Marx, '90, p. 558. 

Chibionidae. 

Cliibiona riparia Koch. One specimen was taken in the vicinity of 
TobiD Harbor (IV). 

Geographic Range. Maryland. Marx. '90, p. 512; New Hampshire, 
8ioB80n. '98, p. 247; Colorado, Banks, '95, p. 422. Dist, Columbia, Marx. 
'9fi. p. 155. 

Agelenidae, 

Tcgeimria derhami (Scop.). Found in the hardwood forest on the 
ridge east of the tamarack swamp (V, '04) back of tbe Club House at 
Washington Harbor. Emerton, '02, p. 06, and Marx, '90, p. 516. 

Geographic Range. I^brador; Gaepe, 'Quebec; "A common house 
spider in North America and Europe," Emerton, '94, p. 411; New Hamp- 
shire, Slosson, '98, p. 247; Colorado, Banks, '95, p. 422; Calif., Collidge, 
ran. Ent., 39, p. 375. Marx. .'96a, p. 190. Dist. Columbia, Marx. '96, p, 
155. Indiana, Fox, '93, p. 268. Probably introduced (Emerton). 

COflotes sp. A s]>ecimen, too young for epeciflc identification, was 
taken in a rock clearing (I, 2) near the Lighthouse. It spins a pocket 
just about large enough for its body beneath loose rocks. 

Oiciirina arniata Keys. Where the soil was very thin on the jack 
pine ridge (I, 5) one specimen was taken under a stone and a female 
was found with a white disk-like cocoon containing a lat^ number of 
young white spiders. A specimen belonging to this genua was found 
along the trail through the balsam forest in leaf mold at Siskowit Bay 
IV. 4), hnt it is too young for specific determination. It spins a small 
pocket-like web beneath flat stones. Pi-equents the dead leaves of 
forests (Emerton). 

Geographic Range. Labrador; New Hampshire; Penn. ; District of 
Columbia; Virginia; Lalie Superior; Minnesota; IlliDois; Colorado, 
Slarx, '90. p. 516 ; '96«, p. 190, '92 ,p. 155. 

Pisauridae. 

Dolomcdca idoneus Montg. This large spider was taken at the log 
cabin of the Washington Club at T^ike Desor (VII. '04) (A. 139). 

Geographic Range. Lake Champlain; Conn. (Emerton) ; Penn.; 
Michigan. 



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256 MICHIGAN SURVEY, 1908. 

Lycoaidae. 

Lycosa frondicoJa Enier. One Hpecinien was taken about camp at 
the Light-house (I, 7). 

Geographic Range. Conn., Marx, '90, p. 561 ; New Hampshire ; 
Slosson, '98, p. 248; Penn., Stone, '90, p. 426; Michigan; l>ist. Columbia, 
Marx, '96, p. 160. Indiana, Fox, '93, p. 269. 

]jycoaa pratensia Emer. On a gravelly beach near the Light-houee (I, 
1) dragfiiDg an egg-caae with it, in the rock clearing (I, 2) and on 
the dry Jack Pine Ridge (I, 5) were the situatioDB in which this species 
was taken. Emerton, '02, p. 69. 

Geographic Range. Anticosti. Quebec ; White Mts., N. H. ; MasB. ; Conn. ; 
Porcnpine Mountains, Mich.; Lake of the Woods; I^aggan, Alberta. 
Marx, '90, p. 563 and Emerton, '94, p. 422. 

Lycosa kochi Keys. This was an abundant species, found upon the 
beach near the Light-house (I, 1) ; in a rock clearing adjoining the 
beach (I, 2) (G, 71), and about the camp on Siskowit Bay (V, 3) where 
one had been captured by a wasp. One with an egg-case attache*! 
was buried under half an inch of soil on a rock ridge (I, 2). Emerton, 
'02, p. 74. 

Geograph\c Range. Mass.^ Conn.; Penn.; New Jersey; Bist. 
Columbia; Michigan. Emerton, '^, p. 486; Stone, '90, p. 426; Marx, 
'96. p. 160. 

Pardosa glacialis Thor. Found quite abundantly running about over 
the wet sphagnnm on the north shore of Forbes Lake (II, 5) ; man.r 
were carrying cocoons. Others were secured among the open cassandra. 
tamarack and spruce zone about a pond (V, 11). Several other speci- 
mens were taken running about with cocoons in the Cladonia-Jitniper 
procumbena and bearberry belt on a sloping rock beach (V, 2). Emer- 
ton, '02, p. 80. 

Geographic Range. Gi-eenland; I^abrador; White Mts., K, H.; Massu- 
chusetts (Emerton) ; Conn. ; Laggan, Alberta ; Emerton, '94, p. 
425; New Mexico,, Psyche, 9, p. 123, Marx, 96a, p. 197. Found 
near Ann Arbor, Mich, in a tamarack swamp (Miss Jean Dawson), 
thus clearly indicating the boreal island character of such a habitat. 

Pardosa- groenlandica Thor. This was apparently the most common 
Kpeoies of spider .collected. Many were found running about over th« 
rocky, gravelly or sandy beaches (I, 1) with cocoons (G. 16, 38, 39, 30. 
46). A specimen was also taken on the open heath beach on the south 
shore near Siskowit Bay (V, 2), Emerton, '02, p. 79. 

Geographic Range. Greenland; Labrador; Anticosti Island, Quebec; 
White Mrs,, above the tree limit among stones (Emerton), N, H.; Lake 
of the Woods; Laggan, Alberta; Idaho; Colorado; Washington ; Oregon ; 
Alaska. Emerton, '94, pp. 400, 423; Banks, '98, p. 16, '95, p. 430; Marx, 
96a, p. 197. 

Pardosa stemalis Th.^=hUeoJ<i Em. .411 the specimens of this specieH 
were taken in open areas in a small CJadonia clearing near the beach 
on the north side of Conglomerate Bay (I, 2), and crawling about over 
the nests of the ant Foi-mica fusca (V, 3) ; the heath beach near Sis- 
kowit Ba.v (V, 2) (A. 1071 and in the open area about our camp on 
Siskowit Bay (V, 3) (G. 22ri). Most of the females carried cocoons. 



E(X)UX}Y OF ISLE ROYALE. 257 

Geographic Range. ColoraiSo, Banks, '95, p. 429. Mt. Washington, 
N. H, On mosses and lic&ens (Emerton). 

Pardoaa lapidicina Emer. This was also a beaoh spider (I, 1) (G. 
25, 38 (2)). The cocoous are very large Id proportion to the size of 
the female and are flattened. Emerton, '02, p. 78. Lives among stones 
(Emerton). 

Geographic Range. Gaspe, Qviel>ec ; Massaehnsetts ; Connecticut ; 
Pennsylvania. Emerton, '02, p. 79, states that this species "lii'es 
nmong stones in the hottest and drvest places from Connecticut to 
Labrador." Man, '90, p. 565; Stoned '90, p. 431. 

Pardoaa tachypoda Thor, Found running over the bare rocks on the 
top of the jack pine ridge (I, 5), and carrying cocoons (A. 21). Emer- 
ton, '02, p. 81. 

Geographic Range. Labrador; Mt. Washington, X. H.; Adirondack 
Mt8., N. Y.; Manitou, Colo. Emerton, '85, p. 493, and "94, p. 401. 

Salticidac — AttiAae. 

Phdddippus borealig Bks. One specimen of this jumping spider was 
found ander loose stones on moist earth on the jack pine ridge (I, 5). 

Geographic Range. New Hampshire, Banks, '95, p. 96, Blosson, '98, 
p. 249; Maine; Kew York, Banks (in letter). 

Habitat Preferencea. Reviewing the preceding list of mites and 
spiders the following habitat preferences appear to be Indicated: 

I. Beach, rocky, bouldery or sandy. 
Lycoaa pratensia (also dry openings). 
LJ/coaa kocki. 

Pardoaa groenlandica. 

Pardoaa aternalia (also dry openings). 

Pardoaa lapidicina. 

Pardoaa gUicialis (also in sphagnum bogs). 

II. Dry openings, rock or shallow soil, heath-juniper — Cladonia jilaut 
society represented by 1, S; V, 2. 

Rhyncolophna aimplcx. 

Laciniua ohioejiaiH (in swamp also). 

Onaphoaa brumalts. 

Draaaua ncglecUis (in swamp also). 

Coeletea sp. 

Cicurina arcuata. 

Lycoaa piatensts, 

Lycoaa kochi (also beach). 

Pardoaa atemaUs (on beach also), 

Pardosa tachypoda. 

Pardoaa glacialis. 

Phidippua borealia. 

III. Wet places — as sphagnum swamps, 
Epeira patagiata (cassandra zone). 
Pardoaa glacialia (also dry openings). 
Draaaua neglectua (also dry openings). 
Luciniia ohiocnaia (also dry openings). 



rfbyGOOgIC 



25S . M[CHIGAN SURVEY, 1908. 

IV. MeBophytio forest— balsam -Bpruce or hardwoods. 
Amarobius bi'itnetti. 

Tegenaria derhami. . 

V. About Camps. 
Linifphia phrygiana. 
DolomeCeg idoneua. 

From tlie above tubiiltition it is evident tlint, if the coUectionn are 
representative, moBt of tbe spiders prefer the open places, the beach, 
rock openings or oiien parts of .swamps, tbe most marked preferenee 
Iteing for dry opening. It thus appears that as the forests encroach 
upon these areas the spider habitats become more restricted. The 
genus Pardona seems quite characteristic of the open places. The 
general Arachnid successions are thus suggested in outline as follows: 
from beach types and rock openings to the forest; inland from the 
aquatic types and swamp forms to the forest. Particular attention is 
directed to the following habitats which deserve special attention for 
their bearing ou succession; these ai-e the bii-cb-a8i>en border and 
clearing wciety, and ^chides or openings in the forest and the forested 
swamps. An examination of the literature clearly shows that tbe 
habitats of spiders have received bnt little attention. This is an ex- 
i-ellent field for study and one certain to give interesting and valuable 
results. 

Geographir; "Xotcs. The following nine species of Isle Eoyale spiders 
Lave been recorded from Labradoi*: Gnaphoaa brumalis, Tetragnatha 
cxtcnaa, Tegenaria derhami. Ciairiiia arciiata, Epeira patagiata. Par- 
dosa glacialia (also Greenlaud), Pardoaa groenlandica (also Greenland), 
J'artloaa lapUUciua and Pardosa tachypoda. 

The following fourteen species Iiave l>een reported from New Hamp- 
shire: DranmiJt wylcctua, Onaphosa briftnalis, Linyphia pkrpgiana, 
Tetragnatha cxtenaa, Epeira patagiata, Clubiona riparia, Tegenaria der- 
hami, Dolunirdea idonem, Lyvom fnntdieola, Lyvoaa p^-atenaia. Par- 
doati glacialia, Pardoaa gnynhiidk'a, Pardosa taehypoda and Phidippus 
borealia. 

The following s{>ecies are found in the mountains of Colorado : Onap- 
hosa bni-malia, Epeira patagiata and Pardoaa groeii land tea, (Banks, '95). 
They also occur in New Ilainpxhire, Labrador, and fre(iuent oi«n places. 



ukveuksceh. 
Banks. N. 
l,s!i:t. The Spider Fauna of the rpi)er Coviiga Lake Basin. Pro. 

Acad. Nat. «ci., Philn. 1S!I2. pp. 11-81. 
18i>4. On the lo'cosidae of Colorado. Jour. X. Y. Ent. Soc, 2, pp. 

40-52. 
lS9u. The Arachnida of Colorado. Ann. N. Y. Acad, of Sciences, 

pp. 417-4:14. 
ISllii. Some New Attidae. Caw. Ent., 27, pp. !W-102. 
IS'JS. Aracbuida from the Malaspinn Glacier, Alaska. Ent. News. 

!l, p. 1(). 
1904. A Treatise on the Acarina or Mites. Pro. V. 9. Nat. Mus., 

28. PI-. 1-114. 
1907. .\ Catalogue of the Acarina. or Mites, of the Uaited States. 

Pro. U. S. Nat. Mns., 32,'pp. 595-625. i,000<^[e 



KCOLOGY OF ISLE ROYALE. 259 

C'hamberlin. K. V. 

1908, Kevisiou of North Aiiiericiin Spidei-s of the Fauiilv Lvoonidao. 
Pro. Acad. Nat. Sci., I'hila., lOOfci, pp. l58-ai8. 
Comstock, J. H. 

1903. A Classification of Xorth American t^piders. Ithaca, N. Y. 
Eraerton, J. H. 

1882. New England Spidera of the Family Therididae. Trans. Conn. 
Acad. Arts and ScienceB, 6, pp. 1-86. 

1884. New England Spiders of the Familv Epeiridae. Trans. Conn. 

Acad., 6, pp. 295342. 

1885. New England Lycoeidae. Trans. Conn, Acad., 6, pp. 481-505. 
1888. New Kngland Spiders of the Family Ciniflonidae. Trans. Conn. 

Acad., 7, pp. 443-458. 
1890. New England Spiders of the Families I>ras9idae, Agalenidae 
and Djsderidae. Trans. Conn. Acad., 8, pp. 166-206. 

1892. New England Spiders of the PamilT Thomisidae. Trans. Conn. 

Acad., 8, pp. 3S9-381. 
1895. Canadian Spiders. Trans. Conn. Acad., 9, pp. 400429. 
Marx, G. 

1890. Catalogue of the Described Araneae of Temperate North 

America. Pro. U. S. Nat. Miis., 12, pp. 497-594. 
1S96. A Revision of the Araneae of the District of Columbia. Proi-. 

Ent. Soc. Wash., 2, pp. 148-ltil. See also, ^. pp. 199-201. 
1896a. A Contribution to the Stndy of the Spider Fauna of the 

Arctic Regions. Proc, Ent. Soc. Wash., 2. pp. 196-200. 
Montgomery, T. H. 

1904. Descriptions of North American Araneae of Families Lycosidae 

and Pisauridae, Pro, Acad. Nat. Sci. Phila., 1904, pp.'261-323. 
Pettit, R, H. 

1901. Insect and Animal Life on the Upper Peninsula Experiment 
Station. Bull. 186, Mich. Agr. Exp. Sta., p. 28-42. 
I'iersig, R, 

1901. Hrdrachnidae. Das Tierreich. 13 Lieferung pp. 1-272. Ber- 
lin. 
Rutbven, A. G. 

1906. Spiders and Insects from the Porcupine Mountains and Isle 
Royale, Michigan. Ann. Rep. Geol. Sur. Mich, for 1905, pp. 
. 100-100. 
Stone, W. 
1890. Pennsylvania and New Jersey Spiders of the Familv Licosidae, 
Pro, Acad. Nat. Sci. Pbil., 1890, pp. 420-434. 
SlosBon, A. T. 

1898. List of Araneae taken in Franconia, New Hampshire. Jour. 
N. Y. Ent. Soc, 6, pp. 247-249. 
Weed, C. M. 

1893. A Descriptive Catalogue of the Harvest Spiders (Phalangiidae) 

of Ohio. Pro. U. S. Nat. Mus., 16, pp. 543-563. 

Useful Bibtiograpbies. 
Banks, N. 

1900. A liist of Works on North American Entomologj. pp. 77-83. 
Bull. 24, r. S. Div. of Ent. V. S. Dept. of Agriculture>o|c 



260 MICHIGAN SURVEY. 1908. 

XJndenvood, L. M. 
1887. The Progress of Axaclmology in America. Amer. Nat. 21, pp. 
963-975. 



Entomobryidac. 

Tomofvnis nigor Beurl. Specimeos of this spriag-tail (G. 140) were 
ttikCD J«I,v 24, in the balsam -spruce forest (I, 3). 

This species has an extensive range; through Europe, Siberia. Alaska, 
California, Minnesota, Isle Eovale, Michigan, cf. Folsom, 1902, Proc. 
Wash. Acad. Sciences. IV. p. 97; Guthrie, 1903, The Collembola of 



Minnesota, p. 



HEMIPTERA. 



Coriia sp. N.vmphs were taken in the rock pools on the beach (I, 1> 
(G. 7.1, 74. 7n) ; in Rummer Lake {III. 5), (G. 175) ; and in the pond 
in a tamarack -spruce s\ram]>s (V, 11) (120. A). 

Belo8tomatidae. 

Bclostoma sp. Nvmphs, but no adults, were secured at Sumner Lake 
(III, 5) (77 A.) ; and in the pond in the tamarack-spruce swamp (V, 
11) (126 A) on August 16. 

Saldidae. 

Salda ligata Say. On August 10 these shore bugs were running about 
in numbers on the bare rock beach, just beyond the reach of flie waves, 
on the south shore near the mouth of Siskowit Bay (V, 2) (106 A). 
"Common over eastern United States. These specimens are darker 
than most in my collection but seem to agree very perfectly with de~ 
Bcriptions of Say and Uhler." H. Osborn. 

Eydrohat'tdae. 

Gerris remigis Say. This member of the surface film fauna was found 
ou July 14 on the bulrush zone and delta near the bead of Rock Harbor 
(III. 3); in rock pools at Scovill Point on July 19 (33 A), where 
it was very abundant and represented by unwinged adults and nymphs; 
in rock pools on the south shore near Siskowit Bay (V, 2) on August 
9 and 14 by adults and nymphs (103 A. 117 A) ; and on LakeDesor (VI [. 
04) on .\ugu8t 20 (139 A) by both young and adults. 

Oerris mfoscvMlattis Latr. This species of water strider, in com- 
jiauy with O. remigis. was taken from rock i)ooIb at Scovill Point (IV, 
I) on July 19 (33 A), The specimens are winged. Also from the rock 
licach pools on the south shore (V, 2) on August 14 (117, A), and from 
a pond surrounded bv a tamarack-spruce bog (V, 11) on August 16 
(126 A>. 

Gerris marginaitiK Say. This third species of strider was found only 
at Sumner Lake (111. 5) ou July 28 (G. 175), and was represented by 
iivmphs and adults. , -- i 



ECOLOGY OP ISLE ROYALB. 261 

Oerria 8p? Nymphs were taken io a rock pool at Tobin Harbor 
tlV, 2) on July 19 (30 A), and oq a small stretim at the head «f a 
bog (V, 5) near Siskowit Lake {95 A). 



Aradus abbas Berg. One Hpecimen was taken on August 7 about 
ramp on Siskowit Bay (V, 3). 

Lygaeidae. 

],ygva pratensis L, Taken about camp at the Light-houee (I, 7) 
July 11. "One of the dark colored yarieticB. The speciea has a wide 
distribution in both Europe and North America." H. Oeborn. 

Pamera sp. Also taken about camp at the Light-house (L 7) on July 
2S. "Apparently an undcscribed species." H. Osborn. (136 A). 

Cicadidae. 

Tibicem rimoaa Say. var. This cicada was abundant upon the hot 
jatk pine ridge (I, 5) and among the birches at its base. Adult 
specimens and a nymph skin were taken ou July 8 (O. 28), 10 (G. 44) 
and 17 (G. 108) ; at Neutson'M resort (IV. 5) ou'jnly lil (44. A) ; in the 
i-ock clearings (I, 2) on July 13 (G, 08) ; on the rock ridges on the 
MeCai^oe trail (II, 3) on July 25 (G. 147)*; in the clearing about the 
Light-house (I, 7) on July S; and near the head of Rock Harbor (III, 6) 
on July 17 (G. 111). The sjiecies thus showed a decided pi-eference for 
the open dry situations. Prof. H. Osborn writes concerning the speci- 
mens sent to him for determination ; "These specimens agree closely with 
a variety of rhnosa occuiTing at Ft. Bridger, Wyoming." 

Concerning the habits of this sjiecies Osborn ('HO, p. 190) states that 
in northwestern Iowa it occurs "on prairie land remote from timber, 
thus indicating a habit quite different from the other members of the 
genus." The occnrrence ot this gjiecies in the more or less open place 
upon Isle Royale is thus in harmony with its prairie habits and shows 
that these rock openiugs may contain not only forms of northern faunal 
affinities but also those from the western plains. The occurrence of 
these western species in open places in the northern forest region is 
aualt^ous to the soiithei-n prairie sjwcies found in dry or sandy places 
in the south-eastern forests. 

JaHsideuc. 

RnthoKcopiis pitivi Pn>v. This leaf hopjter was taken from a rock pool 
uiion a small island in Tohin Harbor (IV. 2| on July 19. 



Ortheaia ap. This bark louse was taken July 1!). at S<'ovill I'li 
(IV, 1). '"rndescribed so far as I can discoypr." H. ttsborn. 



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MICHIGAN SURVEY, 1908. 



RErEREXCE3. 



Gillette, C. P., and Haker. F. 0. 

1895. The Hemiptera of Colorado. Bull. 31. Tech. Sep. I. Colo. 

Agr. Exp. StatioD. 
Osborn, H, 

1896, Observations on the Cicadidae of Iowa. Proc. Iowa Acad, of 

Sciences 189"), III, pp. 194-2<Kl. 
.1900. Remarks on the Ileniipteroua Fauna of Ohio, with a Ppelini- 
inary Kecord of Species. Eighth Sep. Oliio State Acad. Bci.. 
pp. 60-79. 
1901. Additions to the List of Ohio Hemiptera, ?finth Rep. Ohio 
Acad. Sci.. pp. 3(»-37. 

1904. A Ftirther Confribntion to the Hemipteroiis Fauna of Ohio. 

Ohio Naturalist, 4, pp. 99-103. 
Torre Bueno, J. R. de la 

1905. A List of Cei-tain Families of Hemiptera Occurring uititiii 

Sereutv Miles of Sew York. Jour. !N. Y. Ent. Soc., 13, jip. 
29-47, '{also, 12, pp. 251-253). 
VanDuzee. E. P. 

1894. A List of the Hemiptera of Buffalo and Viciuitv. Bull. Buff. 
Soc. Nat. Sci., 5, pp. 167-204. 

XEl-ROPTEIt.V. 

Myrmeleon tdae. 

Myrniclcott hmtuiriiJatiis DeG. Aut-lion. An nppaientiv full gi-own 
ant-lion was taken on one of the bare burued over ridges on the McCai^oe 
Trail (II, 3), on July 23. None of the adult insects were seeu and this 
was the only specimen and funnel obsen'ed. 

Geographic Range. Maine; Keene, N, H.; Isle Royale, Mich.; Illinois; 
Washington, D. C; North Carolina; Texas; Colo.; Oregon; Calif, cf. 
Psyche, 9, p. 95. 

ODONATA — DRAGON f LIES. 

The Dragonfly collection seciii-ed by the party is an interesting one in 
several respects as will be seen by reference to the geographic distribu- 
tion of the varions species. MokI of the specimens were collected by 
H. A. Gleason; the writer and other nieuibera of the party also secured 
some. Vi'e are indebted to Mr. E. B. ^Villiamson of Bluffton, Ind- 
iana, for the determination of the aeries. 

AgiioiUdac. 

Lcatnn utigtiinilntw* Hag. A single female, apparently of this species 
was found in the clearing at Neutson's resort (IV, 5) July 21. 

Geographic Range. Nova Scotia; Quebec; Maine; Mass.; Rhode Is- 
land; New York; Penn.; New Jei-sey; Ohio; Indiana; Tenn, ; Illinois; 
Iowa; Missouri; Wyoming; Montaua; California; Corunna, Ann Arbor, 
Porcupine Mts., Mich. 

Xehalvnnia irene Hag. Two males of this species were found in or 
near a tamarack-spnice-spiiagnum swamp (I, 6) on July 10. This is a 



ECOLOGY OF ISLE ROYALE. 263 

swamp-land species which iisimll.v flieb slowly among the low voRPtaHon. 

Geographic Range. Ontario; Maine; New Hampshire; Slaxs. ; New 
-lersey; Florida; New York; Teiin. ; Ohio; Indiana; Illinois; Iowa; Wis- 
consin; South Dakota; Ann Arbor (Kavunangh I/ake, VII, -i, Wik 
Porcupine Mts,, Mich. 

Enallagma ej^ulans Hag. One male was taken flying over the water 
near the boat landing at Lake Hemr (VII. '04) on .Viigiist 2(1. 

Qeographic Range. Ontario; Maine; New York; I'enn,; Maryland: 
District of Columbia; Virginia; Ohio; Indiana; Illinois; Missouri: 
Texas; Comnnn, Huron Kii'ei', near Portage Lake, Aug. Ill, '03, Wash- 
tenaw Co., Kavanangh Lake, July 10. '03, Chelsea, Mich. 

Enallagma hageni Walsh. This species was quite abundant about the 
sedge mat^iu of Sumner Lake (III, 5) where eight males were taken 
between July 24 and 2!). Three other males were taken on August 16 
in the cassandra and sedge zone aliout a pond neiir Siskowit Bay (V, 11). 

Geographic Range. Newfoundland; Quebec; Ontario; Maine; New 
Hampshire; Mass.; New York; Delaware; Maryland; Ohio; Indiana; 
Illinois; Iowa; Missouri; Wisconsin; South Dakota; (July 10, '03, Kav- 
anangh Lake) Chelsea, Porcupine Mts., Mich. 

Oomphidac. 

Ophiogoinpkiis cotiibriniif Selya. One female was taken, .Vugust 7, 
flying low, at the outlet of Siskowit Ijake (V, !>). This stream falls rap- 
idly providing a rapid water habitat which furnishes favorable condi- 
tions for most Uomphids. The streams on the island are small and How 
through much swamp land so that there is but a slight development of 
the rapid water habitat. 

Geographic Range. Hudson Bay; Quebec; Maine; New Hampshin', 

Aesehnidnr. 

Anax jimins Drury. Nymphs only were taken. Cf. Needham i-epoit. 

Oeographic Range. Central America northward to Newfoundland and 
Alaska; Bermudas; West Indies; Hawaiian Islands; Tahati; China; 
Kamtsehatka. This is a very extensive range, cpiite exceptional among 
Dragonflies, and perhajts only snrimssed by the cosmopolitan Pantala 
flavescetui. 

Aeschna sitchensiit Hag. One female was taken along the i-oad through 
the hardwoods on the Desor trail (III. '04) on August 21. (Psyche, 
1890, p. 35.^). 

Qeographic Range. Sitka. Alaska ; Saskatchewan ; Newfoundland ; 
(Williamson, '06, p. 135); Pequaming. Michigan, Calvert, Knt. News. 
15, p. 288. 

Aeschna species? The remaiaing Aiwchnids cannot be satisfactorily 
determined at present. A male was taken in the sedge and cassandra 
zone bordering a pond (^',11) on .\ugnst 10. A male, which had been 
jtatrolling a small stream flowing from a tamai-ack swamp (V, 5), was 
taken on August 8, -\ teiieral female was taken .Inly 13 on a i*ock ridge 
(1,2) and in the clearing about the camp on Siskowit Bay (V,,"?) ,[231|; 
one male was taken .\Hgust 15. and a female on August 12, Kighf males 
were taken at Snmner Lake (III, 51 between July 20 and 2it. A female 
was taken in a rwk clearing near the lighthouse (I, 2) on July 13 ('*{)-)- 



264 MICHIGAN SURVEY, 1908. 

1'i'i-c-c' nymphs skins were found nt tlie margin of the sedge zone of 
Stunner I^ake (III, 5). ronipare Walker, 'OS, who hds examioed tlie 
Me Royale Bpecitnens. 

CorduUdae. 

Tetragoneuria apinigera Selys, One specimen wag captured in a low 
rock opening at the shore, near the head of Rock Harbor (near III, 3) 
OD July 14; and a female was taken floating npon the water in the 
west cove at the head of the Harbor (III, 6). Tbe third Bpecimeu, a 
female, was secured from n rock ridge near the head of the Harbor 
near III, 2, July 21. (132.) 

Geographic Range. Maine; Sew Hampshire; Mass.; tieorgia; Detroit, 
.Mich,; ^'aucouver Island. 

CorduUa ahurtleffi ScuAd=a€nca I/. Three specimens, two males and 
one female, were taken at the edge of the water in the sedge zone on 
llie north side of 8umner Lake (III, 5) on July 2<.t. (184. 78A.) 

Olographic Range. Nova Scotia; Newfoundland; Ontario; Maine: 
New Hampshire; Penn. ; Saskatchewan; Fort Resolution; Mackenzie; 
British Columbia; Alaska; Northern Asia; Europe; Algeria. 

Somatochlora elongata minw Calvert. Only one male of this interest- 
ing species was found, it was flying about the mouth of a small 
stream at the head of Rock Harbor (III, 3) (165) on Julv 26. Cf. 
<;:>lvert, Ent. News 1898. !l, p. 87. 

Oeograph ic Range. Quebec ; Maine ; New Hampshire ; Michigan ; 
Wyoming. 

LihelUiUdae. 

Celithemia eponina Hagen. Nymphs only taken. Cf. Needham report. 

(I'eographic Range. United States east of the Rocky Mountains and 
southern Canada (Ontario). 

Leucorhinia hudi*onica Selys. Three females were taken, in the sedge 
zone of a tamarack swamp (V, 5) on August 8. (i)6A). Sympctrum 
'•htrusum occuri-e<i abundantly iu the siime locality. 

Ocographic Range. Quebec; Nova Scotia; New Brunswick; New- 
foundland; Maine; New Hampshire; Mass.; Lake Winnipeg; Sask- 
atchewan Kiver; Fort Resolution. Mackenzie; Alberta; British Colum- 
bia; Alaska. 

Leucorhinia proxitna Calvert. About the margins of Sumner I>ake 
(III, 5) these dragonflies were very abundant on July 18, 24, and 
29, and 19 males and 5 females were taken in the sedge zone. About the 
lake this zone was quite extensive, as shown by the photogi-aphs. Most 
:if our collecting of insects was done at the northeast end where, with 
the aid of boots, an excellent swamp collecting ground was found. The 
ground was very wet, and spongy, and treacherous in places, on account 
<)f these soft spots. This species also occurred abundantly about the 
margins of a similar pond near Siskowit Bay (V. 11), where it was 
iisMicinted wilh Eiiallagnia hoydii. Acxrhiia and ygiiiiK'triim rubivuii- 
tliilitm obtruHum. There is an interesting rorrelatiou lietween the geo- 
^i'ii[>hic range of this genus and of its close ally ^ti/nipetriim (both are 
primarily bcireal. throughout both hemispheres) and the geographic de- 
vi'liipiiHiil (if those lijibitat conditions which they prefer. 



ECOLOGY OP ISLE ROyALE. 265 

Oeographic Range. Xova Rcotia; Ontario; Maine; New Hampshire; 
Maee.; Quebec; Vancouver Island; Kalso, British Columbia; Wanhing- 
ton. 

Lucorhinia inta^ia Uagen. Symphs onlv taken. Ct. Needham report. 

Oeographic Range. Nova Scotia; Maine; New Hampshire; Maaaa- 
chusetts; New York; New Jersey; Penn, ; Ohio; Michigan; Ontario; 
Indiana; Illinois; Wisconsin; Iowa; Bouth Dakota; Nevada; Wafhin;!:- 
ton. 

Sympctrum ruhictindithtm obtrtt»um (Hag.). In the open area about 
the camp on Si^owit Bay (V, 3) this species was \'ery abundant. Eight 
-males and eight females were collected on August 11 and 12. A male 
was also taken near the head of Siskowit Bay (VIII, '04) on August 
13 ; and 3 mates and 1 female were taken in the sedge zone of a tamarack 
swamp (V, 5) on August 8. l^e nrnnber of specimens taken is not a 
fair index of the abundance of this species as an effort was made only 
to secure representative forms. Tlie open areas where the heath and 
jiinipercladon:a plant MKriety were the representative types of vegeta- 
tion, seemed to afford feeding grounds for this species and they were 
very abundant in such places. It is in just such situations that the 
small forms of insect life are most abundantly seen on the wing. 

Geographic Range. Nova Scotia; Ontario; Maine; New Hampshire; 
Mass.; Penn.; New Jersey; North Carolina; New York; Ohio; Indiana; 
Illinois; Wisconsin; Colorado; British Columbia; Washington; Coruona, 
Ann Arbor, Porcupine Mts., Tsle Royale. Mich. 

lAbelliila quadrimaculata L. A single male specimen represents this 
species. It was collected about the Lighthouse clearing (I, 7) on Julv 
25. (153). 

Oeographic Range. Newfoundland; Nova Scotia; Ontario; Maine; 
New Hampshire; Mass.; New York; Quebec; New Jersey; Penn.; Ohio; 
Indiana; Illinois; Wisconsin; Wyoming; Montana; Idaho; Utah; Wash- 
ington; British Columbia; Alaska; Northern and Central Asia; Northern 
Europe; Asia Minor; Corunna, Isle Boyale, Mich. 

Oeographic Notes. Attention is called to the geographic range of the 
following species : 

1. Opkiogomphua coluhrinua. Hudson Bay; Quebec; Maine and New 
Hampshire. 

2 Aegchna sttcherMis. Sitka, Alaska; Saskatchewan; Michigan; 
Newfoundland. 

3. Somat. e. minor. Maine; Quebec; New Hampshire; Wyoming. 

4. CorduHa sckurtle/fi i=^€n€a L.) Newfoundland; Nova Scotia; 
New Hampshire; Mackenzie; British Columbia and Alaska; Northern 
Asia; Europe; Algeria. 

r». Tetra. spinigera. Maine; Geor>[ia; Mich.; Vancouver Island. 

6. L€titx>r. hudionica. Newfoundland; Nova Scotia; west to Winni- 
peg, the Mackenzie basin and British Columbia. 

7. Libel, qttadrimanilata. Newfoundland and Alaska; New Jer- 
sey; northwestward to Wyoming; Washington; British Columbia; 
Northern Europe; Asia. 

Prom the above it is seen that seven of the Isle Royale draftonflies are 
decidedly representative of the region from Labrador to Alaska (wjd> 



2M MICHIGAN SURVEY. 1908. 

more especially of the eastern part of this area), largely aorth of the 
17. S, boundar.Y. There Ib also a marked tranacontinental tendency. As 
these forms do not now occur abundantly even in the mountain regions 
of the went, it if likely that many have spread northweat in post- 
Glacial times vith the Xortheastem Biota, rather than from the reipons 
south of the western glaciated area with its relatively arid climate. 

Of these seven species the following four: Cordvlin aenea (C. schurt- 
leffi), Leucorhitm- hudaonica and Libellvta quadrimaculata, are Asiatic 
(Northern) and European — thus circumpolar. Ab to the geographic 
origin of these forms very little can be said, as the taxonomic relations of 
the Odonata, from a geographic and ecologic standpoint, has never been 
attempted. Attention, however, should be called to the fact that so far 
as known, these are all forms that frequent quiet waters. 

There is an interesting correlation between the geographic range of 
the genera Lmi^orMnia and Stfinpetnim and the geographic development 
of the habitat conditions which they frequent. Both are circumpolar 
in the subarctic r^ion. This same area (especially In America) also 
furnishes the greatest almost continuoug tract of lake, pond and swamp 
conditions found upon the earth. In North America at least, the haae 
leveling of the region. 'its imperfect di-ninage due to glaciation, and its 
cool climate are the important or dominant factors in the production 
of this extensive area of favorable habitats for these genera. It is very 
probable that many animals, dependent upon such conditions, will show 
a similar correlation. 

The powerful flight of the larger species suggests that the present 
distribution of the above listed circumpolar species may have taken 
place under conditions similar to those which exist today. Thus 
the habitat preferences and the present geographic distributioD of the 
species all suggest a faiinal interchange via Alaska and Siberia. Such 
a change might have occurred during Glacial, inter-Glacial or post-Gla- 
rial times, but at present we have no criteria or evidence by which to 
determine such relations. 

The migratory habits of certain species of dragonflies also has a 
direct bearing upon the extensive range of certain species. One Isle 
Royale species, IJbelluIa quadrimaailata, has long been known to mi- 
grate (cf. Dragon Flies and Mosquitoes, 1890, p. 161). 

REFERENCES. 

Calvert, P. P. 

1893. Catalogue of the Odonata-Dragonflies of the Vicinity of Phil- 
adelphia, with an Introduction to the Study of this Group 
of Insects. Trans. Am. Ent. Soc, 20, pp. 152a-272. 
1905. The Fauna of Xew England. No. 6. List of Odonata. Occ. 
Papers Bost. Soc. Nat. His, VII, pp. 1-43. 
Cnrrie, B. P. 
1901. The Odonata (Alaska Harriman Expedition). Pro. Wash. 

Acad. Sci.. 3, pp. 217-223. 
1905, Dragonflies from the Kootenay District of British Columbia. 
Proc. Ent. Soc. Wash., 7, pp. 16-20, 



,dbyGoo<^Ie 



ECOLOGY OF ISLE ROTALE. 267 

Hagen, H. A. 

1875. SynopsiB of the Odonata of America. Pro. Boat 8oc. Nat. 
Hist., 18, pp. 20-96. 
Harvey, P. L. 

1898. ContributiotiB to tlie Odonata of Maine. Ent. News., Ill, 9, 

pp. 59-88. 
Osburn, B. C. 

1906. The Odonata of Brttisb Columbia. Ent. News, 16, pp. 184-196. 
Kellicott, D. S. 

1894. List of the Dragonflies of Coruoaa, Michigan. Can. Ent., 26, 
pp. 345-347. 

1899. The Odonata of Ohio. Spec. Paper Ohio Acad. Sd., No. 2, 

pp. 1-114. 
Needham, J. G. 

1901. Aquatic Insects in the Adirondacka. Bxill. 47, N. T. State 

Museum, pp. 429-540. 
1903. Life Histories of Odonata, sub-order Zygoptera. Bull. 68, N. 
Y. State Museum, pp. 218-276. 
Needham, J. G. and Hart, C. A. 

1901. The Dragonflies (Odonata) of Illinois. Part I, Bull. III. 
State Lab. Nat. His., 6, pp. 1-94. 
Walker, E. M. 

1906. A First List of Ontario Odonata. Can. Ent., 38, pp. 106-110, 

149-154. 

1907. A New Somatochlora, with a Note on the Species known from 

Ontario. Can. Ent,. 39, pp. 69-74. 

1908. A Key to the North American Species of Aeechna found North 

of Jlexito. Can. Ent., 40, pp. 377-391. 
"Williamson, E. B.' 

1900. The Dragonflies of Indiana. 24th Ann. Rep. Dept of Geol. 

and Nat. Resources of Indiana, pp. 229-333. 

1906. Dragonflies (Odonata) Collected by Dr. D. A. Atkinson in 

Newfoundland, with Notes on some Species of Somatochlora. 
Entom. News, 17, pp. 133-139. 

1907. A Collecting Ti-ip North of Sault Ste. Marie, Ontario. Ohio 

Naturalist, 7, pp. 129-148. 
ig07a. List of Dragonflies of Canada. Ohio Naturalist, 7, pp. 148- 
150. 

LBPIDOPTBKA, 



Cameadeg Itostonicnsis Qrt. 
Bemaris definia Bdv. 
Hetnaria tlnifsbe Fabr. 
Papitio glaitcua turnua L. 
Colias philodice Godt. 
Argynnia cyprig Edw, 
Argynnia atlantia EWw. 
Brenthig fm;/rina Cramer. 

Brenthis heUona Fabr. ^-- i 

. Phyciodea tharos Dm. oigtizedbyLjOOglC 



S68 MICHIGAN BURTBT, 1908. 

11. Eufffynia j-album Bd, — Lee. 

12. Euvanessa antiopa L. 

13. Aglaig milberti Godt, 

14. Tatwssa atalanta L. 

15. Vanessa hiintcri Fabr. 

16. Vanessa cardiii L. 

17. BaSilarchia arthemia Dm. 
J8. Anosic plcxippus L. 

19. Epidemia dorciix Kby. 

20. Epidemia cpixanthe Bd, — Lee. 

21. Cu-pido saepiolus Bd< 

22. Amblyscirtes vialis Edw. 

Noctuidae. 

Cameades iostoniensis Qrt. A moth of this species was taken aboat 
camp on BiBkowit Bay (V. -t) on August 9 (G. 222). 

Qeographic Range, "Middle, Bastem and Xortliern States", Massa- 
chusetts; New York; Michigan; Canada. 

Sphingidae. 

Bemarit definis Bdv. One specimen taken in the clearing about the 
camp {I, 7) on July 8 (G. 36). 

Oeographic Range. "Canada, Hudson Bay Territory, Maine to Georgia, 
westward to Mifsonri. Iowa" (Smithi. Michigan. 

Hemaris tkyshe Fabr. One specimen in the open abont the camp on 
Siakowit Bay (V, 3) on August 3 (G. 195). Food plant Viburnum. 

Geographic Range. "Ijabrador, Canada ; southward to Florida ; west- 
ward to the Mississippi" (Smith); Michigan. 

Papilionidae. 

Papilio glaiictis tumtis L. Tumns ButterJiy. Nine specimens were 
taken along the beaches 'I, 1) on July 9, one in tlie clearing about 
the Light-house (I, 7) on July 8. and another on July 14, in a rock 
clearing near the head of Bock Harbor (near III, 3). The Isle Royale 
specimens of this species, when compared with specimens from Ann 
Arhor, are much dwarfed ; the right fore wing of three specimens meas- 
nring 41, 42 and 47 mm. respectively. Most of the specimens secured 
were old worn males. This species is known to become dwarfed both in 
the far north (Alaska) and upon mountains (White Mountains, X. H.). 
A number of animals show this dwarfing tendency to the northward 
when they are of southern origin. Scudder, '99, p. 158. 

Oeographic Range. Newfoundland into Florida; U. 8. generally; 
California into Alaska except in southern British Columbia. This is a 
species of southern origin; its relatives are Bouth American. 

Pieridac. 

CoUan philodirr Godt. Clouded Sulphur Butterfly, This species was 
not taken in 1905 but is recorded by Ruthven ('06, p. 103) from the south 
end of the island. 



ECOLOGY OP ISLE ROTALE. 266 

Qeographic Range. Anticosti, Quebec; Ontario; Xewfoundland; 
Maine; New Hampshire; Mass.; New York; Penn.; West Virginia; Kew 
Jersey; Florida; Ohio; Illinois; Indiana; Kansas; Nebraska; Texas; 
Dakota; Iowa; Colorado. Seudder, '99, p. 24. 

yympMlidae. 

Argynnis ci/prix Edw. One specimen was taken on the burned over 
ridges (II, 3) on July 25 {G. 1-17) ; another August 14 on Solidago, in 
an open area near the beach (V, 2), (A. 115). Determined by Dr. J. 
Fletcher and Dr. Wm. Barnes. 

Qeographic Range. A western Rocky Mountain species. 

ArgynMJs atlantis Edw. Atlantis Butterfly. One specimen was token 
on the rock ridge near the head of Rock Harbor (III) on July 21 (0. 
132) ; two in the clearing at Neutson's resort (V, 5) on July 21 (G. 
121) ; one from the burned over rock ridges on the McCargoe trail (III, 
3) on July 25 (G. 146, 147) ; and five from the clearing about the camp on 
Siskowit Bay (V. 3) on August 4, 9, 10 and 16. 

Seudder eays : "The favorite resorts of this butterfly are grassy fields 
skirting the mountain streams, and it differs slightly from other species 
of the genus, preferring sunny woodland nooks to open country." Seud- 
der, '99, p. 213. 

Qeographic Range. Labrador; Newfoundland; Nova Scotia; Rupert 
House, Quebec; Ontario; White Mts., N. H.; Catskill and Adirondack 
Mts.; N. Y. ; Indiana ; Michigan ; Iowa ; Wisconsin ; Lake Winnipeg ; 
Colorado; Mackenzie and Hudson Bay. 

BrenthiB myrina Cram. Myrina Butterfly. One specimen was taken 
at Sumner Lake (III, 5) on July 18 (G. 120); a second one upon a 
Solidago flower near a small stream flowing from the tamarack swamps 
(V, B) on August 8 (G. 97). Seudder remarks that this species "fre- 
quents low, moist meadows and roadsides in their vicinity," Seudder, 
'99, p. 317. 

Geographic Range. Nova Scotia and New England ; south on the 
Mountains to Maryland, Virginia and North Carolina; Indiana; north- 
ern Illinois; Wisconsin; Iowa; Colorado; Utah; Montana; Alberta; 
British Columbia; Sitka, Alaska; Saskatchewan; Nipigon, Ontario; 
Hudson Bay. Has a near relative in Asia. B. amphisaphe. 

Brcnthis beUona Fabr. Bellona Butterfly. One specimen, no data. 
Seudder, '99, p. 311. 

Geographic Range. Quebec; New England; Ontario; Penn.; New 
Jersey; Virginia; northern Ohio; Indiana; Alberta; Colorado; Great 
HIave Lake; Lake yWinnipeg and Hudson Bay. ' 

Phyciodea tharoa Dru. Tharos Butterfly. One specimen was taken 
on the beach (I, 1) on July 10 (G. 209). Seudder, '99, p. 121. 

Qeographic Range. Southern Labrador into Florida ; west to 
Texas, Mexico and the Sierra Nevada; British Columbia; Alberta; Sas- 
katchewan; Mackenzie River and Hudson Bay. 

Eiigonia J-album Bd. — J^ec. White J. Butterfly. Two specimens 
were taken August 23 and 24 in the clearing at Washington Harbor 
(I, '04) and at the Siskowit Camp (V. 3) on August 16. Seudder, '9!), 
p. 7. This butterfly, according to Seudder, frequents "high open wcod- 



270 MICHIGAN SURVEY, 1908. 

land, and on hilly roadsides in the vicinity of woods," Perhaps mi- 
grates. The butterfly hibernates, cf. Scudder, '97, pp. 139-144. 

Geographic Range. No. Labrador; >'ova Scotia; Ontario; mountains 
of Penn. ; Indiana; Wisconsin; British Columbia; Alaslia. Cloaelj 
related to the European E. van-album. Probably of Asiatic origin. 

Euvanesga antiopa L. Antiopa Butterfly. Not secured in 1905 but 
recorded by Buthven ('OG, p. 103) fnm the south end of the island. This 
is a -wide ranging species from Gautamala and Mexico northward over 
most of the United States and southern Canada; Alberta; British Co- 
lumbia; Alaska; northern Asia and Europe. Scudder, '99, p. 1. This 
species is probably of Asiatic origin. The butterfly hibernates. 

Aglaia milberti Godt. Milbert's Butterfly. One specimen was takeD 
in the clearing at the Light-house on Rock Harbor (I, 7), (G. 36). 
Scudder, '99, p. 330. Butterfly hibernates. 

Geographic Range. Labrador; Newfoundland; New Brunswick; On- 
tario ; Nova Scotia ; New Hampshire ; New York ; northern Ohio ; In- 
diana; Montana; Colorado; Arizona and New Mexico, on the moun- 
tains; Alberta; British Columbia south to central California; Great 
Slave Lake; Mackenzie; liake Athabasca; Hudson Bay. A distinctly 
northern and mountain species. 

Vanessa atalanta L, Bed Admiral. Two specimens were taken in 
the light-house clearing (I, 7), (G, 15), This butterfly hibernates. 
Scudder, '99, p. 79. 

fJeograpkic Range. Southern Labrador; Newfoundland; Hudson 
Bay ; Alberta; British Columbia ; of general distribution over 
United States and extending southward on the mountains into Guata- 
mala; Europe; Northern Asia and Africa. The extensive southward 
distribution on the mountains is worthy of note. This species probably 
originated in Asia. 

Vanessa hunteri Fabr, Hunter's Butterfly. A much worn specimen 
was taken July 19, on the beach (I, 1). (G. 29), and on July 21 in the 
<'learing at Neutson's resort {IV, 5), (G. 121). Scudder, '99, p. 114. 
Butterfly hibernates. Larva feeds on the Pearly Everlasting, Anophali» 
ninrgaritarca B. & IL (Dr. J. Fletcher), but the plant was bot found 
upon Isle Royale. 

Geographic Range. Nova Scotia; Quebec; Ontario; Minn.; British 
Columbia ; United States generally ; Mexico ; Central America, and along 
the Andes perhaps even to Patagonia; Antilles; Canary Isles. This 
species, like the preceding, has an extensive southern distribution along 
the mountains. Of North American origin. 

Vanessa cardui L. Thistle Butterfly, Four specimens were taken 
July 21, in the clearing about Neutson's resort (IV, 5), (G. 121) ; two 
more upon the beach (I, I) on July 10; a wing was found among the 
drift on the beach (I, 1) ; several from the clearing at the Light-house 
(I, 7) on July 7, 10 and 22 (G. 26. 45, 104, 133) ; one in the clearing 
about the camp on Siskowit Bay (V, 3) on August 7 {G. 212) and 
in the cassandra and sedge zone about a pond on Siskowit Bay (V, 
11) on Augusi: 16. Scudder, '99, p. 106, This butterfly hibernates. 
This species, with Argynnis atlantis and Basilachia arthemis, were the 
most abundant butterflies upon the island. 

Geograpkic Range. — This butterflv has the most extensive, range of 

iX-oogFc 



EKX)LOGY OF ISLE HOTALE. 271 

any known species. "With the exception of the Arctic regions and Sonth 
America, it is distributed over the entire extent of every continent." 
(Scnddpr). This species is very abandant in Sonthern Europe; con- 
tinually invades northern Europe but cannot establish itself. It Bwarms 
In immense numbers both in Europe and in America (Calif.). Fletcher, 
'(12, p. C6; Famham, '!)5. p. 150; Scndder, '76. This species is probably 
of North American origin. Its inability to withstand, even in the adult 
stage, the winters of northern Europe and northern North America, and 
it» powers of flight sugfr^Bte that this species, which is probably of 
southeaBtem North American origin, reached the old world not only 
by way of the north but also across the Atlantic Ocean. Specimens 
have been taken at sea 200 miles from the Cape Verde Islands in the 
Atlantic. 

Banilarchia arthemis Dm. Arthemis Butterfly. This butterfly was 
very abundant along the beaches, where they were frequently taken on 
Conglomerate Bay (near I, 5) and where two specimens were taken 
on July 10; five othCr specimens were taken along the beaches on Julv 
19 (G. 29) and one on July 11 and 17 respectively (G. 47, 107) ; also 
one specimen in the Light-house clearing (I, 7) on July 8. 10, 24. 26. 
and i^o on July 17. A single specimen was taken along the path at 
the outlet of Sisltowit Lake {V, 9) on Angust 7 (G. 215). This was a 
burned over area and was relatively open and especially so on the rock 
exposures. Three specimens were taken in the clearing about the camp 
on Biskowit Bay (V, 3) on August 9 (G. 222). It was also abundant 
in rock clearings near the head of Rock Harbor (near III, 3) on July 
14 fG. 97). 

This purple black butterfly with its oblique white band is a con- 
spicuous fovm along the beach, on the rock openings, and in the clear- 
iogs and burnings. Scudder, '99. p. 225. 

OcQifraphio Range. Newfoundland; Nova Scotia: Quebec; Ontario; 
northern New England; New Hampshire; northern and western Mass.; 
Catskill and Adirondack Mts., N. Y. ; mountains of Penn. ; southern 
Mich. ; southern Wisconsin ; northern Indiana ; Minnesota ; Alberta ; 
British Columbia; Fort Simpson, Mackenzie. Of North American origin, 
cf. Field, '04, p. 1. 

Anosia pl&tippug L. Milkweed Butterfly. Two dead specimens and 
one yet alive were found upon the beach at the head of a small cove 
south of the Light-house (I, 1) on July 6 (G. 19). On the following 
day about a half dozen dead specimens were also found under 
simitar conditions. This drift must have been cast up several days 
previously, as was indicated by its stage of decay. A single bright 
colored fresh looking specimen was taken near the head of Siskowit 
Bay (VIII, '04) on August 13, and is in striking contrast to the faded 
specimens preserved from the shore drift. Scudder, '99, p. 95. 

The food plant of the caterpillar is milkweed, one species of which, 
Asclepias incarnata L., was found along Washington Kiver (II, '04), 

This species is not, in all probability, a permanent resident of the 
island, as it cannot endure the winters of easteim Canada. Scudder 
('93, p. 52) has expressed the opinion that this species cannot survive 
the winter north of the Gulf States and that those individuals found " 
further north reach there each rear as migrants from the south, or the 



i72 MICHIGAN SURVEY, 1908. 

immediate descendante of such migrants. It is -well known that these 
butterflieB congregate in vast flocks in the fall and migrate, some think 
to the 80Uth (like birds), others that tbey wander about aimlessl; antil 
killed by the approaching winter (Tntt, '02, p. 127). This wandering 
tendency, however, would tend to scatter them as they died off slowly 
by exhaustion. While it seems incredible to think of a soiitherly 
migratory instinct, yet the meteorological conditions developing in the 
far north might give a southerly direction to the wandering move- 
ments. 

The occurrence of specimens in the shore drift is of interest in 
connection with the wandering habit of tins butterfly. Specimens have 
been picked up on the beaches of Lake Michigan (Needham, '00, p. 6) ; 
Lake Erie (Moffat, '01. p. 48) ; and Lake Ontario (Bowles, Can. Ent. 
Vol. 12, p. 134; and they have been observed flying over Lake Erie. 
Such facts as above cited suggest that this member of the Isle Royale 
fauna is restocked each year by migrants, which are probably mope 
likely to come from the southern rather than the northern shore of 
Lake Superior. If Scudder's opinion is correct, an annual extension 
of range from the Gulf States to Isle Koyale — over 1,200 miles — certain- 
ly shows remarkable powers of dispersal. 

Geographic Range. — This species has a very extensive range in this 
hemisphere from northern Patagonia in South America, northward 
through the tropics, West Indies, over most of the United States and 
southern Canada to British Columbia, Hudson Bay and Lake .\thabaftka. 
Through man's influence this species has become almost world-wide in 
its range. It has been recorded from the south Pacific 500 miles from 
land (Tutt, '01, p. 40). Originally it was of American origin. 

Lycaenidae. 

Epid^mia dwcas Eby. A single specimen of this was taken on August 
16 in the Cassandra and sedge sone of an open bog (V, 11), (A. 136). 
Determined by Dr. James Fletcher. 

Geographic Range. — Michigan (Isle Royale) ; Nipigon, Ontario; Mani- 
toba ; Saskatchewan ; Athabasca ; .Alaska. 

Epidemia epixanthe Bd. — Lee. Exipanthe Butterfly. Pour specimens 
were taken on the sedge zone on the north shore of Sumuer Lake (1X1* 
5) on July 29. 

This is a swamp species about which Fiske ('01, p. 50) writes: "It 
confines itself closely in its jouraeyings to the swamp or bog in which 
its early stages are passed, and rarely indeed ventures upon higher 
ground. It loves best of ail an open, mossy morass, such as are found 
scattered throughout New England, usually surrounding some small 
pond caught in a hollow between the hills, and formed by the moss and 
«ubaquatic plants which, constantly encroaching upon the water, are 
bonnd in time to cover it over." 

Geographic Range. Newfoundland; Quebec; Ontario; Maine; New 
Hampshire; northern Indiana; Iowa; Kansas and Nebraska. 

Ciipido sacpiolus Bd, Greenish Blue Butterfly. In all seven speci- 
mens of this interesting western species were taken ; one about the 
camp at the Light-house (I, 7) on July 7, (G. 26); a second in the 
clearing on Benson Brook (II, 1) on July 25 (G. 148) and five speci- 



ECOLOGY OP ISI^E ROYAI^. ' 278 

mens in the clearing about Neutson's R«8ort (IV, 5) on July 21 (G. 
121). 

Geographic Ran^e. — Michij;aD (Isle Royale only) ; Great Slave Lake; 
Mackenzie Baein; British Columbia; Montana; Colorado; NevBda; Cali- 
fornia. Cf. Elrod, '06, p. 136;Care.r, '06, p. 451. Isle Royale ia alec 
the most eastern record for this species. 

Seaperidae. 

Ambhfscirtea viali$ Ed-n-. Vialis Skipper. Three specimens of this 
skipper were taken on Jnly 11 (G. 49). 23 (G. 133), and 28 (G. 179) 
in the clearing about the Light-house (I, 7). 

Geographic Range. Quebec ; Maine ; New Hampshire to Florida and 
westward to Texa*; Nevada; Alberta; Manitoba (Fletcher), and 
British Colnmbia. 

Geographic 'Notes, After the preceeding geographic records had been 
secured, the following notes on the distribution of butterflies in Can- 
ada were received from Dr. James Fletcher, of Ottawa, Canada. His 
letter contains so many interesting features that I have thought it de- 
sirable, with Dr. Fletcher's consent, to publish it in fnll, supplementary 
to the data already given, rather than to scatter the records. 

"In just running through your letter I see that I can answer it with- 
out turning tip any records. The geographical range in Canada of the 
following butterflies is as follows: Papilio glaucua tumua — from the 
Atlantic to the prairie region, common; across the prairies into British 
Columbia, rather scarce, and not to my knowledge crossing the interior 
elevated plateau which is a semi-arid region. North of this Jn the moun- 
tuins it reaches right to the Pacific Coast. South of the north part of 
Vancouver Island its place is taken by Pap. eurymedon and P. rutulus 
arizonensig as named by Mr. W. H. Edwards. 

"CoUas philodice — very abundant from the Atlantic to the Lake 
Superior region, where its place is taken by C. eurytheme, of which one 
form, the variety eriphgle resembles philodice very closely and although 
it is claimed that it is a form of eun/tkcme it resembles philodice so 
closely that it cannot always be separated unless the locality is known. 

"^rgynnia atlantia — this occurs in what we consider the typical form 
from the Atlantic coast to the Great I^kce. West of that the black 
markings are rather lighter and the color is brighter. In the Rocky 
Mountains I believe what we have been calling eUcta is merely a form 
of atlantia. That at any rate extends to the main chain of the 
Rockies, but I have never seen it further west than the Arrow Lakes. 

"BrenthAa myrina and lellxma — from Atlantic Coast to the interior 
plateau of British Columbia. 

"P. tharoa — from the Atlantic Coast to British Columbia, running 
north to the coast, probably with the main chain of the Rockies. In 
Vancouver Island and the Fraser River Valley its place is taken by P. 
pra-tensia and the same areas are inhabited by Brenthia cpithore in 
place of tdUma. Pratensia however extends west into Manitoba. 

"Ettffonia j-album, E. antiopa, A. milberti, V. atalanta and V. cardui 
occur over the whole of our country from Atlantic to Pacific and from 
the southern border to the arctic regions. 



274 MICHIGAN. SURVEY. 1908. 

"V. h'unteri also occurs right to the coast but is very much rarer 
west of the Great Lakes thau the other species meDtioned. I have it 
from Xora Scotia and also took it on Vancouver Island. 

"BasilarcfUa arthemis extends from the Atlantic coast to the Kootenai 
Lakes, when its place is taken hy B. lorquimi. Anoaia ptexippua a mi- 
grant and may turn up at any place where Aaclcpias grows, but is 
much rarei- in British Columbia. 

"Epidemia epixantJie. — ^This is the only species I have some doubts 
about. Ttiere is no doubt that some of the records of epixanthe should 
be of the rare and little understood species dorcaa which occurs in the 
Lake Superior region and into Manitoba. It is easily distinguished 
from epi-vanthe by its slightly larger size and the brilliant orange wash 
on the under surface. Bptxanthe I have only actually taken myself in 
Ontario. Dorcaa I have from Nepigon on Lake Superior the Bruce 
peninsula and from Manitoba. West of that the form, for it is hardly 
a variety, florus which is really only a dimorphic form of helloidea 
occurs, and has I think, sometimes been recorded as epixanthe. The 
reference of fiorvs to dorctis instead of hclloides as a variety, which 
was done by Dr. Dyar, has in my opinion no reason in it at all, 

"Amilyscirtea vialis.— This is nowhere very common but extends from 
Atlantic to the Pacific coast. I have specimens from Halifax, Nova 
Scotia and have taken it in Vancouver Island. It is more abundant 
perhaps in the Lake Superior region than any other where I have col- 
lected." 

The butterfl.y fauna of Isle Royale may well be compared to that of 
the White Mountains of New Hampshire on account of the large number 
of species common to both localities (cf. Scudder, '97, pp. 71-87). Much 
the same resemblance holds for northern New England in general. So 
far as United States is conce;-ned Isle Royale is the western outlier 
of the distinctly northeastern or Canadian biotic type. Perhaps the 
Black Hills will show similar affinities, but farther west a marked 
Rocky Mountain influence becomes apparent. It is of interest to note 
that six of the eighteen species hibernate as butterflies and another 
spreads eacli season into the region. This number includes the species 
which are of the most northern nmge and one (F. cardtii) which is 
cosmopolitan. Four of the species, B. j-album, P. cardui, B. arthemia 
and A. plexippua are known to flock, or migrate. There can be but 
little doubt but that these characteristics are important factors in an 
understanding of their geographic range, and are probably adaptations 
which permit these species to maintain themselves in the region. Such 
adaptations may have originated in response to the environment or the 
possession of them have allowed the species to enter the region already 
adapted to it. 

When the above listed species are grouped geographically they fall 
into the following classes : 

1. Of general geographic range from Labrador, Newfoundland, north- 
ern New England, southward on the Adirondacks, Oatskills and Ap- 
palachians, westward through northern Ohio, Indiana, Wisconsin to 
Alberta, and southward on the Rocky Mountains, British Colombia to 
Alaska. 



ECOLOQY OF ISLE ROTAI^. 27S 

1. Argynnis atlanttt, 

2. Brenthia myrina. 

3. Brentkig bellona. 

4. Eugonia j-album. 

5. Vaneaga milberti. 

6. Baailarchia arthemia. 

These forms ma; well \m: called members of the Canadian biota for they 
-only exteud a short distance southward on the lowlands of the United 
States, but reach much farther Bouth at higher altitudes. 

2. Much the same uorthem limit as group 1, but reaching much 
fiarther south of the above soathern lowland limit. 

1. Papilio glaucua tumua. 

2. Colias phUodice. 

3. Phpciodea tharoa. 

4. Euvaneasa antiopa (Asiatic). 
&. Taneasa atalanta (Asiatic). 

6. Vaneata hwnteri. 

7. Vanessa cardui (cosmopolitan). 

8. Anoaia plexipjma (Nearly cosmopolitan). 

9. Epidemia epixanthe. 
10. AmhVyacirtea vtalia. 

3. Species of distinctly Rocky Mountain or Rocky Mountain and 
Pacific coast distribution, and reaching their eaBtern limit in the Lake 
;Snperior region. 

1. Epidemia dorcaa, 

2. Cupido saepiolua. 

3. Argynnia cypria. 

4. Of very extenpive getigraphic range, Asiatic or Cosmopolitan. 

1. Pyrameia cardui. 

2. Anoaia plexippus. 

3. Vatteasa atalanta. 

4. Euvaneasa antiopa, 
I¥obable geographic origin: 

1. Papilio glaucus tumua, S. American. 

2. Colias philodice, American. 

3. Argynnia atlanta, Asiatic. 

4. Brenthia myrina, Asiatic. 

5. Brenthia brllona, Asiatic. 

6. Phyciodea tharos, American, 

7. Eugonia j-album, Asiatic. 

8. Euvaneasa antiopa, Asiatic. 

9. Vanessa milberti, American. 
"lO, Vanesaa atalanta, Asiatic. 

11. Vanessa huntert, American. 

12. Vaneaaa cardui, Amercian. 

13. Basilarchia arthemia, American, 

14. Anoaia plexippus, So. American. 

15. Cupido saepiolua. West No. Amer. 

16. Epidimiii cpiratithr. Eastern U, S. 
:17. Epidemia dorcas, W. No, Amer, 

18. Amblyscirtea vialis, No. Amer. -. 

D,gt,zedby(jOO<^[e 



276 MICHIQAN SURVEY. 1908. 

In discBBsing the geographic origin of the batterfiies oommon to 
the old and new worlds, Scndder Beldom attempts more than a hemis- 
pherical location. In discussing the origin of Americnn faunae that 
are both boreal and Asiatic, it is well to recall that geologically speak- 
ing the American boreal and arctic are larffcli/ of rerciit origin in the 
northern regiOTit. It is therefore not uulikclv that manv of these forms 
which it has been customary to consider boreal are in reality not so, 
but from high altitudes — frdm the North American Cordilleras or from 
the Himalayas, where high altitude and low teaiperatare existed long 
before the Ice Age. With the development of an Ice Age, there was 
a great increase of this low temperature, lowland habitat and when 
once the glacial climate declined a vast area was open for invasion — 
an area of such great extent that we have become thoroughly accustomed 
to think the fauna has originated there. It has thus become customary 
to speak of them as of northern origin, in spite of the fact that we know 
that they are almost entirely post-Olacial migrants from the south. 

EEFERBNCES. 

Carey, M. 

1906. On the Diurnal Lepidoptera of the Athabaska and Mackenzie 
Begion, British America. Pro. U. B. Kat. Mus., 31, pp. 425- 
457. ■ 
Dyar. H. G. 

1903. A List of North American Lepidoptera. Bull. 52, U. S. Nat. 

Mas. 

1904. The Lepidoptera of the Kootenai District of British Columbia. 

Pro. U. B. Nat. Mus., 27, pp. 779-938. 
Elrod, M. J. 
1906. The Butterflies of Montana. Univ. of Montana, Bull. 30, Biol. 
Series No. 10. 
Engel, H. 

1908. A Preliminary List of the Lepidoptera of Western Pennsylvania 
collected in the vicinity of Pittsburg. Ann. Carn^e Mna., 
6, pp. 27a36. 
Farnham, G. D. 
1895. (Migration of Pyrameis cardiii in Calif.) Ent. News, 6, p. 
150. 
Field, W. L. W. 

1904. Problems of the Genus Basilarchia. Psyche, 11, pp. 1-6. 
Fiske, W. P. 

1901. An Annotated Catalogue of the Butterflies of Sew Hampshire^ 

Tech. Boll. No. 1, N. H. Agr. Exp. Station. 
Fletcher, J. 

1902. The Painted Lady Butterfly. Thirty-second Ann. Rep. ' Ent. 

Boe. of Ont., 1901, pp. 54-57. 
MoCFat, J. A. 
1901. Anosia archippus, yet Again. Thirty-first Ann. Bep. Ent. Soc. 
Ont., 1900, pp. 44-51. 
Necdham, J. G. 
1900. Insect Drift on the Shore of Lake Michigan. Occ. Mem. Chicaga 
Entomol. Soc., 1, pp. 1-8. , , 



BCOLOGT OF ISLBI BOTAl^. 277 

Bnthven, A. G. 

1906. Spiders aod Insects from the Porcupine Moantains and Isle 
Rojale, Michigan. Ann. Rep. Mich. Oeol. Burr, for 1905, pp. 
100-106. 
Scadder, 8. H. 

1876. A Cosmopplitan Butterflj. Amer. Nat, 10, pp. 392-396; 602- 

611. 
1889. Coamopolitaa Butterflies. Psyche, 5, pp. 190-192. 

1897. Frail Children of the Air. Boston. 
1899. Everyday Butterflies. Boston. 

Skinner, H. 

1898. A Synonymic Catalogue of the North American Bhopalocera. 

Amer. Entomol. 8oc. 
Tutt, J. W. 

1902. The Migration and Dispersal of Insects. London. 



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ECOLOGY OF ISLE HOTALE. 



ANNOTATED LIST OF THE ilOLLrSCA OF ISLE KOYALE, MICH- 
IGAN. 

BY BftTANT WALKER, 

1. Introduction. 

This list coataJDS all the species of molluBcs collected on Isle Boyale, 
both in ld04 and ld05, and so represents the complete fanna bo far as 
known. For the details as to the localities represented bv the col- 
lection made in 1904, reference should be made to the Report of that 
expedition (pp. 96-99). 

The collection made in 1905 was much larger than that of the preced- 
ing year, both in species and IndiTidnals. While eleven of the species 
■ collected in 1904 were not found in 1905, no less than forty-two ad- 
ditional species were obtained, making the total nnmber of species now 
known from the island seventy-two. The species added in 1905 are in- 
dicated by an * on the list. In compiling the completed list, it has 
been: deemed of aufflcient interest to add both the general range of 
each species and its distribution so far as known in the state of Mich- 
igan. 

2. Faunal Afjinities. 

The mollnscan fanna of Isle Royale becomes of additional interest 
when considered in conection with Dr. W. H. Dall's recent work on the 
''Land and Fresh Water Mollusks of Alaska and Adjoining Regions," 
(Harriman Alaska Expedition, Vol. XIII, 1905), which is a complete 
digest of onr present knowledge of the land and fresh water molluacn 
of North America north of latitnde 49° North, and practically of our 
entire boreal fauna. So far as available, the general range of the species 
given in that work has been followed in this list as being the most "up- 
to-date" attainable. 

Of the seventy-two species recognized from Isle Royale, forty-nine 
are included by Dr. Dall in his list of boreal species. 

The remaining twenty-three species not cited by him are: 

Strohilops Virgo (Pils.) 

Vertigo tridentata Wolf. 

Euconulus chcrsintts poJygyratus Pils. 

Zonitoides exigua (Stimp.) 
ylgriolimax campestris (Say). 
:J>alUfera hemphilU (W. G. Binn.). 

•J^alUfera doraalia (Binn.). 
^Lymnwa pilsbryana Walker. 
^hysa aayii Tap'p. 

,Physa aplectoides Sterki. 

Ancylus sp? 

Amnicola lustrica Say. 

Mmculium 8€curis Prime. ,-. , 

37 D,gt,zedby(jOOg[C 



282 MICHIGAN SUBVBY. 1908. 

PUidium afflne 8terki. 

Pigidium aargenti Sterki. 

Pigidium roperi Sterki. 

Pisidium subrotimdum Bterki 

Piaidium spJendidulum Bteriii. 

Piaidium pauperculum Sterki. 

Pisidium medianum Sterki. 

Piaidium punctatum aimplea; Sterki, 

Of these Lynrnga pilsbryana is, bo far as yet known, peculiar to Isle 
Royale. The remainder have, as a rule, a general distribution all over the 
state. !?ltrohilopa rirgo and Zonitoidea exigva, however, are charaoterie- 
tic boreal species and are apparently rarely found south of the Saginaw- 
Orand Valley. While, of course, it is possible that Isle Royale marks 
the northern limit of the range of these species, in view of the fact 
that nearly all of them are known to rajige through the Upper FeDiasnIa 
and that practically nothing is known of the fauna of the north shore 
of Lake Superior, the probability is that most of them range further 
north and should be included in the boreal fauna of North America. 
Wbile the Isle Royale ftinna is thus to be considered purely boreal 
in its character, it must be understood that it is not in any material 
respect different from that of the Upper Peninsula and of the northern 
part of the Lower. The occurrence of such species as ValUmia costala. 
Vertigo tridentata and Physa aplectoidea must be considered rather as 
evidence that these forms have been overlooked in the region immediate- 
ly south rather than that they are restricted to Isle Royale. And the 
existence of Lymnwa pilsbryana and the peculiar forms of Lymtuta 
atdgnalie, Phmorbia bicarinatua and P. campanulatua, if ultimately foand 
to be confined to the island, should rather be ascribed to long isolation 
under peculiar local conditions than as indicating any essential difference 
from the fauna of the surrounding region. 

Considering the fauna of the Upper Peninsula and Isle Royale as a 
whole, it will be found to include two elements. The first and larger 
one consists of species having a general range through the northern 
United States and southern portion of Canada, Just how far to the 
north most of them range is as yet undetermined. These species, although 
found in the boreal region, cannot be properly said to be distinctly 
boreal species. Accompanying tliewe species of geueral distribution, is 
a smaller element of purely boreal species, which are characteristic of 
the northern region and whose range to the south i.« as a role quite 
restricted. This purely boreal element is represented on Isle Royale 
by the following species : 

1. Aranthinula harjm (Say). 

2. Yitrina limpida Gould. 

3. Vitrea hinneyana (Mse.). 

4. Zonitoidea CTigua (Stimp.). 
a. Pyramidula asteriacna (Mse,). 
B. Lymrwa megaaoma Say. 

7. Lymnaa emarginata Say, 

8. liymnfra pilsbryana Walker. 

So far as the land species of Isle Royale are concerned, there are 
but few of special interest, most of them being of general distribution 



ECOLOQT OF ISL& ROTALE. 283 

in the Upper Feninsola. The occnrrence of Polygyra alboUibriB (which 
was not found in 1904) in abundance causes a feeling of surprise that 
Polygra fratema (monodon auct.) was not found. In Michigan, its 
range is coincident with that of alboldbria and, according to Dall, it is 
fonnd as far north as James Baj, Hudson Bay. 

All the specimens collected of Pyramidula cronkhitei were of the 
anthonyi form as in 1904. 

Among the fluviatile species, however, several forms either pre- 
viously undiscovered or new to the fauna of the state were found and 
some interesting facts in regard to their distribution were developed. 
The division of Lymnaa staffnaUs Into three very distinct forms coin- 
cident with the character of their habitat is very striking and specially 
noteworthy. 

The fact that all the larger species of Lymn(Bid€te from Siskowit 
Lake: Lymtuea stagnali«, Planorbis bioarinatua and PUtnorbi^ cam- 
panulatus, are well marked varieties peculiar to that lake, is very 
significant and points to some specially farorabie envlroamental condi- 
tions, which apparently are not present to affect the facies of these 
species on other parts of the island. What these are, if not already de- 
termined, would be an interesting subject for future investigation. 

The acqnisitton of the additional material, which has enabled the 
specific distinctness of LymjuEa ptlabryana to be determined, is a matter 
of congratulation. 

The occurrence of the beautiful little Phyaa apUctoidea adds a new 
species to the fauna of the state and affords another instance of ap- 
parently anomalous distribution, which so frequently puzzles the 
student. Originally described from Ohio, it has hitherto escaped at- 
tention in southern Michigain, and its discovery on Isle Boyale was 
wholly unexpected. The remarkable form of PJanorbia bicarinatui 
from Siskowit Lake was one of the most interesting novelties discovered 
by the expedition and is a noteworthy addition to the fauna of the 
state. 

"All the Pisidia, except Pisidmm idahoense and Pisidium aargenti, 
are represented by small, and, as it seems, characteristically northern 
forms, slight and generally of light or pale color. Some are not very 
characteristic and apparently little different from each other and 
were rather difBcuIt to work up," (Sterki.) 

The writer again acknowledges his indebtedness to Dr. V. Kterki for 
the identification of the Sphwria and Pisidia and to Dr. H, A. Pilsbry 
for the determination of the sings. Dr. H, A. Gleason, who collected 
most of the specimens, and Sir. Chas. C. Adams have kindly interpolated 
the details as regards the local habitats and distribution of the 
different species. The field numbers by Mr. C. C. Adams are indicated 
bv the letter A; all others are Dr. Gleason's, except a few lots collected 
bV Mr. N. A. Wood. 
' Detroit, April 1, 1909. 

3. Annotated List. 

1." Polygyra alboTabria (Say). 

Range: "Eastern United States, from Georgia and Arkansas to the 
Saskatchewan." (Dall). iX.OOglC 

Michigan : Generally distributed. 



2M MICHIGAN 6URVBT, 190S. 

Distribution on Isle Rovale: Station 1, Sub. 1, I^abe and Bav 
Beaches, Nambers 19, 32, 50 ; I, 5, Jack Pine Eidges, Nob. 19 A, 23. 33. 
81, 187; I, 7, Light-house Clearing, Nos. 34. 42; II, 1, Ransom Clear- 
ing, No. 150; II, 2, Tamarack Swamp, So. 113; II, 3, Kock Ridge 
Clearing. Nos. 51, 145; III, 4, Trail to Sumner Lake, Nob. 88, 93. 13S. 
174; III, 5, Sumner Lake, No. 120; HI, 6, Southwest Coves, Rock 
Harbor, No. 91; IV, 5, Neutson's Resort, No. 121; IV, 8, Trail to 
Greenstone Range, No. 128; V, 2, Heath Zone and Beach, Nos. 101 A. 
107 A; V, 3, Rock Clearing at Camp, No. 233; VIII, '04, Upper End of 
Siskowit Bay, No. 232. 

A "dead" shell was found in driftwood cast upon the beach at the 
head of Tonkin Bay (No. 19) and {No. 32) in a small creek at the 
head of Conglomerate Bay. The animal was dead but the body was 
still within the shell. In both of these numbers the ehells were be- 
yond their normal habitat. 

Live examples (No. 50) were seldom seen but the abundance of the 
dead ones npon the rock ridges and open Gladonm clearings make it 
apparent that they are abundant here. Numbers, 23, 33. 81, 187. 51. 
145, 88, 93, 138, 174, 128, 101 A, 107 A, 232 and 233 were all taken from 
that association. Thence they wander in small numbers to the moister 
places, such as the lighthouse clearing (Nos. 34 and 42), Ransom 
Clearing (No. 150), or even in the tamarack swamps (No. 113). 

Although not found at alt in 1904, this species was collected in 
1905 in considerable numbers, which show a wide variation both in 
size and in the thickness of the shells. The 124 mature specimens col- 
lected varied in height from 14 to 21.5 mm., and in width from 21.75 to 
30.75 mm. The average being 17 by 25.86 m/m. The accompanying dia- 
grams, 61-62, show the variation in height and greater diameter. 

A series of 42 from all parts of the Upper Peninsula vary in height 
from 14 to 20 mm., and in width from 22, 75 to 30 mm., with an average 
of 17.20 by 26.28 mm. While a series of 183 from all parts of the Lower 
Peninsula vary in height from 12 to 24.25 mm., and in width from 18.75 
to 34.25 mm., with an average of 18.10 by 27.11 mm. 

It would appear from these series that the average Isle Royale shell 
is slightly smaller than the average specimen from the Upper Peninsula, 
and considerably smaller than the average Lower Peninsula example. 
The range of variation in the Isle Eoyale series is somewhat greater than 
in the Upper Peninsula series, but much less than in that from the 
Lower Peninsula. 

2. AcaiUhinula harpa (Bay). 

Range: "Northwestern Scandinavia, Northeastern America, British 
America near Hudson Bay, Southeastern Alaska and the easternmost 
margin of Siberia." (Dall.) 

Michigan : Petoskey and Charlevoix in the Lower Peninsula, and 
Ontonagon County and Isle Royale in the Upper. 

Isle Royale: I, 5, Jack Pine Kidges, No. 19 A; II, 1, Ransom Clear- 
ing, No. 150 ; V, 2, Heath Zone and Beach near Siskowit Bay, S(^. 118 
A, 129 A, 130 A; V, 4, Trail through Balsam-Birch Forest, No. 236. 

Number 150 was collected under logs resting on the ground in an 
open clearing near the lake; No. 236 was in leaf mold in the dense 



ECOLOGY OF ISLE EOTALE. 285 

shade of tbe balsam forest; No. IS A vaa taken from under a atone; 
and No. lis A, 129 A, 130 A from under mats of Cladonia. 

3." Strobilops Virgo (Vila,). 

Range: "Canada to Northern Alabama, and west to JXinnesota and 
Kansas," (Pilsbry) , 

Michigan : Upper Peninsula and northern conntiea of tbe Lower. 

Isle Rovale: I, 5, Jack Pine Ridge, No. 81; V, 2, Heath Zone and 
Beach near Siskowit Bav, Noa. 129 A, 130 A; III, '04, Deaor Trail, No. 
142 A. 

In damp aoil (No. 81) under loose rocks at a depth of 2-6 inches; 
Ko. 129 A and No. 130 A from under Cladonia; and No. 142 A from 
leaf mould and rotten logs in the maple forest. 

4." Bifidaria tap-paniana (C. B. Adams). 

Range : "Ontario to Gulf of Mexico, west to Iowa and Kansas, south- 
west to Arizona, bnt not known from the southeastern Atlantic States, 
Virginia to Florida." (Vanatta and Pilsbry). 

Michigan: Oenerallj distributed in Lower Peninsula, Isle Boyale. 

Isle Boyale: II, 1, Benson Brook, No. 150. A single specimen, the 
first record of this species from the Upper Peninsula. 

Under a prostrate log in an open place near the lake. 

5. Vertigo ovata Say. 

Range: "Eastern United States from Maine to Texas and north- 
ward." (Dall). 
Michigan : Generally distributed. 
Isle RoyaJe: Not collected in 1905. See Report 1904, p. 97. 

6. Vertigo gouldU Binn. 

Range: "Northern United States east of the Socky Mountalna and 
northward," ( Dall.) . 

Michigan : Generally distributed. 

Isle Royale: Not collected in 1905. See Report Exped. 1904, p. 
97. 

7." Tertigo tridentata Wolf. 

Range: "Quebec and Maine to Minnesota, aontb to Illinois and Ohio." 
(Pilsbry). 

Michigan : Ann Arbor, Grand Rapids and Isle Royale. 

Isle Boyale: V. 2, Heath Zone and Beach near Siskowit Bay, No. 
130 A. 

A single specimen found under Cladonia. The first record from the 
Upper Peninsula. 

8. Vertigo sp? 

Isle Boyale: I, 2, Natural Bock Clearings, Light-house Peninsula. 
Two unidentifiable fragments, 

9. Virtrina limpida Gld. 

Bange: "Central New York and northward from New Brunswick to 
Alberta and Hudson Bay." (Dall). 

Michigan : Upper Peninsula and northern coanties in the Lower. 

Isle Royale: V, 2, Heath. Zone and Beach near Siskowit Bay, No. 
130 A. A single dead specimen was found under Cladonia. 

10. Vitrea hinneyana (Morse). 

Bange: "Quebec and Maine to Northern Michigan and British Colum- 
bia." (Dall). ,--. , 

X.ooglc 



286 HICHIQAN SURVEY. 1908. 

Slicbigan; Upper Peningnla and northern counties in Lower. 

Isle EoyaJe: I, 3, Balsam-Spruce Forest, No. 140; I, 5, Jack Pioe 
Ridges, Xos. 19 A, 81 ; V, 2, Heath Zone and Beach near Siskowit Bay, 
No. 130 A; III, '04, Deaor Trail, Noe. 141 A, 142 A; V, '04, Eidge back 
of Club House, Noe. 144 A, 147 A. 

In moist soil under loose rocks (Nos. 81 and 19 A) or in the loose 
leaf mold under the balsam forest. No. 140 ; No. 130 A under CladorUa; 
Noe. 141 A and 142 A, from leaf mould or rotten logs in yellow birch 
or maple forest and Nos. 144 A and No. 147 A from under bark, leaves 
and among moss in the forest. 

11. Euconulus fulvua (Dr.). 

Kange: "Holarctic, and widely distributed southward," (Dall). 

Michigan: Generally distributed. 

Isle Royale: I, 5, Jack Pine Ridge, No. 19 A; V, 2, Heath Z<Hie and 
Beach near Siskowit Bay, Nos. 129 A, 130 A ; V. '04. Ridge back of the 
Clnb House No. 147 A. No. 15 A from under a stone; Nos. 129 A and 
130 A from under Cladonia. 

12. Euconulus chcrainut polygyratus (Pila.). 
Range: Northern United States and Canada. 
Michigan : Generally distributed. 

Isle Royale: I, 5, Jack Pine Ridges, No. 19 A; V. 2, Heath Zone 
and Beach near Siskowit Bay, No. 129 A; III, '04, Desor Trail, No. 
142 A ; V, '04, Ridge back of Club House, No. 147 A. 

Pound under a damp stone (No. 19 A) ; under Cladonia (No. 129 A) ; 
in the forest nnder leaf mould, barb, moss or decaying logs (No. 142 A 
and 147 A.). 

13. Zonitoides arborea (Say). 

Range: "North America generally and Japan." (Dall). 

Michigan : Generally distributed. 

Isle Rovale: I, 2, Natural Rock Clearings, No. 65; I, 3, Balsam- 
Spruce Forest, Nos. 140, 141 ; I, 5, Jack Pine Ridges, Nos. 19 A, 81, 102 ; 
II, 1, Ransom Clearins, No. 150; II, 2, Tamarack Swamp, No. 113; V, 
2, Heath Zone and Beach near Siskowit Bay, Nos. 118 A, 129 A, 130 A; 
V, 4, Balsam-Birch Forest, No. 236; III, '04, Desor Trail, Nob, 142 A, 
149 A ; V, '04, Ridge back of Club House, No. 147 A. 

In the thin soil collected under bearberry on the dry rock clearing 
north of the lighthouse (No. 65); under loose rocks in the jack pine 
lidges (Nos. 19 A, 81, 102) ; under log in an open, sunny place (No, 150) ; 
or in leaf mold in the dense shade of the balsam forest; nnder Cladonia 
iNos. IIS A, 129 A, i:iOA) ; and in the dense forest among litter (No«*. 
142 A, 147 A, and 149 A). 

li. Zonitoides exigiia (Stimp.), 

Range: "Quebec and Ontario, New England, New York, Alleghan.T 
Co., Pa. and Michigan." (Pilsbry), 

Michigan: Upper Peninsula and northern counties of the Lower. 

Isle Royale: III, '04, Desor Trail, Nos. 141 A, 149 A; V, '04, Back 
of Club House. No. 144 A. 

Found only in the dense forest among litter. 

15. Zonitoides miUum. (Morse). 

Range: "Eastern United States and Canada, Manitoba." (Dall.) 

Michigan : Generallv distributed. -^-. ■ 

lyCoogle 



ECOLOGT OF ISLE KOTALE. 287 

Isle Royale: V, 2, Heath Zone and Beach near Siskowit Bay, No. 
130 A. A single specimen under Cladonia. 

16. AgrioUmax campeatris (Binn.). 
Bange: "Entire United States." (PiUbry). 
Michigan : Generally distributed. 

Isle Rovale: V, 2, Heath Zone and Beach near Siskowit Bay, No. 
133 A, Found only ander Cladonia. 

17. I'nWfcra hcntphim (W. G. Binn.). 

Bange: Mountains of Georgia, North Carolina, and Eastern Pennayl- 
vania, and Michigan. 

Michigan : Isle Borale, Ontonagon Coantv and Ann Arbor. 

Isle Royale: Not collected in 11)05. See Report Bxped. 1904, p. 96. 

18.* PalUfera- dorsalis (Binn.). 

Range: New England, New York and Michigan. 

Michigan: Isle Royale. Eaton and Marqnette Gonnties. 

Isle Royale: III, '04, Desor Trail, Nos. 142 A, 149 A, 

Taken only in the dense hai'dwood forest among litter. 

19. Pyramiduia aUemata (Say). 

Range: "Eastern North America as far north as Nova Scotia, Lower 
Canada and the international boundary." (Dall). 

Michigan: Generailv distributed. 

Isle Royale: HI, '04, Desor Trail, Nob. 142 A, 143 A, 149 A; V, 'Oi 
Ridge back of Club Honse, No. 144 A; VIII, '04, Upper end of Siskowit 
Bay, No. 232. 

From the Htter of the maple forest (No. 142 A and 144 A and 149 
A). 

20. Pjframidula cronkhitci anthonyi (Pilsbry). 

Range: '*Kan6aiJ. northward to Great Slave Lake and from New Eng- 
land to the Sierra Nevada and south to Arizona." (Da!!). 

Michigan: Generally distributed in the Lower Peninsula. 

Isle Royale: I, 2, Natural Rock Clearing, No. 78; BalBam-Spruce 
Forest, Noa, 140, 141 ; T, 4, Tamarack and Arbor-vitie Swamp, No. 182 ; 
I, 5, Jack Pine Ridge, Nos. 19 A, 81, 102; II. 1. Ransom Clearing, No. 
150 ; Y, 2. Heath Zone and Beach near Siskowit Bay, No. 129 A, 130 A ; 
V, 4, Trail through Balsam-Spruce Forest, No. 236; III, '04, Desor Trail. 
Nos. 141 A, 142 A, 149 A; V, '04, Back of Club House, Nos. 144 A, 14r 
A. 

As stated in the 1904 Report, this form is apparently replaced 
throughout the Upjier Peninsula by the var. catskillcnxis Pils. 

This species shows a wide range of habitat, and may occur under or 
in decaying 1(^« (No. IS, 150); under loose rooks (19 A, 81, 102); in 
leaf moid (140, 141. 236). Also found under Cladonia (129 A and 130 At 
and in the litter of the hardwood forest (141 A, 142 A, 144 A, 147 A, 
and 149 A). 

20a. var. albina (Ckll.), 

Isle RoTale: This form occurred in 1905 at I, 5, Jack Pine Ridge, No. 
]9 A; HI, '04, Desor Trail, No. 141 A; V, '04, Back of Club House, No, 
147 A. 

Found under stones (No. 19 A) and in the hardwood litter (No. 141 
A and 147 A.) ' /^- i 

21. Pyramiduia astenscua (Morse). i.,000QIc 



288 MICHIGAN SURVBT, 1908. 

Banger "Maine; Provinces of Quebec and Ontario, Canada." (Dall.) 
Also Northern Michigan. 

Michigan : Isle Royale, Ontonagon Coanty and Charlevoix. 

iBle Royale: Not collected in 1905. See Report Exped. 1904, p. 97. 

11* H elicodisais parallehig (Say). 

Range: Eastern United States, Florida and Texas, north to Manitoba. 

Michigan: Generally distriboted. 

Isle Koyale : V, 2, Heath Zone and Beach near Siskowit Bay, Nos. 
118 A, 129 A, 130 A. 

All found among or under Cladonia, upon the rock beach slope. 

23, Punctum pygmaiam (Dr.). 

Range: "United States generally; Quebec; Manitoba; Victoria, Van- 
couver Island, Europe," (Dall), 
Michigan: Generally distributed. 
Isle Royale : Not collected in 1905. See Report Exped. 1904, p. 97. 

24. Spkyradivm edcniulum (Dr.). 

Range: "Northern Europe, Asia and America." (Dall). 
• Michigan : Generally distributed. 

Isle Royale: Not collected in 1905, See Report Exped. 1904, p. 97. 
One of the specimens under No. 8 may belong here. 

25.* CootUicopa luirica (Mull,), 

Range: "Europe, North Africa and Asia Minor; Siberia; Kamchatka; 
most of North America," (Dall). 

Michigan : Generally distributed. 

Isle Royale: 11, 1, Ransom Clearing, No. 150; V, 2, Heath Zone and 
Beach near Siskon-it Bay, No. 130 A. 

No. 160 was taken under a fallen log in an open place near the lake 
and No. 130 A under Cladonia upon the rock beach. 

26." Vallonia pulchclla (Mull.). 

Range: Europe; North Africa; Southern and Western Bib^a to the 
Amur; Madeira; the Azores; North America from Manitoba to Florida 
and Montana to Nova Scotia." (Dall). 

Michigan: Generally distributed. 

Isle Royale: A single specimen only occurred in the collections, the 
exact locality of which is uncertain, 

27.* Vallonia ooaUita (Muller). 

Range: Europe; Northern United States and northward. 

Michigan: Owobho, Monroe and Isle Royale. 

Isle Boyale : II, 1, Benson Brook, No. 150. Apparently a rare speciea 
in Michigan, and this the first record in the Upper Peninsula. Under a 
fallen log in an open sunny place near the lake. 

28." Succinea ovaiis Say. 

Range: "From Ijouisiana to Hudson Bay and eastivard to New Eng- 
land and Gaepe, but not west of the Mississippi valley." (Dall), 

Michigan: Generally distributed. 

Isle Royale: Y, '04, Tamarack Swamp, No. 145 A. A single dead 
shell was found at the margin of a small stream flowing from the 
swamp. 

29.* Sucdnfu retusa Lea. 

Range: "Northern United States, from Kentucky northward to 
Canada and British America." (Dall). , OoO*^?lc 



ECOLOGY OF ISLE R6TALB. 289 

Michigan: G«Deral1y diBtributed. 

lale Eoyale : Only a single specimen was collected, the exact locality 
of which waB loet. 

30, Carjfchium exile canadense Clapp. 
Hange: I^ortbern United States and Canada. 

Michigan : Generally distributed north of the Saginaw-Qrand Valley. 
Isle Eoyale: Not collected in 1905. See Report Exped. 1904, p. 97., 
where it is listed as Carychium exile. 

31. Lt/mn<sa stagnaUa (L.). Fig. 63. 

Range: "Europe; the Caucasus; Western and Northern Asia; the 
Northern United States; Canada and British America." (Dall). 

i^Iichigan : Generally distributed. 

Isle Royale : 

Variety A. (Fig. 63, No. 1.) II, 1, Month of Benson Brook, No. 167; II, 
4, McCargoe Cove. No. 53; III, 2, Small Island in Rock Harbor, No. 89; 
III, 3, Bulrush Zone, Head of Kock Harbor, Nos. 161, 162, 168; III, 4, 
Sumner Lake Trail, on Bock Harbor, No. 166 ; HI, 5, Sumner Lake, No. 
155; III, 6, Southwest Coves of Rock Harbor, No8. 91, 95; North side of 
Rock Harbor. No. 110; IV, 6, Small Island in Tobin Harbor, No. 123; 
Washington Harbor (Wood). 

Variety B. (Fig. 63, No. 3.) I, 1, Lake and Bay Beaches, Nos. 32, 50, 57; 
3rd Cove below Camp on Light-house Peninsula, No, 7 ; II, 1, Mouth of 
Benson Brook. No. 54; III, 4, Head of Sumner Lake Trail, (Wood) ; V, 
1, Beach at Siskowit Bay, No. 200. 

Variety C. (Pig. 63, No. 6.) V, 6, South shore of Siskowit Lake, Nob. 
199, 210, 211, 217. 

Notes on the babitats of this species are giv«i in detail in the chapter 
by H. A. Qleason. 

This lai^ and widespread species seems in tbis coontry at least, to 
exhibit its greatest variability in the La^e Superior B^on. Two well 
marked varieties have already been described; one var. higleyi Baker 
from Michipicoten Island on the north shore, and the other var. aanct/p- 
mariw Walker from the St. Mary's River. It is apparently one of the 
inost abundant species on Isle Royale, where three very distinct for;>ia 
are represented, none of which are typical and none exactly coincident 
with any of the described varieties. Al! the specimens collected fall 
into one of these groups which are apparently correlated with definite 
local conditions. For present purposes they may be designated as 
varieties A. B. and C. Variety A (Fig. 63, No. 1) most closely approaches 
to the usual North American form known as var. appreasa Say (Fig. 63, 
No. 4) from which it differs mainly in the pear-shaped rather than regu- 
larly rounded aperture. It is characteristic of the quieter waters of the 
long, narrow harbors which are such a remarkable feature of the island. 
The same form has also been collected in the St. Mary's River near the 
Neebish Rapids. Variety B is an inhabitant of the shores exposed to the 
more violent wave^ of the main lake. Correlated with these conditions 
the shell is smaller, with a short spire and a relatively large body whorl 
for the accommodation of the large foot necessary to enable it to re- 
tain its hold upon the rocks, among which it lives. This form (Fig. 63, 
No. 3) is more nearly related to the var. higleyi (Fig. 63, No. 5) from the 
north shore, but is apparently much smaller. It is about the size of the 



200 MICHIGAN SURVKT, 1908. 

Tar. sanctwmaria (Fig. fi3, No. 2) but quite different in the shape of the 
spire. Variety C. (Fig. 63, No. 6) is the largest in cubic capacity yet 
known from this country. It was found only in Siskowit Lake, whose 
quiet waters and especially favorable conditions have conduced to the 
production of this unusually fine, thin, inflated form. 

32.* Lymnaa megasoma Say. 

Range: "Northern New England, Canada, and British America to 
Lat. 57° N." (Dall). Also Northern Michigan, Wisconsin and Minnesota. 

Michigan: Isle Hovale, St, Mary's River and Roscommon County. 

Isle Royale: IV, 3, Bayou at Tobin Harbor, No. 124, (Adams). A siD- 
gle large, but dead, specimen was found in a pond-like bayou which was 
connected with Tobin Harbor by a verj- narrow and short outlet. 

33. Lymnwa emarginata Say. 

Range : "Northern United States east of the Mississippi, Canada and 
northward." (Dall), 

Michigan : Shores of the Qreat Lakes north of Saginaw Bay and some 
inland lakes from Roscommon County northward. 

Isle Royale: I, 1, Lake and Bay Beaches, Nos. 24, 50, 57, 58, 59, 74, 
118, and 125; 3rd Cove west of Camp on Light-house Peninsula, No. 7; 
III, 4. Head of Sumner Lake Trail, (Wood); V, 1, Beach at Siskowit 
Bay, No. 200. 

This is a characteristic and abundant species along the lake beach 
where it was found associated with var. B. of Lymnwa stagnalis. The 
same form, but with a rather heavier ^ell, is very abundant along the 
shore of Mackinac Island. Specimens from one rock pool, No. 58, are 
peculiar in being longitudinally striped with white like Lymnaa refiexa 
zebra Tr^on. Those from another, Nos. 58 and No. 74, are nnicolored. 

34. Lymtura piUbryana Walker. 
(Nautilus, XXII, p. 4, PI. I, fig. 2, 8-11, 1907). 
Range : Isle Royale is the only known locality. 

Isle Royale: X, '04, Washington Harbor, No. 1 (Wood). TTiis form, 
so far as the collections show, is apparently confined to the west end 
of the island. It was doubtfully referred to LymruBa aumassi Bd. in 
the Report Exped, 1904, p. 97. A larger suite of specimens from the 
original locality in Washington Harbor was collected in 1905, including 
a few full grown examples which confirm its specific distinctness. 

35.* Lymrura ohrusaa Say. 

Range: "Northern United States and Northward." (Dall). 

Michigan: Generally distrlbnted. 

Isle Royale: III, 2, Small Island in Rock Harbor, No. 89; III, 3, 
Bulrush Zone at western end of Rock Harbor, No. 164. 

36." hymnaa catascopium Say. 

Range: "Northern United States to Rocky Mountains, Canada and 
northward," (Dall). 

Michigan : Shores of the Great Lakes and connecting rivers and lower 
waters of tributaries in northern counties. 

Isle Royale: North shore of Bock Harbor, No. 110; III. 3, Bulrush 
Zone at western end of Rock Harbor, Nos, 160, 163, 164; III, 6. South- 
west Coves of Rock Harbor, No. 91; IV, 3, Island No. 14, Tobin Harbor, 
No. 30 A; IV, 6, Small Island in Tobin Harbor, No. 123; V, 6, South 
shore of Siskowit Lake, No. 220. , , , 

iX.ooglc 



ECMJLOGT OF ISLE ROYALE. 291 

Apparently iDoet frequent ia flhallow water in placee aheltered from 
the wavea, but No. 220, a single very young specimen, was collected on 
the nniJer surface of a water-lily leaf. 

With one exception, the specimens from all these localities are alike 
and belong to the common, rather short, lake form of this apeciee. As- 
sociated with this form at Station IV, 6, was a very thin, elongated 
form with the characteristic sculpture of catastxypium, which is closely 
related to, but much more fragile than, the elongated form, which is 
characteristic of the lower Great Lakes. 

37. Iitmncta sp? 

Isle RoTale: III, 3. Bulrush Zone at western end of Rock Harbor, 
No. 163 ; iV, 2, Island No. 14 in Tobin Harbor, No. 126. 

At boih these localities occurred a few dead, fragmentary and more 
or less decayed specimens, which could hardly be referred to any of the 
species listed above, and yet were too imperfect to saccessfully identi^. 

38. Phym sfKfii Tapp. 

Range: Northern United States and Canada. 

Michigan : Generally distributed. 

Isle Royale: I, 1, Lake and Bay Beaches, Noe. 50, 57, 118, 125; 3rd 
Cove west of Camp on Light-house Peninsula, No. 7; III, 4, Head of 
Sumner Lake Trail, (Wood); V, 1, Beach at Siskowit Bay, No. 200; 
Washington Harbor, (Wood). 

The specimens from Washington H8rb<H: Eire of normal thickness and 
more nearly typical in shape than those from the other localities, which 
are unusually thin. 

More detailed notes on the local distribution of this species are given 
in the chapter by H. A. Gleason. 

30.* Phiyta gyrtna Say. 

Range: "The United States east of the Mississippi, Canada and north- 
ward." (Dall). 

Michigan : Generally distribnted. 

Isle Royale: II, 5, Forbes Late, Noe. 71 A, 90; III, B, Sonthwest 
Coves of Kock Harbor, No. 91. 

Specimens No. 71 A were found on driftwood In water a few inches 
deep. 

The specimens from Forbes Lake are a very large, inflated form. Those 
from the other locality are much smaller and may be one of the varying 
forms of No. 41, though closer to typical gyrina than those included 
under that head. 

40.* Physa aplectoides Sterki. 

Range: "Tuscarawas County, Ohio, and elsewhere." Sterki. 

Michigan: Isle Royale. 

Isle Royale: V, 11, Tamarack Swamp, No. 128 A. 

Taken from foot-print pools in the Sedge and Buck Bean Zone 
about a small ■poaA. 

The occurrence of this minute species so far from its original locality 
in Ohio, was one of the surprises of the 1905 collection. It is a very 
distinct form resembling a young Aplexa hypnorum in shape but beauti- 
fnlly sculptured, especially on the apical whorls. The identification ia 
based on comparison with topotypes of aplectoides received from Dr. Y. 
Sterki. .-. , 

lyCoogle 



292 MICHIGAN SURVEY, 1908. 

41. Phyaa sp? 

iBle Boyale: II, 1, Benson Brook, No. 149, 167; III, 2, Island at 
West end of Rock Harbor, No. 89; ,11, 3, Bulrush Zone at western end 
of Rock Harbor, Nos. 161, 162, 163, 164 ; North shore of Rock Harbor, 
No. 110; III, 5, Stunner I^ake, Nos. 77 A, 78 A, 79 A; IV, 5, Neutson's 
Resort at Bock Harbor, No. 44 A; IV, 6, Island in Tobin Harbor, No. 
123 ; IV, 7, Head of Tobio Harbor, No. 127 ; V, 1, Beact near Siskowit 
Bay, No. 200; V, 6, South shore of Siskowit Lake, Nos. 220, 221; V, 
9. Outlet of Siskowit Lake, No. 238; V, 11, Swamp near Siskowit Bay, 
Ko. 126 A. 

Under this head are included nearly all the Physw from the harbors 
and interior waters which, although exhibiting considerable rariatioD 
in shape and size, appear to be variation of a common form. Most o( 
the specimens are Immatnre. The few mature examples at first glance 
would naturally be referred to Physa heterostropha Say, but the uniform 
sculpture of the apical whorls, which becomes more or less obsolete on 
the body whorl of the mature shell, forbids their reference to that species. 
The sculpture is that of gifriiia and the form may ultimately referred 
to that species as an extreme form, but the shape of the immature shell, 
its small, acute spire and deeply impressed suture is quite ditfer^it 
from that of typical gyrina. In the present chaotic state of the nomen- 
clature of the American species of Physa it would seem the better policy 
to refrain from any attempt at specific identification than to run the 
risk of adding to an already , over-burdened synonymy. 

42. Aplexa hypnorum (L.). 

Range : "Northern Europe, Asia and America, Northern United 
States and Canada," (Dall). 
Michigan : Generally distributed. 
Isle Royale: Not collected in 1906. See Report Bxped. 1904, p. 98. 

43. Planorbia trivolvw Say. 

Range: "Entire Atlantic Drainage of North America; and the Mis- 
sissippi Valley and northward." (Dall). 
Michigan: Generally distributed. 
Isle Boyale : III, 5, Sumner Lake, Nob. 78 A, 135. 
in a small pool in the sedge zone of a tamarack swamp. 

44. Planorbia bicartnatua Say. 

Range: "The United States East of the Rocky Mountains; Eaatem 
Canada; Oregon." (Dall). 

Michigan: Generally distributed. 

Isle Boyale; III, 2, Small Island in Rock Harbor, No. 89; III, 3, 
Bulru-iah Zone at western end of Rock Harbor, Nos. 159, 160; III, 6, 
Southwest Coves of Rock Harbor, No. 91. 

Dredged from the mud Itottom at the upper end of Rock Harbor, near 
mouth of a small stream, in 3-5 feet of water (Nos. 159, 160). 

44a, var. strtatuH Baker. 

Isle Koyjile: III. 3, Bulrush Zone at western end of Bock Harbor, 
No. 102; III, 5, Sumner Lake, Nos. 78 A, 79 A. 

Dredged from the mud bottom of the small sluggish stream at the 
head of Book Harbor (No, 162) and from the margin and sedge zone 
of Sumner Lake (No. 78 A, 79 A.). 



3y Google 



ECOLOGT OP ISLB ROTALB. 293 

44b," var. fvyalensis Walker. 

(Nautilns, XXII, p. 9-10, PI. I, fig. 11, (1909). 

Isle Eojale: V, 6, Sonth shore of Siakowit Lake, No. 210. 

This novel and very distinct form, characterized by its very acate 
carinie and rough, irregularly corrugated surface, was one of the most 
interesting discoveries of the expedition. 

In the mud among loose stones at a depth of about 1 foot. 

45. Planoriia campanulatus Say, 

Range: "Atlantic, Mackenzie and Hudson Bay water sheds and north 
to Great Slave Lake." (Dall). 

Michigan : Generally distributed. 

Tale Boyale: III, 5, Sumner Lake, Nos. 78 A, 79 A; V, 6, South shore 
of Siskowit Lake, Kos. 210, 211. 

In mud and among loose stones at a depth of about one foot (Nob. 
210, 211) and in small pools in the Sedge 7one (Nos. 78 A, 79 A). 

The specimens from Sumner Lake are the usual form. Those from Sis- 
kowit Lake are a peculiar variety resembling the rare Planorhis multi- 
volvis Case, in having apparently the apical whorls elevated above the 
line of the body whorl. Unfortunately the upper surface of all the speci- 
mens collected is so eroded that it is impossible to determine juat what 
degree of elevation the spire of the perfect shell attains. 

46. Planorlis exacuous Say. 

Eange: "Northern United States, east of the Bockies; Canada, etc., 
Bouth to New Mexico." (Dall). 

Michigan : Generally distributed. 

Isle Boyale: III, 2, Small Island in Bock Harbor, No. 89; III, 3, 
Bulrush Zone at the western end of Bock Harbor, Nos. 159, 160, 161, 
162. 

In the muddy bottom of a small stream Sowing into Bock Harbor and 
in the Harbor itself, at a depth of 2-5 feet (Nos. 159 to 163). 

47. Planorhis parvus Say. 

Bauge: "Eastern North America from. Florida to North Lat. 67-°, 
and the Yukon Drainage System." (Dall). 

Michigan: Generally distributed. 

Isle Boyale : I, 1, Rock Pool, No. 2, Light-house Peninsula, No. 59 ; 
III, 2, Small Island at Bock Hart)or, No. 89; III, 3, Bulrush Zone at 
western end of Bock Harbor, Nos. 159, 160, 163, 164; III, 5, Sumner 
Lake, No. 79 A; III, 6, Southwest Coves of Bock Harbor, No. 91. 

As with the last species (Nos. 159, 160,, 163 and 164). 

48. Planorbis hirsutus Old. 

Range: '"Washington, D. C, northward east of the Mississippi." 
(Dall). 
Michigan: Generally distributed. 
Isle Boyale: Not collected in 1905. See Report Exped. 1904, p. 98. 

49. Ancybis sp? 

Isle Boyale : I, 3, Balsam-spruce Forest, No. 140. 

"In the damp leaf mold in the dense shade of the balsam-spruce 
forest." 

A single broken specimen, too much damaged to identify spocifically, 
was the only one obtained. There is apparently some mistaJie in regard 
to the locality where this specimen was found. /^ -,nl,> 

50. Talvata tricarinata Say. v^.UOyiL 



294 MICHIGAN SUHTBT, IMS. 

Bange : "From New England and Virginia westward to Missouri and 
northward." (Dall). 

Michigan: Generally distributed. 

Isle Royale; III, 2, Small Island in Rock Harbor, No. 89; III, 3, 
Bulrush Zone at western end of Rock Harbor (Nos. 160, 163). 

In the mad in deep water at the head of Bock Harbor (Nos. 160, 
163). 

51. Yalvata leteisii Currier. 

Range : "Northern United States from Atlantic to Pacific and North- 
ward." (Dall). 

Michigan : Generally distributed. 

Isle Royale; Not collected in 1905. See Report Exped. 1904, p. 98, 
cited as Valvata aincera lewi»ii. 

52. Valvata aincera nylanderi Dall. 

Banj;^: Northern United States from Maine to Wisconsin. 

Michigan : Isle Royale and Marquette County. 

Isle Royale: III, 2, Small Island in Rock Harbor, No. 89; III, 3, Bul- 
rush Zone at western end of Rock Harbor, Nos. 159, 160, 163, 164; V, 
6, South Shore of Siskowit Lake, No. 220. 

With Valvata tricarinata at the head of Bock Harbor and in shallow 
water in Siskowit Lake (No. 220), especially abundant on the lower 
side of water-lily leares. 

53. Amnicola limoaa (Say). 

Range; "Virginia to Wisconsin and Hudson Bay." (Dall). 
Michigan : Generally distributed. 

Isle Royale: V, 6, South shore of Siskowit Lake, No. 220, living in 
company with the preceding species under water-lily leaves. 

54. Amnicola histrica Pile. 
Range : Northern United States. 
Michigan; Generallv distributed. 

Isle Royale : III, 2, Small Island in Rock Harbor, No. 89 ; III. 3, Bul- 
rush Zone at western end of Kock Harbor. Nos. 159, 160, 163, 164. 

In muddy bottom in deep water (3-5 feet deep) at the head of Rock 
Harbor. 

55. iMmpailis Juteola (Lam). 

Range : "Entire Mississippi drainage and north to the Bed River of 
the North." (Dall). 

Michigan: Generally distributed. 

Isle Royale: V. 6, South shore of Siskowit Lake, Nos. 210, 211, 218. 

Some of the specimens collected are very similar to the form from the 
Beaver Islands. Lake Michigan, referred to Lampsilis ioreaJis Gray, but 
they are connected by intermediate specimens with the more tyi>ical 
form and seem I'iither referable to this species than to horcalis. 

56. Anodonta grandis footiana Lea. 

Range: Northern United States and northward. 

Michigan: Generally distributed. 

Isle Royale; II. 4. McCargoe Cove. No. 52; III. 2, Small Island in 
Rock HRPhor, No. 89; III, 3, Bulrush Zone at it^estern end of Bock 
rrarlior, No, 168 ; III, 4, near head of Trail to Sumner Lake, Bock Har- 
bor. No. 93; III, 5. Sumner Ijake, No. 154; III. 6. Southwest Coves of 
Rock Harbor, Nos. 91, 91, 156; South Side of Rock Harbor, Nos. 109; 



ECOLOGY OF ISLE ROTALE. 205 

V, 6, South Shore of Siskowit Lake, Nob. 210, 211, 218; Sargent Lake, 
No. 112. 

Abundant in all of the takea, especially on sandy or gravelly Iwttom 
in the smaller coves sheltered from the waves. 

57. Anodonta marginata Say, 

Bange: "Drainage of the St. Lawrence River basin, including the 
Lakes." (Dall). 

Michigan : Generally distributed. 

Isle Koyale: II, 1. Benson Brook Clearing, No. 80 A; II, 5, Forbes 
I^ake, No. 90; III, 5, Sumner Lake, Nos. 94, 120, 135, 139, 154, 186, 
(Wood) ; IV, 3, Bavon at Tobin Harbor, No. 124; V. 6, South shore of 
Siskowit Lake, No. 210; VII, '04, Lake Desor, No. 139 A. 

58.* Sphaerium gimile (Say). 

Range: "United Staites east of the Mississippi Biver; Canada, Mani- 
toba." (Dall). 

Michigan: Lake Michigan and Lake Superior. 

Isle EoyaJe: III, 5, Sumner Lake. Only two immature valves taken. 

59.* Sphaerium loalkeri Sterki. 

Range: Lake Michigan and Northward. 

Michigan : Generally distributed. 

Isle Royale: III, 2, Small Island in Rock Harbor, No. 89. A single 
fragmentary specimen is doubtfully referred to this species by Dr. V. 
Sterki. 

60." Muscutium securts (Prime). 

Range : Northern United States, Maine to Minnesota. 

Michigan: Generally distributed. 

Isle Royale: III. 3, Bulrush Zone, Rock Harbor, No. 160; III, 5, 
Sumner Lake, Nob. 77 A, 78 A, 79, 176 ; V, 9, Outlet of Siskowit Lake, 
No. 238. A few examples only of a small form. Some immature ex- 
amples from the latter locality "may be the same." 

In small, shallow pools in the outlet of Siskowit Lake (No. 238) and 
at the margin and Sedge Zone of Sumner Lake (No. 77 A, 78 A). 

61.* Pisidium idajioenge Roper. 

Range: Idaho; Washington; Alaska; Lake Michigan; Lake Saperior. 

Michigan: Lake Michigan and Lake Superior. 

Isle Royale: IIL 2. Small Island in Rock Harbor, No. 89; III, 3, 
Bulroflh Zone, Rock Harbor, Nos. 159, 160. 162, 163. Rather common. 
The Isle Royale form is similar to that from Lake Michigan and is neither 
as lai^e nor as inflated as the typical form. 

Dredged from a muddy bottom in 2-5 feet of water at the head of 
Bock Harbor and in a small stream (lowing into it. (Nos. 159, 160, 
162. 163.) 

62. Pisidium variabile Prime. 

Range: "Eastern United States, north of Virginia; Colorado anil 
northward." (Dall). 

Michigan : Generally distributed. 

Isle Royale: III, 5, Sumner Lake, 77 A; III. 2. Small Island in 
Rock Harbor, No. 89; IIT, 3, Rnlrush Zone, Rock Harbor, Nos. 160, 
162, 163; V, 9. Outlet of Siskowit I^ake, No. 338. 

With the last at the bead of Bock Harbor, and also in the small 



lyCoogle 



296 MICHIGAN SURVEY, 1908. 

pools with gravel bottom in the outlet from Siskowit Lake and at the 
margin of Sumner Lake. 

62a.* Tar. brevius Sterki. 

Range: ' "Michigan; Minnesota and Keewatin." '(Sterki.) 

Michigan : Upper Peninsula and northern part of the Lower Pen- 
insula. 

Isle Rojale: III, 3, Bulrush Zone, Hock Harbor, Nos. 160. 164. All 
the examples both of the typical form and the variety are "Bmall and 
mostly immature." 

With the typical form in the mud bottom in 3-5 feet of water at the 
head of Kock Harbor. 

63." Pisidium affine Sterki. 

Range: "Great Lake Region, Michigan to New York; Minnesota. 
Illinois and Ohio (Ohio River Drainage)." (Sterki). 

Michigan: Generally distributed. 

l^e Royale: III, 5, Sumner Lake, Nob. 77 A, 79 A. A few ex- 
amples, "quite small," from the margin and Sedge Zone. 

64.* Pisidium sargenti Sterki? 

Range: "Northern United States, New Y(ffk to Minnesota." (Sterki). 

Michigan: Generally distributed in Lower Peninsala; Isle Royale. 

Isle Royale: III, 5, Sumner Lake, No. 176. Two specimens only, 
which "may be Piaidimrt sargen-ti. One example is large, especially in 
contrast with the small forms of tbe other species. In the smaller 
specimen the hinge is partly reversed." 

65.* Pisidium 8(mt€llatum Sterki. 

Range: Northern United States, Michigan to Washington and 
northward. 

Michigan : Generally distributed. 

Isle Royale: III, 2, Small Island in Rock Harbor, No. 89; III. 3. 
Bulrush Zone, Rock Harbor, Nos. 159, 160, 163, 16i; "Small, north- 
em variety. The most common species and somewhat variable." 

In the mud and silt bottom in 10 inches to 5 feet of water at the 
upper end of Rock Harbor (Noa 159, 160, 163). 

66.* Pisidiwn roperi Sterki. 

Range: Northern United States, Maine to Minnesota. 

Michigan : Generally distributed. 

Isle Royale: III, 5, Sumner Lake, No. 78; IV, 8, Trail to Green- 
stone Range, No. 128. "Small, but good and characteristic." 

67.* Pisidium ventricosum. Prime. 

Range: Northern United States, Maine to Michigan and northward. 

Michigan: Western part of the State, Kent County to Charlevoix 
County ; Marquette County ; Isle Royale. 

Isle Royale: III, 5, Sumner Lake, Nos. 77 A, 79 A; III, 2, Small 
Island in Rock Harbor, No. 89; III, 3, Bulmsh Zone at Rock Harbor, 
Nos. 163, 164. 

In 10 inches of water in the Potamogeton Zone at the mouth of a 
creek at the upper end of Rock Harbor (No. 163) and at the margin 
and in the sedge of Sumner Lake. 

68.* Pisidium snJtrotundam Sterki. 

Range: "New England; Anticoeti Island to Michigan." (Sterki). 

Michigan : Kent, Marquette and Ontonagon counties and Isle Royale. 



ECOLOOT OF ISLB ROYALS. 297 

iBle Boyale: I, 6, Sphagnum-Sproce Bog, No. 116; IV, 8, TWul to 
Greenstone Range, No. 128; III, 3, Bnlrusli Zone at western end of 
Kock Harbor, Nos. 159. 160; I, 4, Tamarack and Arbor Vita Swamp, 
No8. 181, 1^; V, 5, Tamarack Swamp, No. 237; V, 9, Outlet of Siakowit 
Lake, No. 238. A "form" of this species "common and somewhat vari- 
able." 

Am<mg dead leaves and sedges at the bottom of shallow pools in a 
tamarack swamp (No. 116). In silt and debris on the bottom in 4-S 
feet of water (Noa. 1H9, 160); small sphagnam-lined pools, seldom 
exceeding six inches in depth in dense shade (Nos. 181, 182); among 
sphagnnm and Utricularia in smalt streams and pools, mostly in the 
sun (No. 237) ; in shaded, shallow pools with gravelly bottom (No. 
238). 

69.* Ptstdiian rotundatwn Prime. 

Range: Northern United States, Maine to Minnesota and north- 
ward. 

Michigan : Generally distributed. 

Isle Royale: I, 6, Sphagnum-spruce Bog, No. 116; III, 3, Bulrush 
Zone, Eock Harbor No. 160; V, 5, Tamarack Swamp, No. 237; "Few 
and probably none mature." 

Among dead leaves and sedges at the bottom of shallow pools, in 
shade (No. 116) ; among sphagnum and Utricularia in small, shallow 
streams and pools, mostly in the sun (No. 237.) 

70.* Piaiddum splendidulum Sterki. 

Range: Northern United States, Maine to Michigan. 

Michigan : Generally distributed. 

Isle Royale: III, 5, Sumner Lake, Nos. 77 A, 79 A, 176. At margin 
and in the Sedge Zone. 

71.* Piaidium pauperculum Sterki. 

Range: Northern United States, Maine to Minnesota. 

Michigan : Generally distributed. 

Isle Royale: III, 3, Bulrush Zone at western end of Rock Harbor, 
No. 164. A few specimens of a small form. 

72,* Piaidium medianum Sterki. 

Eauge: Northern United States, Maine to Wisconsin. 

Michigan: Generally distributed. 

Isle Royale: III, 5, Sumner Lake, Nos. 77 A, 78 A, 79 A; III, 2, 
Small Island in Rock Harbor, No. 89; III, 3, Bulrush Zone at western 
end of Rock Harbor, Nos. 160, 164 ; V, 9, Outlet of Siskowit Lake, 
No. 238. 

Near the mouth of a small creek, on a silt and mud bottom at a 
depth of 5' feet (No. 160) ; In shallow, shaded pools with gravellv 
bottom (No. 238). 

73.* Pisidium pwustatwn aimples; Sterki. 

Range: "Michigan, Wisconsin and Illinois." (Sterki.) 

Michigan: Carp Lake, Emmet Co. and Isle Royale. 

Isle Boyale: III, 3, Bulrush Zone at Western end of Rock Harbor, 
No. 160. 

74.* Piaidium milium Held. 

Range: Europe; Maine and Michigan. 

Michigan: Generally distributed. ,-. , 

39 DijtizBdbyLlOOglC 



298 MICHIGAN SUHVET, 1908. 

Isle Royale : III, 3, Bulmsh Zone at western end of Bock Harbor, 
No8. 160, 162. !No. 162 occurred in a smaJi creek near its mouth, on a 
silt and debris-covered bottom at a depth of 3 feet. 

75. Pisidivm. ahditum Haldenian. 

Range: "Nortb America, from Honduras, north to Alaska." (Dall). 

Michigan ; Generally distributed. 

Isle Royale: Not found in 1905." See Report Exped., 1904, p. 98. 

76. Pisidmm sp.? 

Isle Royale: Undetermined forms of Pisidium were collected in the 
following localities: III, 3, Bulmsh Zone at western end of Rock 
Harbor, Nos. 169, 160, 163; V, 5, Tamarack Swamp, No. 237. 



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ECOLOGY OP ISLE ROYALB. 



REPORT OX THE ISLE ROYALE ORTHOPTERA OP THE 1905 
EXPEDITION. 



1. General Remarks. 

The orfhoptera secured by the University Museum expedition of 1905 
to Isle Royale eonsist of representatives of 13 spepieB. Of these, one 
is a cosmopolitan roach, tlie Croton-bug, BlatteUa germanica, doubt- 
less introduced in nierchaudise through man's agency. The others, with 
a single exception, are boreal Acridians characteristic of the cooler 
parts of central and eastern North America, whose presence in this 
locality was either known or to be confidently expected. The exception, 
MeUmoplas ala«kantis, is a species hitherto known only from the North- 
west, whose presence on Isle Royale, in sufficient numbers to make it 
appear to be the dominant species of its genus there, was, to say the 
least, unlooked foK It would be of much interest in this connection to 
know the status of this species on the adjoining northern mainland. 

Some striking variations are present in the representatives of certain 
species and will be found noted in the following list. In general, there 
seems to be a tendency toward an increaiied duskiness of coloring as 
shown by Camnitla ptAlueida, Hippiscus tuberculatus. and Circotettix 
vemiculatus (all geophilous species), due perhaps to humidity, per- 
haps to environmental coloration; also, as compared with eastern ma- 
terial, the specimens are of relatively large size, particularly in the 
cases of Melanoplus ej^tremus and faaciatus. 

The locust societies represented consist of campestral and thamnoph- 
ilous groups, though all of the species are to be obtained in the clear- 
ings or about their edges, rather than in the forest. 

The thanmophile species are Chloealtis conspcrsa and abdominalis, 
Melanoplu« fasciatus and httroni. The campestral species, notwith- 
standing the generally forested character of the country, are more 
numerous and consist both of phytophilous and geophilous species. In 
the damper situations occur iStenohothrus curtipeiin^K, Mecostethiis Une- 
attis, Alelanoplus extreniug and femoratns. In drier places Camnv-la 
pelluGtdaf Meldtwpliia aUiakantis, and Bippiacvs tuierculatus are to be 
found. Characteristic of the bare rock ridges is the saxicolous Circo- 
tcttiat vemiculatus. The two latter species, with their strikingly colored 
wings and noisy Sight, are the most conspicuous members of the orthop- 
terouB fauna of the island. 

The 1904 expedition secured examples of a Tettigid (Tettix acndiais) 
and of a wingless Ix)custarian — Ceuthophilus aecUisus. The B|>ecie8 of 
CeutlMphihis usually inhabit damp, dark places, under bark, in hollow 
logs, etc., and are among the most characteristically sylvan of our oi^>ol(^ 



tboptera. 



300 MICHIGAN SDEVBT. 1908. 

The lot Qimibers refer to the field numbers of the collectors, thtwe 
without a letter to H. A. Oleaaon and those followed by an A. to C. 
C. Adams. 

2. Annotated List of Spedcs. 

Blattidce. 

1. BlottelUi germanica Linn. Croton-bug. Station I, 7, camp. Lot 
179, July 28. A single female with o6theca. Probably introduced Id 
merchandise. 

AcridHdae. 

2. Stenoiothrus citrtipennis Harr. Stations I, 7, camp; V, 3; T, a; 
I, '04 and VIII, '04. Lota 99 A., 134 A., 137, 228, 232. July 24 to 
Aug. 24. 

This species is a characteristic inhabitant of moiat, grassy or se^ 
meadowa. It waa taken in the caasandra and hnnunock zone at V, 5. 
and in a clearing near end of Wendigo road at Washington Harbor. 
Both long — and abort-winged forms were secured. 

3. Cttloealtis abdormnalis Thom. Stations II, 3; IV, 5; and I, '04. 
Lots 121, 143, 154 A. July 21, 25, Aug. 24. A young male in 5tli stage 
on July 21. 

4. Chloealtis con»perta Harr. Stations I, 1 ; II, 3 ; and I, '04. Lots 
22, 143. 141, 147, 354 A. July 6 to Aug. 24. A young miile in 5th stage 
on Jnly 6, Some of the males of this species show indications of the 
more closely reticulated spot in the tegmina so characteristic of aii- 

■dominalis. 

The species of this genus are dwellers in thicket and woodland edges, 
ovipOBiting usually in decayed, though sometimes in firm, wood. 

5. Mecostethua lineatva Scudd. Stations- II, 5; V, 5; V, 11; and I, 
'04. Loft 91 A.. 136 A., 154 A., 180. July 8 to Aug. 24. 

Numerous immature examples of this genus are referred to thi« 
species with some doubt, and it is quite poBsible that aoine of them be- 
long to an allied species. The hind tibiae of these young specimens are 
markedly fuscous. 

6. Camnula pcUucida Scudd. Stations I, 7; IV, 5; V, 3; and I, '04. 
Lots 121, 133, 222, 228, 154 A. July 21 to Ang. 24. Young in 4th and 
5th stages on July 21 and 22. 

The representatives of this species, like those of some others, are im- 
usually dark in color, a phase of coloration probably correlated with the 
hnmid climate or soil background. This is a campestral species, occur- 
ring plentifully from Atlantic to Pacific oceans in the boreal zone, 
usually upon dry, upland soil. 

7. Hippiacus tuberculatua Palis. Goral-winged Locust. Stations 
IV, 5; IV, 9; V, 9. Lots 121, 122, 215. July 21 to Aug. 7. 

This species is represented by 6 males, 4 females, which differ marked- 
ly from typical eastern examples in being deeply infuscated, the hind 
tibiae coral red except on basal half of outer side, and in baying tbe 
posterior process of the pronotum more produced. 

This large locust is an inhabitant of clearings on dry soil. Itn bright- 



ECOLOGY OF ISLE ROTALE. 301 

colored wings (whence the name of Coral-winged Locast ia derived) 
render it a conspicuona object during its powerful and usnally snatained 
flight. 

8. Circotettix verruculatus Kirbv. stations I, 5; I, 7; II, 3; III; 

IV. 5; V, 2; V, 3; V, 9. Lota 121, 131, 132, 144, 108, 147, 179, 208, 201, 

213, 215, 222, 239, 27, 107 A., 135 A. July 20 to Aug. 16. 

This BpecieB is represented bj nnmeroua specimena, in- general very 
dark in color, which were secured in the cladonia zone, the beach heath 
zone, rock clearings and ridges. It is a typically aaxicolons locnat, de- 
lighting to sun itself on outcropping ledges of rock or the neighboring 
patches of bare soil and usually presents a very cloae resemblance in 
coloration to its background. 

9. Melanoplm aJaakanus Scudd. Stations I, 7; II, 3; IV, 5; IV, 9; 

V, 2; V, 9: V, 11; and I, '04. Lots 55, 121, 122. 133, 137, 144, 146, 147, 
166, 179, 215. 216. 107 A.. 154 A., 136 A. July 21 to Aug. 24. 

This Bpeciea, described from Alaska, is apparently the dominant form 
of the genus occurring on Isle Boyale, to judge from its abundance in 
the material examined and the localities whence derived. It is recorded 
from clearings, along trails, rock ridgea, the beach heath zone, etc. As 
the original description was based on a small aeries of apecimena, 
measurements follow to show the range of size in the material at hand. 
The coloration varies much individually, recalling that of femur-rub- 
riim. in cool, moist regions. 

Length of body: c? 20-23.5; 9 , 23-29. Hind femora: &, 12-14; 2 , 12- 
14.5. Tegmina:d", 19-20.5; 2, 17.5-23 mm. 

10. Melanoplua extremus Walk.' Stations II, B; V, B; V, 11. Lota 
180, 99 A., 136 A. July 8 to Aug. 16. 

This species was found in the cassandra and sedge zone of the 
swamps, and on the jack pine ridge. It ia a typical inhabitant of moist 
meadows throughout the Canadian zone from Alaska to Nova Scotia. 
The examples secured are of large aize and measurements are ap- 
pended. 

Lei>gthof bodv: c?, 20-21; 9,26 29. Hind femora: d*, 11.7 12.7;5 ,13.5- 
14.6 Tegmina: c?. 11.5-14.5; 2, 13.5-15.5 mm. 

11. Melanoplua fasaatua Barnst.-Walk. Stations IT, 3; III; V, 2; 
V. 3; W, 5; V, 9; IIL '04. Lola 121, 131. 114. 146, 193, 207, 208, 212, 

214. 231, 213, 222. 239, 101 A.. 107 A., 135 A.. 143 A. Jnly 20 to Aug. 16. 
This is a common and widely distributed species in the procumbens, 

heath and cladonia zones. It is a tbamnophilous species in the east, 
and typically short-winged, but in the central part of the continent ex- 
amples with fully developed wings and tegmina are not rare, and both 
forms are represented among the material secured. The average size is 
considerably greater than that of specimens from the east and measure- 
men ta are appended. 

Length of body: c?, 19-23; 2, 23-27. Hind femora; d", 10.7-12; 2, 12- 
13.5. Tegmina; c?, 11.5-18-5 (average 12.5); 2, 10.5-18 mm. 

12. Melanoplus femoratM Burni. Station I, 7, camp. July 24. Lot 
137, a single male. 

13. Melanoplua huroni Blatchley. Stations I, Light-bouse Penin- 
sula; I, 5; III. 5; IV, 5; IV, 7. Lots 35, 44, 121, 131. 183, 35 A. July 

8 to 29. Seven females from dry, aspen-covered, burned-over ridge, rock ,1 -, 
clearing and ridges. cV 



302 MICHIGAN 8URVBT. 1908. 

The Melanopli have been determined fron^ adult examplea Bolely. 
There are in addition numerous immature specimens, in several stages, 
representing at least three epeeies and possibly more, whicb cannot be 
identified with certaiaty at present. 



3. Station List, 1905 Collections. 

I, 1. Lake Superior and Bay Beaches. 

Ckloealtis conapersa, {No. 22). 
1, 5. Jaelj Pine Bidge. 

Circotettix tx-rnirulatiis, (108, 27). 

Melanoplus huroni, (44). 

I, 7. Camp at Light-bouse Clearing. 

Blattclla germanica, (179). 
Stenobotkrva curtipentUs, (137). 
Camtuila pellucida, (133). 
Circotettix verrueulatua, (179). 
MeUinoplus alaskanua, (133, 137, 166, 179). 
Melanoplus femoratus, (137). 

II, 3. Bock Bidge Clearings on McCargoe Cove Trail. 

Chloealtis consperM, (143, 144, 147). 
Cliloealtia abdominalts, (143). 
Circotettix verructilatus, (147, 144). 
Melanoplus alaskanus, (144, 146, 147, 55). 
Melanoplus fasciatus, (144, 146). 

II, 5. Forbes Lake. 

Mecostethus lineatus, (ISO). 
Melanoplus extremus, (180). 

III. Western End of Hock Harbor. 

Circotettix verruculatus, (131, 132). 
Melanoplus fasciatus, (131). 
Melanoplus huroni, (131). 

III, 5. Sumner Lake. 

Melanoplus huroni, (183). 

IV, 5. Clearing at Neutson's Resort. 

ClUoealtis abdominalis, jav., (121). 
Camnula pellucida, (121). 
Hippiscus tubcrculatus, (121). 
Circotettix verruculatus, (121). 
Melanoplus alaskanus, (121). 
MeUinoplus huroni, (121). 
Melanoplus fasciatus, (121). 
rV, 7. Head of Tobin Harbor. 

MelanoplHs huroni, (35 A.). 

IV, 9. Mountain Top. 

Hippiscus tuhercuJatus, (122). 
Melanoplus alaskanvs, (122). 

V, 2. Heath Zone and Beach. 

Circotettix rcrniculatus, (A. 135, A. 107). 

Melanoplus alaskanvs. (107 A.). 

Melanoplus fasciatus, (101 A, 135 A, ^PT A^.Qqqq[(> 



ECOLOGY OP ISLE ROTALE. 303 

V, 3. Rock Clearing at Camp on Siakowit Bay. 

Stenobothrus curtipennig, (99 A., 228). 

Camnula pelludda, (222, 228). 

Circotettix verruculatua, (239, 222, 208, 201, 212). 

Melanoplus faaciatus, (193, 201, 208, 212, 214, 222, 239, 231). 
V, 5. Tamarack Swamp. 

SteTiobotkrus curtipennig, (99 A.)- 

Mecoetethua lincatus, (91 A.). 

Mekmoplus extrcmus, (99 A.). 
V, 9. OoHet to Siskowit Lake. 

Sippiacus tuberculatua, (215). 

Melanoplus alaakanua, (215). 

Melanoplua fasctatua, (215). 

Circotettix verruculatua, (215). 
V, 11, Tamarack-Spince Swamp. 

Mecoatethus Uneatua, (136 A.). 

Melanoplus alaalcanut, (136 A., 216). 

MeUinophia extremua, (136 A.). 
I, '04. Clearing ou the Shore of Washington Harbor. 

Stenohothrua curtipennia, {154 A.). 

Chlaealtia abdominalia, (154 A.). 

Chloealtis conapersa, (154 A.). 

Mecottethus lineatua, (154 A.). 

Camnula pellucida, (154 A.). 

Melanoplua alaakanua. (154 A.). 

Melanoplua ep. indet. 9 . 
Ill, '04. Trail along the Top of Greenstone Range. 

Melanophia faaciatua, (143 A.). 
VIII, '04. Western End of Siskowit Bay. 

Stenobothrua curtipennia, (232). 



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ECOLOGY OP ISLE ROTALB. 



NEUROPTEEOID INSECTS FROM ISLE ROYALE, MICHIGAN. 

DR. JA^ES G. NEEDHAM, COBNELL UNIVERBITX. 

A small bat intereBting collection of aquatic larvae of dragonflies, 
Btonefllee and mayfiies was obtained from Isle Royale, and the fine 
Btonefly, Pteronarcya dorsata Say, from the Northern Peninsula. Among 
the dragonfly larvae were two that are hitherto nndescribed, a species 
of Syrwpetrum too immature for description, and the cast skins and the 
young larva of a species of Somatochlora, described below. The list is 
as follows: 

Odonata. 

1. AnOfte Junius Drury. A young larva from Isle Boyale was col- 
lected on August 14 (No. 120 A) in a rock pool on the beach (V, 2) ; 
and another on July 29 at Summer Lake (III, 5). 

2. Aeachna sp.? perhaps constricta Say. Represented by both cast 
skins and nymphs from Sumner Lake (III, 5), Nos. 170, 221, 72 A, 
77 A, 78 A, 79 A; a rock pool on the beach {V, 2) on August 14; in the 
stomach of a duckling loon {Gavia imber) from Siskowit Lake (V, 6) 
August 10, No. 108 A; and from the mai^n of a swamp (V, 11) 
on August 16 (No. 126 A). 

3. Aeachna sp? A second species, represented by a single young 
larva, was taken in a rock pool (V, 2) on August 14 (No. 120 A).' It 
has lateral spines on segments B-9 of the abdomen, that of 5 (usually 
absent) being very small. , 

4. Somatochlora sp? Perhaps 8. forctpata Selys. (This suggestion 
as to the species is based solely on the fact that this species is known 
to occur commonly at Duluth, Minn.). A single young nymph in 
alcohol, July 26, from the head of Rock Harbor, No. 162; and a cast 
skin (No. 89) from Rock Harbor (III, 1) July 14, 1905. Being new to 
science, a description drawn from the cast skin is herewith offered. 

Length 23 mm., abdomen 13 mm., hind femur 7 mm., width of head 6 
mm., of abdomen 7 mm. 

Body stout, hairy on all margins. Antennae very hairy, and also the 
legs, especially the tibiae externally. Head with a ruff of stiff rough 
recurved hairs overspreading the abruptly narrowed bind angles. La- 
bium stout and wide, its hinge reaching posteriorly well between the 
bases of the fore legs. Median lobe prominently angulate in the mid- 
dle ; mental setae alwut thirteen each side, the outermost eight of these 
in a close-set uniform series; some of the smaller inner ones more or 
less out of line. liateral setae eight; hook small, hardly longer than the 
setae, hat much stouter; teeth crenulately recurved and densely spinu- 
lose margined. 

The wing cases reach posteriorly to the tip of the 6th abdominal seg- 
ment. Dorsal hooks on abdominal segments 3-9, beginning with a r 



Google 



806 MICHIGAN 3UBVEY, 1908. 

rudiment on the third and regularly increasing in size backward to the 
ninth, where somewhat surpassing the level of the tip of the lOth seg- 
ment; the; are spinelike ou segments 4-6, but somewhat laterally flat- 
tened and distinctly decurved at the tip on segments 7-9. Lateral spines 
on segments 7 and 8 straight and sharp, directed straight x>osteriorlj, 
at bas0 very slightly angulate with the lateral margins of their seg- 
ments, that of the 9th segment about two-thirds as long as its segment 
and about twice as large as that of the 8th segment. The marffins of 
all the abdominal segments are hairy, especially posteriorly, and there 
is a dense fringe across the ventral spical border of the 9tfa segment. 
The superior appendage is slightly shorter than the inferiors: these are 
triangular and sharp pointed ; the laterals equal the supwior in 
length, and are stout and cylindric, and abruptly pointed. 

The larvae of the four American species of Somatochlora now known 
may be distinguished as follows: — 

1. Lateral setae of the labium eight: lateral spine of the 9th 
abdominal segment more than half as long as its s^ment, and 
the dorsal hook of that segment larger than its predecessors. 
8. forcipata, supposition. 

Ijateral setae of the labium seven : lateral spine of the 9th 
abdominal segment less than half as long as its segment, and the 
dorsal hook of that segment not larger, usually smaller, than some 
of its predecessors 2. 

2. Abdomen less than one-fonrth longer than wide: lateral 
spines of the abdomen, short and broad, eqnilateral triangles. 
8. tenebrosa, supposition 3. 

Abdomen more than one-half longer than broad: Lateral spines 
of the abdomen longer than more acute. 

3. Dorsal hooks of the abdomen of equal size on segments 6-9. 
Dorsal hooks largest on the 7th and Sth s^ments, that of the 

9th segment smaller 8. Nnearts. 

6. Cordulia shurtlefji Bcudder. This fine species is apparently com- 
mon on Isle Boyale, being represented br DTmphs, Kos. 79 A, from Sum- 
ner Lake (III, 5), July 29; No. 120 A. from a rock pool on beach {V, 2) 
on August 14; and Ko. 126 A. from a swamp (V, II) on August 16. 

6. Celithemig eponina Hageu. Two large and four small larvae. 
From a rock pool (Y, 2) on August 14, No. 120 A. 

7. Leucorliinia intacta Hagen. A nnmber of larvae in bad conditioo, 
apparently this species, ou July 29, No. 78 A, 79 A, from the sedge sone 
of Sumner Lake {III, 5). 

8. Enallagma sp.? A number of broken larvae: Sumner Lake (III, 
G), July 29, Hob. 79 A and 126 A; and a rock pool on beach (T, 2) on 
August 14. (No. 120 A) ; also from a swamp (Y, 11) on August 16 
{No. 126 A). 

Plecoptera. 

1. Arsapnia vemalis Newman. A number of specimens of both 
sexes, Nos. 24. 46, and 80 (I, 1), July 6, 11, and 14. Very abundant 
upon the cliff at the shore. 

2. IsopterjjjT cydippa Newman. One specimen from the balsam-birch 
forest (V, 4). on August 14 (G. 236.) ,. , 

,„.CoogIc 



ECOLOGY OP ISLE ROYALB. 307 

Pteronarcys doraata Say. A feir larvae from Otter Eiver, in Baraga 
County Michigan, collected by A. G. Eutliven, No. 30791, U. of M. 
Museum. 

Ephemerida. 

1. He.ptagenia sp.? One pinned female subimago from Isle Royale 
in August, whollr undeterminable; another specimen from Tobin Har- 
bor on Jiilv 'Id. 

2. Sipklunts sp.? probably S. altemata Say. A larva from the 
sedge zone of Sumner Lake (III, 5) Ko. 78 A, on Jaly 29. 



Neuroptera. 

1. Sialig infumata Walker. One larva and one adnlt were taken on 
July 26, bv H. A. Uleason, (No. 160) near the head of Bock Harbor 
(in, 3). 



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MICHIGAN SURVEY, 1908. 



DIPTERA OP THE 1905 UNIVERSITY MUSEUM EXPEDITION TO 
ISLE ROYALE. 

PROFESSOB JAMES S. HINE, OHIO STATE UNIVEESITT. 

Mnch interest always attaches to a collection of insects from north- 
ern regions and when Mr. Cbas. C. Adams wrote and asked me to work 
Tip the Diptera of the 1005 Isle Eojale Expedition, I gladly accepted. 
The collection is a small ope and includes a number of common and 
widely distributed species, but on the other hand it also includes several 
species of special interest. Moat of the specimens were collected by 
Dr. H. A. Qleason, bnt be was aided by Mr, B. F. Savery. 

As the locality is not so far from midway between the East and the 
West the question naturally ariees as to whether the eastern or the 
western species predominate in the makeup of the fauna. This matter 
is the more interesting to me for the reason that lately I have studied a 
collection of Diptera from Hew England and also one from British 
Columbia. After some study of species of Diptera from boreal regions 
I am convinced that there is not the difference in the eastern and 
western faunas in the North that there is in the South. There are a 
number of species in the collection that are common to New England 
and British Columbia, but there are others that so far are not proven 
to have such a wide range, and it is with the latter that most interest 
attaches in the consideration of our question. 

If the Tabanidae are considered we find that three species may he 
said to be exclusively eastern and one exclusively western, while six 
are distributed entirely across the continent. 

In the family Syrphidae are seven species that may be considered ex- 
clusively eastern, and twelve species that reach clear across the conti- 
nent, but not a single one that is exclusively western. 

Id the Stratiomyidae the single species is eastern. So far ae I can 
find Isle Royale is the farthest west the species has been taken. 

In the Bombyliidae one species is western, and the other reaches 
across the continent. 

In the Theivvidae the single species is western, Montreal being the 
farthest east that specimens have been taken. 

In the Asilidae two species are eaatem and one is western. The west- 
ern species however is hardly typical. 

Id the remaining families are several species that are exclusivelv 
eastern and several that reach clear across the continent, but none that 
are exclusively western. To sum up I find four western species and 
more than a dozen eastern, while there are about thirty that occur from 
the Atlantic to the Pacific. Therefore, although there are many species 
common to Isle Eoyale and British Columbia, the following show that 
the general complection of the Isle Royale Dipterous fauna favors that 
of eastern rather than that of western North America. 



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ECOLOGT OF ISLE ROYALE. 309 

Family Culicidae. 

1. Culex pipieni Linn. The rain barrel Moaquito wafl taken July 
11, (Station I, Sub. 1) and August 3, (V, 3). This is the common mos- 
qnito that breeds in receptacles of standing water and small pools 
generally, and widely dlatributed in this country and in Europe, hav- 
ing been described under various names. I hesitated somewhat in giv- 
ing the specimens a specific name for the reason that they were dropped 
into alcohol when they were collected and lost many of their scales be- 
fore they reached me. 

Family Bimuliidae. 

2. SimuUum venustvm Say. Black Fly. Taken July 14 (I. 5) and 
July 28, fill, 5). In Ohio I have found the larvae of this species cling- 
ing to rocks in swift flowing brooks and at the outlet of a small artifl- 
cial lake where the water passed through an iron pipe and dropped ,a 
foot or two on to rubbish and stones. This miuatnre waterfall seemed 
to furnish ideal breeding grounds for the species, for the larvae were 
there in abundance and the adults were flying about in swarms. The 
type locality for the species is along the Ohio River near Cincinnati, 
but it has been identified from a number of states and from Canada. 
It is a matter of interest to know that the species is a member of the 
geous with the well known and destructive Buffalo Gnat. 

Family Stratiomyidae. 

3. Stratiomyia badia Walker. Judging from the large number of 
specimens taken the species must have been common from July 17-31. 
Ail specimens were taken at the Light-houfe clearing (I, 7). At San- 
dusky the species appears in numbers on flowers of White Sweet-clover 
and milkweeds, and specimens are often seen with the pollen-masses of 
the latter plant clinging to their feet. Tbe Isle Boyale specimens are 
typical in coloration but are slightly larger on an average than other 
specimens I have seen. The type locality is Kew Hampshire but its 
range is known to extend over a large part of northeastern North 
America. 

Family Taianidae. 

4. Cltrysops carhonarius Walker, Specimens were taken along the 
MeCai^oe Cove trail and at the head of Rock Harbor (III, 3), July 11- 
14. This is nsually a northern species but has been taken on the east- 
ern coast of the United States as far south as North Carolina. It be- 
longs to the group without an apical spot and is closely related to mitis, 
the species next considered, and from which it is separated by the pres- 
ence of 8 hyaline spot at the base of the fifth posterior cell. In these 
specimens this spot is very small, sometimes making it difScnIt to say to 
which species they really belong. As a usual thing specimens of oar- 
iomiriua are noticeably smaller than specimens of mitia. 

5. Chrysopa mitis Osten Sacken. A number of specimens taken 
along the McCai^oe Cove Trail, July 11, are of this species. As stated 
above the difference between this species and the former is not always ap- 



310 MICHIGAN SURVET. 1908. 

parent, but the specimens ■with the fifth posterior cell uniformly infus- 
, cated at its base are usually decidedly larger than the othere. This is 
quite noticeably in the Isle Itoyale specimens. The type locality for the 
species is the Lake Superior region, therefore these specimens ehould be 
and are very nearly typical. Specimens of carbonarius from farther 
east usually have a distinct hyaline spot at the base of the fifth pos- 
terior cell and therefore are easily known. 

6. Chrysops fp'gidua Osten Sacken. A single specimen taken Au- 
gust 7, by B, F. Savery (V, 3) answers the description of this species 
very well. Here the abdomen is variable in coloration in a series of 
specimens, but the wing markings are nearly constant. I have never 
observed or heard of the species being so abundant and troublesome as 
other niembera of the genus. Type locality Great Slave Lake and other 
northern regions, but it is now known from as far south as Ohio and 
New Jersey. 

Tabanwf afflnU Kirby. Taken July 2, on Mackinaw Island, Michigan. 
A species with hairy eyes, measuring nearly 30 millimeters in length, the 
abdomen is broadly red on the sides and the palpi are long and slen- 
der. The type locality is Boreal America and the species may be ex- 
pected anywhere from Maine to British Columbia. This specimen is 
typical for the species. 

7. Tabanus epiatatua Osten Sacken. Three specimens taken at 
Light-house clearing (1,7) July 8, 11 and 31. Similar to the last in 
coloration and g^eral appearance, but smaller and the palpi are ro- 
bust. Type locality Ilu^on Bay Territory, but now known to be 
widely distributed in northern United States and Canada south to Ohio 
and New Jersey, 

8. Tabanus lasiophthalmua Macquart. A single specimen taken at 
Rock Harbor, in July, by Adams. The eyes are hairy, the abdomen is 
red on the sides and the sise is near that of cpistatua. The cross-veins 
are margined with fuscons making the wings appear spotted, a char- 
acter which ser^'es to separate it from epiatatua and most other north- 
ern species with hairy eyes. Type locality Carolina, but it is dis- 
tributed over northeastern North America south to Georgia and west 
to Illinois. 

9. Tabanus nivoaua Osten Sacken. Several specimens taken at 
Light-house clearing (T, 7) July 11, 2fi and 28; (V, 3) August 7 and 9. 
Tvcngth about 15 millimeters with a row of large white blotches or 
spots on each side of the abdomen; wings clear hyaline; general color 
blackish. Type locality New Jersey, and known from New York and 
Ohio. 

10. Tabanvs sp. Specimens taken at Light-house clearing (I, 7), 
July 18 and 22. This, I take it. is a distinct species but it may be 
one of Walker's obscure forms and I bestitate to name it specifically 
until more material is available. The size is near that of nivi)sus, but 
the general color is reddish, and the white markings on the sides of the 
abdomen are not so conspicuous. There are a number of other char- 
acters which distinguish it. 

11. Tabatiug illotus Osten Sacken. Specimens taken at Light-houae 
clearing (I, 7). July 7, 11 and 25; and fill. 3), July 14. and AuguBt 
5. Eyes hairy, abdomen with a row of white spots on each side, wings 



ECOLOGY OP ISLE ROTALE. 311 

with the front part o( the basal half clouded with fuscous, but other- 
wiae hyaline. The species is near the si?^ of nivosus and appears 
much like that species. Type locality Hudson Bay Region, but at the 
present time known from Alaska and various parts of the British Pos- 
sessions. Specimens from Isle Royale have the white spots on the 
sides of the abdomen lai^r than in some specimens I have observed. 

12. Tabanus insnetus Oaten Sacken. A single specimen taken by 
B. F. Savery August 9, (V, 3). This is the only species known from the 
western states, with the hairy eyes and ocelligerous tubercle absent and 
therefore falls in the genus Atylotua of some authors. Type locality 
Weber Lake, California, ^'ow known from Alaska, British Columbia 
and several of the northwestern states. 

13. Tabanus astiittis Osten Sacken, (?) Several specimens taken at 
Light-house clearing (I, 7) July 2(i and 2S, and (V, 3} August 7 and 
15, by B. F. Savery. These specimens come nearer agreeing with aatutus 
than any other species I know, but there are some points in which they 
do not agree and for that reason I have named them aatutu^ with a 
question. Walker described a number of species from the far north that 
have never been identified since. It is therefore with much interest that 
I receive such collections as the present. Although a number of species 
have a wide uorth and south distribution in boreal regions every col- 
lection from the north is apt to contain something of interest. 

Family BombiUidae. 

14. Anthrax morio Linn. Two specimens taken at Light-house clear- 
ing (I, 7), July 11 and 26. Anthrax seminigra and morio are be- 
lieved to be synonyms. The species is common to Europe and North 
America and is distributed in the latter country from Maine to British 
Columbia. Nearly the basal two-thirds of the wing is black, the re- 
mainder hyaline. The outer margin of the black is irregular and begins 
on the costa near the apex and proceeds obliquely, gradually nearing 
the base. 

15. Anthrax fulmana Say. A single specimen taken July 26, (IIT, 
3). The whole body of this insect is clothed with dense yellow pile, 
the legs are black and the wings hyaline with costal margin and narrow, 
base black. Type locality Pembina, Minnesota, and besides it has been 
taken in New Mexico, , Washington and British Columbia. 

Family Thereddaf. 

16. Tkeretv, frontalis Say. Two specimens taken at Light-house 
clearing (I, 7) July 29, and (V. 3) August 9. Type locality Northwest 
Territory and specimens are at hand from Montreal, Colorado and Brit- 
ish Columbia. The Isle Eoyale specimens are rather larger in size than 
other specimens I have seen but agree closely in coloration with Colo- 
rado examples. Those at hand from British Columbia are slightly more 
brownish, but the thoracic and abdominal markings are of the same 
form and extent in all. 



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312 MICHIQAN SURVBT, 1908. 

Family Asilidae, 

17. Cyrtopogon chryaopogon I^oew. Taken at Light-house clearing 
(I, 7), Julj 6 and 10. Type locality Massachusetts. Known from 
Montreal, Quebec, New Jersey, New York and Florida. This record ei- 
tends the westward range of the Bpecieg considerably. It ia black with 
tlie beard straw-yellow and the bases of all the tibiae red. 

18. DasyllU a»tur Osten Sacken. Taken at Lighthouse clearing (I. 
7), July 7. The two specimens that I include under this name do not 
fully agree with the original description of the apeciea bnt are nearer 
it than to poatioata^ and as Osten Sacbeu indicates certain variationB Id 
his description the specimens are given this name. In typical astur 
from British Columbia the pile on the anterior dorsum of the thorai 
is largely black and that on the tibiae yellow. In the Isle Boyale speci- 
mens the pile on the anterior dorsum of the thorax is all yellow and 
that on the tibiae is black. Osten Sacken observed that specimens ot 
aatur taken at low altitudes bad the pile on the tibiae black, and as the 
size agrees I belieTe it proper to place the Bpecimens in this way. Type 
locality California, but otherwise known from Oregon, Washington and 
British Columbia. 

19. Aailua onntdotiwt ' Williston. Three specimens taken August 5 
(V, 3) . Known from northeastern North America, as far west as Kan- 
sas. The specimens appear to be typical for the species. 

Family Dolichopodidae. 

20. Eydrophorua pftitomftrius Wheeler. A number of specimnia 
taken July 11 (I, 1). I suspect there are plenty of species of this 
family in the Isle Boyale locality but this is the only one Included in 
the collection sent for study. The type locality is Milwaukee County. 
Wisconsin, and it is also recorded from Texas. These specimens are 
typical, agreeing in detail with the original description and figure. 

Family Syrphidae. 

21. Chryaotoxum ventricosum Loew. Specimens taken July 7, at 
Lighthouse clearing (I, 7), The family Syrphidae is a most attractive 
family of flies and the genus to which this species belongs is one of its 
finest groups. The various species are mostly found in northern regions 
or at high altitudes, and are easily recognized by the oblique yellow 
abdominal markings and elongate antennae. This one is tlie largest 
American species of the genus and was first described from specimens 
taken in the District of Columbia. It is now known from New JerseT, 
Canada and Arizona. 

22. Pyrophaena granditarsua Forster. A female specimen taken 
July 28, at Light-house clearing {I, 7). This is the same species that 
formerly passed under the specific name ocymi. It is common to Europe 
and North America and in the latter country is distributed from Ne* 
England to British Columbia. The two sexes are very different in 
appearance and to some extent in structure, and it is from the front 
tarsi of the male that its specific name is derived. 

23. Platyehirus peltatus Meigen. Taken July 26, [II, 1). Common 

i.Cooglc 



ECOLOGY OF ISLE ROYALE. 313 

to Europe and North America. Widely distributed in northern Noi-th 
Aniei'ica from >'ew Knp^lEmd to Itritish Columbia and Alaska. 

24. Platychirus hi/pcrborciis Stsiefxr. Taken, at Light-house clear- 
inp: (I, 7) July 23 and 26. Type locality Greenland but widely dia- 
trihiited in XoVth Amcriea. As with most species of the genus only 
tlie males can be identified satisfactorily by the known characteps. 

25. Melanostnma atimistatum Williston. Specimens taken July 23 
and 26, at Light-house "olearinp; fl. 7). Type locality, state of Wash- 
ington. Known also from the White Mountains and British Columbia. 

26. Syrphm americanns Wiedemann. One specimen taken Jnly 22, 
at Light-house clearing (I, 7). The species is abundant and somewhat 
variable in coloration and is distributed over nearly the whole United 
States and Canada. The larvae have been observed feeding on the grain 
Aphis. 

27. SyrphuB direraipes Macquart. Specimens taken August i and 
7, (V, 31. Type locality Newfoundland. Distributed from New York 
to British Columbia and Alaska, reaching as far south as southern Ohio. 

28. Syiphiis genualia Williston. Taken Jnly 24, at Lighthouse 
clearing (I, 7), July 25, (IT, 1). Type locality New Hampshire and 
recently reported from Beulah, New Mexico. 

20, i'lj/rphvs ribesH Linn. Specimens taken July 24 and 26 at Light- 
house clearing (I. 7). Common to Europe and North .\merica. This 
is one of the most common members of the family and is almost sure to 
l)e included in local lists of Diptera as it is distributed over nearly the 
whole of North .\merica. The larvae are of importance as they feed on 
various species of plant lice. One often sees a colony of plant lice with 
one of the syrphid larvae in the midst of them, and he cannot help be- 
coming intei-ested if he observes for a short time and endeavors to count 
the number of plant lice a larva is able to devour in a given time under 
favorable conditions. 

30. Hphaerophoria cyliitdrica Say, Specimens taken July 25, 26, and 
2S at Light-house clearing (I, 7) and July 25, (II, 1). Tyjie locality 
PennsyJvania. Common over a wide range and included in many local 
lists. The larvae ai-e reported as feeding on the grain Aphis and on that 
account the species is of interest to the economic entomologist. The 
sexes are quite different from one another and one is not likely to asso- 
ciate them on first acquaintance. 

31. ErhtttliH dimidiatus Wiedemann, Si>ecimens taken July 22, 24. 
25 and 26 at Lighthouse clearing (I, 7), August 4 (V, 3) and July 25 
fll, 1). About 40 Bi)ecimens of this siMK-ies were procured indicating 
that it is as common at Isle Koyale as at other places. The larvae of 
the various species of KristuUs are what are known as rat-tailed larvae 
and are found in shallow water in swamjiy places or at the outlet of 
sewers and drains. Each larva is furnished with a posterior appendage . 
which can be lengthened and shortened at will and which contains the 
posterior parts of the tracheal trunks. At the free end of the appendage 
are the two posterior spiracles which ai-e kept at the surface of 
the water. Thus the larva is fitted so it can remain beneath the water 
and yet get the necessary air for carrying on respiration. The adults 
are common around flowers in autunm, sometimes several species visit- 
ing the same patch of asters or goldenrods as the case mav be, .E. dimi- 

41 ' X.OCH^IC 



314 MICHIGAN SURVBY, 1908. 

iiatiia is found all over eastern North America from Florida to Canada 
and west to Kansas. 

;i2. EHstaiis bastardii Macquart, Specimens taken August 4 (V, 3). 
I hare observed tiiis species at midday when the sun was shining, flvlng 
actively over water and hove taken them in numbers at snch times with 
a net. It ia common over the preater part of northeastern North Amer- 
ica, being found as far south as the District of Columbia. 

33. Hclophihia nimilis Loew. Specimens taken July 2G and 28 at 
Light-house clearing (I, 7). The members of this genus are i)eculiar 
in that the eyes are separated in the male as well as the female. This 
species is often taken in early spring from the blossoms of willow and 
other early flowering plants. The type locality is Georgia bnt it appears 
to be more common northward where its range extends from the Atlantic 
to the Pacific. 

34. Afallota cimbicifonnis Fallen. Specimens taken July 23 and 26 
at Lighthouse clearing (I, 7). Common to Europe and North America 
and widely distributed over the eastern part of the latter country. The 
species has a'resemblance to certain species of EristaUs, but the greatly 
thickened hind femora are distinctive. 

35. Xylota curvipes Loew. One specimen taken July 26, at Light- 
honse clearing (I, 7). The genus Xylota contains upwards of 40 North 
American species which in the main are reasonably easy to separate, 
and for that reason it is an ati:ractive group. Various species are often 
observed resting on toga in damp places or that lie across small streams. 
In many the abdomen is distinctly elongated and the hind femora are 
swollen. X. curvipes is common to Europe and North America, being 
most often taken in northern latitudes. 

3fi. Xylota fraudulosa Loew. One specimen taken August 12 (V, 3). 
Type locality Illinois, but known in northern North America from New 
England to Washington reaching south to Ohio and Nebraska, 

37. Xylota pi(jra Fabr. One specimen taken July 22 at Lighthouse 
clearing (I, 7). Common to Europe and North America and generally 
distributed over the United States and Canada. The adult has been 
reared from a larva taken from under the bark of a pine tree. 

38. Temnostoma aequalia Loew. Specimens taken July 17, 22, 23, 24, 
26 at Lighthouse clearing (I, 7) and July 25 (II, 1). This fly has some- 
what the appearance of the common bald-faced hornet and one usually 
thinks the second time before taking it in his hand. In Ohio various 
species of the genus are to be found around rotten logs where the females 
oviposit and the larvae pass their lives as such. Type locality, English 
River, Hudson Bay Kegion. Otherwise known from New England and 
Colorado. The Isle Royale specimens vary slightly in abdominal and 
thoracic markings but on the whole agree very well with the original 
description. 

39. Temnostoma hombylani Fabr. One specimen taken July 17, at 
Light-house clearing (I. 7). Common to Europe and North America and 
widely distributed in the latter country, having been taken as far south 
as southern Ohio. 

Family Tachinidae. 

40. Pelcteria robiista Wiedemann. One specimen taken July 26 at 
Light-house clearing (I, 7). This species is reported as occuring from 



ECOLOGY OF ISLE ROYALE. 315 

Ai^ntina to Canada and from the Atlantic to the Pacific, Wiedemann's 
types were taken in South America. The Isle Royale specimen has 
less red at the tip of the abdomen than most Ohio specimens. 

The family Tachinidae contains a large number of species and nearlv 
all of them are of more or less interest to the economic entomologist on 
account of their parasitic habits. Many injurious insects have one or 
more Tachinid parasites which aid in holding them in check. 

41. Echi iiomi/ia algetis Wiedemann. Two specimens taken July 14 
(III, 3) and July 26 at Lighthouse clearing (I, 7). The types were 
taken in North America, but the exact locality is not given. Recent 
writers have reported the species from many points in Mexico and north- 
ward. It is said to be parasitic on the larvae of the moth, Hadena ligni- 
color Gnenfe. 

Family Sarcophagidac. 

42. Sarcophaga sarraceniae Riley. Flesh Fly. A specimen taken 
July 11 at Light-house clearing (I, 7). This ia our common flesh fly, 
and is an important scavenger. Type locality MiBsouri. The species of 
Sarcophaga are not well understood in America and it may be that this 
is a synonym. However the name is included in many local lists of 
Diptera which indicates a wide range for the species. 

43. Lucilia caeaar Linn. Carrion Flv. Four specimens taken July 
2.5 (II, 1), July 26 at Light-honse clearing (I, 7) and August 5' (V, 3). 
Known from Europe and America. A very common carrion fly every- 
where. Along the shores of the Great Lakes its larvae feed lately 
upon the carcases of fishes cast on the beach by the waves. 

44. Gallophora virtdesccns Desv. Blow Fly. Three specimens taken 
July 25 (II, 3) and August 7 (V, 3). This is one of the common blow 
flies and is widely distributed in Europe and America. 

45. Cynomyia cadaverina Desv, Two specimens taken July 8 (I, 1). 
Carolina is the type locality but the species is found in most localities 
in the United states and Canada. 

46. Phormia terraenovae Desv. A specimen taken August 4 (V, 3). 
Type locality Newfoundland. Generally distributed over North Amer- 
ica, especially northward. 

Family Muscidae. 

47. Musca domestica Linn. House Fly. A specimen taken August 
7 (V, 3). This species needs no particular comment here. It is found 
In nearly all parts of the world and has lately been proven to be con- 
nected with the transmission of typhoid fever. 

Family Anthomyidac. 

48. Byetodegia aerva Meigen. Five specimens taken July 11, 23 and 
24 at Light-hpuse clearing (I, 7) and August 4 (V, 3). This European 
species has been reported for America, but its distribution is not well 

^understood. I have compared these specimens carefully with Schiner's 
description and find that they agree well, but as the group is very rich 
in species their determination is not alwaj's an easy matter. 



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MICHIGAN SURVEY, 1908. 



Family Scionij/zidae. 



49. Tetanoccra plebeia hoew. A specinieo taken July 26 at Light- 
liOTise clearing; (I. 7). Type locality Jliddle States. Specimens are at 
hand from Britisli Columbia and other localities. The niembei'S of this 
genus are often common in marshy and damp places. 

60. Sepcdon piisUhis Tjoew. Two specimens taken in a swamp in Cass- 
andra and Sedge Zone (V, 2), by Max M. Peet. Type locality Middle 
States. Known from Ohio, New Jersey and White Mountains, New 
Hampshire. The various sj}ecies of this genus are usually found in 
swamps where they may be observed flying over water. 



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ECOLOGY OF ISLE ROYALB. 



ANNOTATED LIST OF ISLE ROYALE HYMENOI'TERA. 

B. G. TITUS, KNTOMOI^OIST, UTAH AGRICLLTUEAL EXPERIMENT STATION. 

The determinations id this gronp were made in Washington at the 
U. K. National MnKPuni, Mr. J. C Crawford and nijself working over 
most of the material together. Mr. Tlieodore Tergande, Bureau of Ento- 
mology', determined the two species of anta represented in the collertion. 
The general collection of anta are reported upon elsewhere by Dr. W. 
M. Wheeler. Mr. W. F. Fiske. at that time in the Foi-est Insect section 
of the Bureau of Entomology, very liindly determined the Ichnenmons 
and Siricids, both of which groups he had been worliing with for sereral 
years. At the time the determinations were made the writer had no 
expectation of wi-iting up the notes or he would have doubtless been able 
to add considerable to their value by making further notes on the speci- 
mens retained at the National Museum. 

Bombiis tcrricola Kby. (Det. Crawford.) Three on flowers of Opu- 
laster (II, 1) (14) : about camp at Rock Harbor several specimens (133, 
lfi6 three, 179, 191) and one Bi>ecimen (222) around camp at Siskowit 
Bay. 

Geographic Range: Originally described from Canada; Kirby, 1S37. 
Also reported by Provancher from region around Montreal and by 
other authors from various localities in Northern United States east of 
the Rocky mountains and as far south as Colorado. I have seen speci- 
mens from Massachusetts, New York, Illinois, Kansas and Colorado. 

Bombus conaimilis Cress. (Det. Crawford.) Two were taken around 
camp at Rock Harbor (36, 45). 

Geographic Range : I>esepibed from New York, Cresson 1864, p. 41 
and reported by Packard, IHQi, p. 112. 

BomliHS sp. One flying over beach at end of Conglomerate bay (31) ; 
one on flowers of D. trifida on a jack pine ridge (23) ; and two about 
camp at Rock Harbor (A. 5, 98). These ail seem to represent one species 
but neither Mr. Crawford or myself care to name it in the present un- 
settled condition of the group. 

Psith'ijru« latitarmis Morrill (Det. Crawford.) Two around camp 
clearing at Rock Harbor (A 36, 45), 

Geographic range : Desc. from Montana by Morril 1903, p. 224. 

Megach'Udac. 

Monumctha albifrotis Kby. (Det. Titns.) One specimen flying over 
■ ridges near Conglomerate bay, (68). 

Geographic range: Desc. by Kirby 1837, p. 270 from "Lat. C5°"; 
again by Cresson 1864, p. 387, 388, as three separate new si>ecies from 
Colorado, Pikes Peak, and Slave Lake. It probably occurs over all the 
region from the Mackenzie river and Upper Hudson bay to the lower 



318 MICHIGAN SURVEY, 1908. 

Rocky Mts. areaa io New Mexico and westward to the Pacific. (Titns 
1906, p. 158, Cockereil 1906) (1 aod 2). Nothing is known of its 
breeding habits. I have specimenR ffom eastern Canada and New Eng- 
land but have seen none from south of New York along the Atlantic 
region. There are mites on the Isle Koyale specimens. 

Xanthosarus mclanophwa Smith. (i)et. Titus). Taken on jack pine 
ridge (108) ; about camp at Rocfc Harbor (133, 160 eleven) and on flow- 
ers of Vampatiula rotuiidifoHa in clearing at Siskowit Bay (148, 202). 

Geographic range: Described from British America, Smith 1853, p. 
91 and known to occur throughout the region of southern Canada, New 
England, New York, and in northern United States to the Pacific coast 
and in British Columbia. This and the following species are leaf-cut- 
ters working esi)ecial!y on the leaves of Rosa spp., the pieces clipped 
out ai-e used in lining their nests which are usually made in old logs 
or dead trees, the female often utilizing the abandoned boring of some 
other insect. The little rolls are often found when splitting I(^b or 
wood in the fall or winter. 

Xmitkosarvs latimanus Say. (Det. Titus). Very frequently taken 
about the camp clearings at Rock Harbor and Sit^kowit Bay (S8, 49, 6S, 
133, 137, 153, 179, 231) ; also at sand beaches at head of Conglomerate 
bay {^1} ; on jack pine ridges (68, 1-8) ; near Neutson's resort (121) ; on 
flowers of OpuJastcr (148); and on flowers of Campanula rotiindifolui 
(202). 

Geographic range: Described from "Arkansas" by Say 1823, p. 81. 
which may mean any where from Missouri to Colorado, It is one of the 
most common species in the United States and Canada, occurring from 
coast to coast and from the Gulf northward. 

Anthemoia sp. near infragilis Creseon (Det. Titus). This specimen 
was taken around camp at Rock Harbor (86). While it bears a close 
resemblance to A. infragilis there are sufficient differences to make it 
a good species and probably new. A. infragilis was described from 
New York and probably occurs in the Isle Royale region, since I have 
seen specimens from Canada (Titus 1906, p. 152). 

Stelidae. 

Ctrlioxys moesta Cresson. (Det. Titus). One on flowers of Cam- 
pnntil^i rotuiidifolia in clearing at Siskowit Bay (202) (V, 2), 

Geographic range : Described from Connecticut, Cresson, 1864, p. 
403; reported by Provancher, 1882, p. 241, 1S83, p. 725 as tristit, from 
Canada. Occui-s westward to Colorado, New Mexico and probably 
Utah. 

Chclynia nitida Cresson. (Det. Titus). One specimen about camp at 
Rock Harbor (26). 

Geographic Range : Desc. from New York by Cresson, 1878, as 
a Htelia and from Canada by Provancher, 1888, p. 322 as Chelnia 
labiata and in Panurgidae. Ashmead, 1896, p. 283, erected the genoB 
MelanostcHs for his species betkcU, which is congeneric with nitida 
(Titus, 1906, p. 161). 



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ECOLOGY OF ISLE ROYALE. 



Andrenidac. 



ffalirtiia IcroiixU Lepeletior. (Det. Crawford). One specimen about 
camp at Kock Harbor (133). 

Ot'Ograpliic range: Described by J^epeletier 1S41, p. 272 from "Am. 
Boreal.*' Occurs at least as far west as Illinois, Robertson, ISitS, p. 146. 

ffalictiis irrsaiiji Lo^■Bl!. (I)et. Crawford). Five specimens on flowers 
of I'hifsocarjitis in Bnnsom clearing (II. 1), (14S). 

Oeograpliio range: Descrilied from Maine by Lovell. 

Proaopidae. 

Pyoso-pin hifKtilis Smith. (Det. Crawford and Titus). Ctae about clear- 
ing at Rock Harbor (lllti), 

Oeograt.liic range: l(e8cril)cd from Hudson's Bay by Smith, 1853, 
p. ^i:^ and occurs from the upper Atlantic coast to at least the moun- 
tains of Colorado. 

Mxjsopis RjR'cies. (Det. Crawford). On flowers of Opulaater (V, 2) 
behind camp at H^iskowit Bay |203) ; in camp clearing at same plac*^ 
1 212) and two unmarked epecimena. There may be two species involved 
here but we were unable to specifically determine them. The species of 
the genus breed in stems of small plants. 

Crabronidae. 

Crabro sittffulans Smith. (Det. Crawford). One specimen on sand 
beach on a jack pine ridge near Conglomerate bay. (lOS). 

(ieograpliic nm'ge: '"Canada and United States." 

t^'olcnius sp. (Det. Titus and Crawford). One specimen about canip 
at Bock Harbor. (170). 

Pcmphrcdonidae. 

Diodontiis atUiiiisi n. sp. Titus (Det. Titus and Crawford). On sand 
beach with Ammophila at end of Conglomerate bay (31). Xotes on 
AiiimOphUa will apply to this s{)ecies. 

2 Leng h 7.1 mm'. Black, with scattered silvery pubescence, espec- 
ially abundant on fai-e; olyitens projecting, with two sharp teeth wide 
at the base, tips of mandibles rnJdish, palpi brown; tegulae brown, 
yellow in front; wings slightly infuscated; tibiae and tarsi reddish 
brown. 

Sphrgidae. 

Ammophila sp. (Det, T, & C). Three on sand beaches at head of 
Conglomerate bay (31) ; one near Tonkin bay (41) ; and one about camp 
at Bock Harbor (13.3). The normal habitat of this species of sand- 
wasp Is on the sand beaches. They fly rapidly about at a height of 
3-iri centimeters over the sand or gravel, alighting only ■ on the 
«and. This gi-oop all store their nests with caterpillars, the holes being 
nsHally in quite hard ground. They are very skillful in removing <»!• 
covering up all traces of the place where they have worked, often going 
to much more lal)or than the occasion would seem to require. Feckham 
and Peckham, 1S98, pp. 6-32, have a very interesting chapter on this 
subject. 



320 MICHIGAN SURVEY, 1908. 

PsamtiiopliiUt sp. (I)et. T. & C.)- One apecinieu from rock clearing 
near outlet of WisUowit lake {V, 9) (215). 

Ccropalidae. 

EntyiHis amevk-uniiH Pal-Beaiiv. (I)et. T. & C). One Bpecimen (2^5) 
captiireii with a Hi>ecimen of Lycosn kochi Keys, (Det. Banks). The 
waap was backing over the gpounrt. dragging the spider, at brief in- 
tervals it drop])ed its prey and ran rapidly back and fortli looking for 
its hole. It apparently had a general idea of tlie direction in which 
llie nest lay hut had to walk right to it in order to be certain of its 
location. Even a couple of uentiuieters was not close enough. Having 
found the nest the wasp searched in the same way for the spider and 
was backing away with it in a direct line for the hole when both were 
cajitured. 

fieograpliic range: Delia Torre gives "United States," It was de- 
scribed by Palisot-Beauvois, 1811, p. 117. Peckham and Petkham, li^DS, 
pp. 125-l(i6, describe the labors of several siwcies belonging to this 
gronp and call them '"The Spider Itavishers." 

Vcspidac. 

Ycspa (Uaholico Saussure, (Det. T. & C). One specimen about camp 
cleai-ing at Rock Harbor {16fi). Described by Sanssure, 1853, p. 
138. Occurs fairly common throughout the eastern United States and 
Canada. There have l>een many errors in determining si>eties of This 
group so that one can hardly state the distribution of any species. 

Eiiwinidae. 

Ancintroceniit ciiprn Sauss. (Det. T. & C). One taken about camp 
at Rock Harbor (133). 

Geographic range: Sanssure 1857, p. 273. Known to occur in 
nortliern United States and eastern Canada. 

Atu-istrocfTiia pertinax Sauss. (Det. T, & C). Two on flowers of 
Hi'rarJriiii) iintoiiiiii in camp clearing at Rock Harbor (105). Tliis 
si)ccies may l>e a true O'Ii/iktus. All of this group are predaceous and 
these probably store their uests with caterpillars. Their habits are 
rnried, some boring in one plant or substance and others using old 
burrows. 

Geographic range: Snusanre. 185G, ]». 216. Known from northern 
and eastern T'nited States and Canada. 

Enmnivs sp. (Det. T. & C). One in Cladonia clearing behind camp 
at Siskowit Ray |2II1|. Tliis genns are the so-called "jug-makers" or 
"mason-wasps" and store their nests with smalt caterpillars. 

Formicidoc. 

Formica sp. (Det. Pcrgande). One specimen in camp clearing at 
Siskowit Bay (23U. 

Ciiinpnnotiis piiii)s>/h-;iinnin Degeer, (Det. Pergaude). One from 
Station IV, 1 |13()), another found running over smmith sand bleach at 
head of Conglnmenite bay (311; one at camp at Rock harbor tl"-** 



ECOLOGY OP ISLE ROYALB. 321 

and others at Siskowit Bay camp (212, 222 eleven, and 232). A very 
common Bi>e<:le8 throughout eastern United States and Canada. 

CJtrifsididae. 

ChrffSOgOna vrticolis Piitton. (Pet. T. & C). One specimen about 
cauip clearing Siskowit Bay (2.10). ■* 

Geograi)hic ran^e: Described by Patton ISJO, p. 07, and aftei-ward 
noted by Aaron 1.SS5, p. 226, from California and Provancher 1887, p. 
215, fi-om Canada. 

Emniidac. 

Gaxteruption incerttis Cresson. (Det. Fiske). One at camp at Sisk- 
owit Bay (239). Mr. Fiske placed this in Focntis which Ashmead makes 
a synonyn of Ganternptioti. 

(icographic range; "Cauadii, Colorado" Cresson. All of this gennH 
that have been bied were foiind jKirasitic on some apeoiea of wasp or bee. 

GasteruptioH tarsitorius Say. (Det. Fiske). One taken at Hiskowit 
hay camp clearing (A. 152). 

Geogi-aphio range: Eastern United States and "Canada." 

Aiilficus ruptareus Cresson, (Det. Fiske). One at Eock Harbor 
(133) ; and others at Siskowit Bay (A. 152, 212 five, 231 two, 239). 

(Jeographic range: "Canada, Colorado" Cresson. 

Some of the 8;>ecies in this gi'oup are parasitic on Cerambycid larvae. 

Ichneumonidac. 

Pmnenis Hp. (Det. Fiske). One about camp clearing Siskowit Bay 
(231). Probably a parasite on some wood-boring coleoptera. 

Fimpla conquisitor Sav. (Det. Fiske). One about camp clearing Rock 
Harbor (179). 

Oeographic range: ''(Canada; TT. S." Ci-esson. 

Doubtless parasitic on a T^pidopterous lar\'a. 

Ephialtcx gigaa Walsh. (Det. Fiske). One about camp at Siskowit 
Bay (153). 

Geographic range: Descrilied from Illinois, also reported from Can- 
ada. 

Rhi/ssa alboiiutciilata Cresson (Det. Fiske). Taken at Rock Harbor 
clearing (106), and at Siskowit Bay camp clearing (A. 152. 153, 212 
six. 222. 231 two. 239). 

Geographic range : "Canada ; U. S," Cresson, 

Species in this genus have been repeatedly bred from the larva of 

xyloi)hagous saw-flies such as Trrocerns. The ovipositors in some species 

, attain the length of six or more inches, with these they are able to reach 

the larva of the host and lay their eggs even when the unsusi>ecting 

victim is living far inside the trunk of a tree. 

Bra<conidac. 

ApantrlcM sp. (Det. C. & T.). One alcoholic specimen without data. 
It would be hnpossible to even superficially determine this from the one 
specimen. ' ..^ 



322 MICHIGAN SURVEY, 1908. 

Oymnoscclus pedulis CrcBBon (IVt Fiske). All taken arouod camp 
clearing Siskowit Bay, {A. 152. 212 six, 222 three, 231, 239 six). 

(jeogra()liic range: Canada, Eastern U- S, 

Parasitic upon some nood-boring Coleoi)tera. 

Mclanobrocon sp. {I>et. Fiske). Two taken at camp clearing Siskowit 
Bay (212). 

SHcUlac. 

Vroceni€ fiavipcimia Kirby, (Det. Fieke), All sitecimens taken in 
camp clearing, or near it, at Siskowit Bay (A. 152 four, 153, ftl, O, 212 
two, 209, 22y, 231 three, 239 seven). Usually flying with a moderate 
velocity about the cJenring, keeping at average height of 2-3 feel 
and not alighting. A few were found climbing up and down trunks of 
balsam trees in which they deposit their eggs. 

Ge<^aphic range: Vancouver's Island, Kirby 1882, p. 380. Occurs 
across the continent. 

Urocrrus fiamcomis Fabricius. (Det. Fiske). All taken in or near 
Siskowit Bay camp clearing and not differentiated at the time from the 
previous species (195, 201. 212, 241 two). 

Geographic range: British America, Fabricius 1781, p. 418; appears 
to be a more northern species in its range than V. ftavipennis tliough 
they are often taken in same localities. 

Tenthridinidae. 

Teiithredo melUna Nort. (Det. MacOillivray). One at Bock Harbor 
(136), others on flowers of Opula^tcr in camp clearing .Biskowit Bay 
(203) and around camp, same place (208). 

(ieographic range: Canada and U. S, 

CimJiicidae. 

Cimbcx amerU-ana. Leach. (Det. T. & C). Taken in rock clearing near 
water's edge on north side Conglomerate bay (106) ; also one lar\-a in 
alcohol which may belong to this or the next species. 

(ieographic range: The earliest record for this species is Alibot 
1792. plate 01, under the name of femoi-ata Linne. Leach described it 
in 1817, p. 33, and since that time many autliors have written upon 
the species which is rather a common insect throughout most of the 
regions in North America where willows are found. The larva of this 
and probably the following species feed on willow leaves. 

Cimhrx violacea Lepeletier. (Det. T. & C). Julv 13. eta. I, 2 
(71 and 55). 

Geographic range : Described by lepeletier 1823, p. 27, from North 
America and reported by Kirby from British America ; also occurs in 
northern United States. 



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ECOLOGY OF ISLE ROTALE. 



BIBLIOGRAPHY. 



AaroD, 1885, Tr. Am. Eat. Soc, xii. 
Abbot, 1792, Drawings Insects Georgia, xii. 

Cockereil, 190G, (1) The Bees of New Mexico, Tr. Am. Ent Soc.,xxxii. 
(2) The Bees of FloriesaDt, Am. Miis. Nat. Hist., 
V. xxii, art. xxv. 
Cresson, 18«3, Proc. Ent. Soc. Phila., ii. 

1864, Proc. Ent. Soc. Phila., vol. ii and iii. 
1878, Tr. Am. Ent. Soo., vii. 
De Geer, ni'i, Mem. Sen-. Ins., iii. 
Pabricius, 1781, Spec. Insecta, i. 
Kirby, ISiST,, Fauna Boreal America, iv. 

1882, List Hvm. Brit. Mnseum, i. 
T-each, 1817, Zool. Miec, III. 
I^peletier, 1823, MonograjA, Tenthredin. 

1841, Hist. Nat. Ins. Hymenop., ii. 
Morrill, 1903: Canad. Entomologist. 
Norton, 1860, Boston Jour. Nat. Hist, VII. 

Paliaot Beauvois, 1811, Ins. rec. en Africa and America, Hymenop. 
Patton, 1873, Canadian Entomologist, xi. 
Packard, 1864, Proc. Epsex Inst., iv. 
Peckham & Perkham, 1898, Solitary Wasp. 
Provaocher, 18S2, Nat. Cadadien xiii. 

1883, Fauna Ent. Canad. Hym. 
1887-8, Addit. faun. Can. Hym. 
Robertson, 1893, Tr. Am. Ent. Soc, xx. 
Say, 1823, Western Quarterly Reporter Cin. ii. 
Saussure, 1856, Etud, fani. Vespidae, iii. 

1857, Rev. and Mag. Zool. (2), ix. 
Titus, 1906, Proc. Ent. Soc. Washington, vii. 



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ECOLOGY OP I8LB ROTALB. 



THE AKTS OF ISLE ROYALE, MICHIGAN. 

BY DK. WILLIAM MORTON WHEELBR. 

Hairard University. 
Subfamily Myrmicinae. 

1. Myrmica hiviinoilis Eniei'y var. cana^ieTisig Wheeler. Sev- 
eral workers from a single colony: CI (I, 2) H. A. Gleason. "Found 
on the dry rock ridgea under the mats of bearberry and also excavating 
nests in the crevices of rocks to a depth of some 8 cm." This is the com- 
mon variety of the subepecira brevinodis at higher elevations in Canada 
and the Eastern States. 

2. Leptothorax acercorum canadenitis Provancher, Workers from 
three colonies: 63 (I, 2), (I, 1), 77 (I. 2), H. A. G. "Abundant in Cto- 
donia clearings and on rock ridges, running about on the surface and 
through the thin deposits of soil. The specimens of No. 73 were from the 
pock pools on the shore just south of Tonkin Bay." This ant, like the pre- 
ceding, extends its range info the Northern and Eastern States, but it is 
by no means common. It is abundant, however, at higher elevations 
(8000-9000 ft.) in the Rocky Mountains and at lower elevations in Nova 
Scotia. 

Subfamily Dolochodcrinoe. 

3. Tapinoma sessile Say. Workers from a single colony: 132 (V, 
2) C. C Adams, "under Oladonia." This is the only I >olichodfirine ant 
which ascends to high latitudes and elevations. I have found it nesting 
under stones at altitudes of over 10,000 ft. near Cripple Creek, Colorado, 
and it is common in the Canadian zone throughout the Eocky Mountains. 
In the Northeastera States it descends to sea-level. 

Subfamily Cnmponotiiiae. 

4. Lasiiis niger L. var. n€onig<'r Emerv. Workers from five col- 
onies: 20 (I. 5) C. C. A., and 75 (I. 1), 70 (I, 5), 82 (I, 5), 83 (I, 5), 
H. A. G., "Abundant on the rock ridges and jack pine ridges (I, 2, 5). 
The nest is always constructed beneath or at the side of a flat or angular 
stone, at a depth of one decimeter or more, A complicated system of 
roomy galleries is excavated with passages l.D-2.5 cm. high by 2-5 cm. 
broad. This ant was seen to capture and kill a beetle. No. 75 H. A. G. 
is material from the rock pools." (Gleason). 

There are in North America three distinct varieties of the circum- 
boreal L, niger, viz., var. tieoniger Emery, sitkdcnsis Pergande and ameri- 
canus Emery. The first and second have the legs and antennae of the 
workers and females covered with suberect hairs, and the hairs on the 
body are also conspicuously abundant. L. nrnniger is small and black, 
sitkaensis much larger and of a lighter brown or yellowish color. ^^JL^ 



32Q MICHIGAN SURVET, 1908. 

americanus is small, like neonlger, but brown aod has few erect bairs 
on the body and none on the legs and scapes. It is clowly related fo 
the palearctie variety alicnxis Porster. and like this form inhabits warm 
and rather dry localities. It is the eonimou form of nigr.r throughout 
the Northern States. L. gitkdciiais occui's in Alaeka, Jlova Scotia and in 
the damp alpine meadows of the Rocky Mountains at altitudes between 
8000 and 9000 ft. L. nconigrr occurs in dryer situations at somewhat 
lower elevations and is occasionally found even near sea-level in isolated 
colonies in our northern woods. Varieties (hybrids?) intermediate be- 
tween neoniger and americanus also occur in thece same localities. 

5. Formica aanguinca attcrva Forel. Workers from two colonies: 78 
(I, 2), 72 (I, 2) H. A. G. "This is one of the commonest species on the 
rock ridges, but constructs its nest either in or under decaying wood. 
On the ridge north of the light house, a nest (72) was made under a 
rather small rotten stick, and the soil beneath was composed mainly of 
finely comminuted fragments of the wood. The second colony (78) had 
constructed a nest in the interior of a large decaying li^," (Gleason). 
This subspecies has been taken hitherto only at Toronto (Porel), on the 
summit of Mt. Washington (Mrs. A. T. Slosson), among the Litchfield 
Hills of Connecticut (Wheeler) and in Casco Bay, Maine (Wheeler). 
It is a decidedly boreal form, approaching the typical palearctie 
sanguinea in size and coloration. There were no slaves accompanying 
the specimens from Isle Boyale, a fact which tends to confirm the con- 
clusions of Porel and myself that this subspecies usually lives in pure 
colonies. 

6. Formica adamsi sp. nov. Worker. Length S.5-5mm. Allied to 
F. rvfa Ij. Head, including the mandibles, nearly as broad as long 
even in the smallest individuals, with straight posterior border, rounded 
posterior corners, and slightly but distinctly convex sides. Eyes large. 
Mandibles 7-8 toothed. Clypeus prominently carinate, with broadly 
rounded anterior border, not produced in the middle. Palpi of moderate 
length. Antennae slender, scapes nearly straight at the base, funicular 
joints all distinctly longer than broad, the basal somewhat more slender 
and longer than the apical joints. Pro- and mesonotum moderately 
rounded, convex, the latter eliptical and nearly twice as long as broad, 
the former a little broader than long. Kpinotiim with subequal base 
and declivity, the former slightly convex, the latter flattened or slightly 
concave; the two surfaces in profile passing into each other through a 
rounded angle. Petiole more than half as broad as the epinotum, in 
profile with convex anterior and flattened posterior surface and sharp 
upper border; seen from behind the border is rounded and but feebly 
or not at all produced upward in the middle. Gaster and legs of the 
usual shape. 

Opaque throughout ; only the mandibles, frontal area and sides of the 
clypeus faintly shining or glossy. Mandibles finely and densely striated. 
Surface of body densely and indistinctly shagreened. 

Hairs and pubescens pale yellow; the latter covering the whole body 
and appendages, not conspicuous except on the gaster, but even on 
this r^on not sufHciently dense to conceal the surface sculpture. Hairs 
short, sparse and obtuse, in several rows on the gastric segments; on 
the thorax confined to the upper portions of the pro- and mesonotum. 
on the head to the clypeus, front and vertex. The hairs on the mandibles 



ECOLOGY OP ISLE ROYALE. 327 

are appreesed and pointed, ob the palpi short bat numerous and conapic- 
nous. I-egs naked except for a series of pointed bristles on the flexor 
surfaces of the tibiae and tarsi and a few blunt hairs on the anterior 
surfaces of the fore coxae. 

Sordid brownish red, the smaller sppcimens somewhat more yellowish 
red. Gaster dark brown, except a large spot on the base of the first seg- 
ment and the anal region, which are reddish yellow. A large spot on 
the pronotuni, one on the mesonotum, much of the posterior portion of 
the head, the distal halves of the antennal funicnli and in many speci- 
mens also the coxae and femora, dark brown or blackish. These dark 
markings are present in the largest as well as in the smallest workers. 
Teeth of mandibles black. 

Described from numerons specimens taken from a simple colony: 115 
fl, 6) n. A. G. A dozen workers taken by myself on Pikes Peak, Col- 
orado, near timber line, at an altitude of lO.EiOO to 11,000 ft. differ from 
the Isle Hoyale specimens only in having the frontal area smooth and 
shining, in having the middle of the petiolar border produced upward 
as s distinct, blunt jxiint, and in the less extensive infnscation of the 
bead, pro- and mesonotnm. Th«(e specimens may be regarded as repre- 
senting a distinct variety, alpina var. nov. Both this and the typical 
adamsi may be distinguished from onr other North American forms of 
the rufa group by their small size, opaque surface and peculiar color- 
ing and pilosity. The following collector's note on the Isle Royale 
specimens adds some elhological characters which are not seen in the 
other small forms of the rufa group known to me: "The nests of this 
ant are one of thij most conspicuous features of the drier tamarack 
Kwanips. They aro rounded-conical in shape. %Q dcm. high or even 
lai^er and with a diameler at the base about equalling the height. They 
are composed within of Sphagnum, but as would be expected with such 
material, without any definite system of galleries. The outer surface 
is thickly covered with leaves of Cassandra, probably to prevent loss 
cf moisture by evaporation from the interior. They are frequently 
placed near or under a bush of the Cansandra, but the same covering is 
used if no Cassandra is near," (H. A. Gleason). 

7. Ff>rmica rufa obsctiriventi-is Mavr, Workers from six colonies: 
4« (I. 1), 47 (I, 1), 63 {I, 2), 76 (I, 2), 114 {I. 6), 14 (112) H. A. G. 
"This subspecies occurs on the rock beaches (I, 1, 46, 47) where it 
forages about on the surface and in crevices but is more abundant on 
the jack pine ridges (I, 5, 63) and on the rock clearings (I, 2, 76)." 

I recently described this subspecies as F. dryas, but an examination 
during the past summer of some of Mayr'e types in Pi-ofessor Forel's 
collection, shows that in so doing I created a synonym. Mayr's original 
description based on specimens from Connnecticut is entirely inade- 
quate, and the list of localities which he later cited for obsniriventris 
shows that he lumped together a number of different forms belonging 
to the rufa group The name ohscurivcntris, therefore, should be re- 
stricted to the form having the characters of my F. dryas. This ant is 
rare in the Eastern and Northern States and evidently belongs to the 
boreal fauna. 

8, Formica fusca L. var. subscricca Say. Workers from 11 colonies: 
23 (I. 5), 102 (V. 2), 131 (V, 2), C. C.'a., and 80 (I, 5), 81 (I, 5). 
100 (I. 5). 102 (I, B), 223 (V, 3), 224 (V, 3), 226 (V, 3). 227 (V, 3) 



328 MICHIGAN SURVEY. 1908. 

II. A. G. Also specimens from a. single rolony ou Mackiimt^ Inland 
(3. H. A. G.)- "A common ant ou the jack piue ridges (T, 5, 8(1, 81. 
]00, 102), It constructs its nests under rocks in moist soil (100) and 
was observed to capture beetle larvae (103). Tlie specimens collected 
in the rock-clearings at Piakowit Bay (V, 3, 223, 224, 220, 227i. con- 
strupted circular, flat-topped craters 6 dcm. in diameter, covered with 
debris of balsam and spruce needles and frequently with growinft plants 
on them." This is the common form of the circumboreal F. fusm- 
throughout Canada and the northern states. At higher altitudes on 
the Bocky Mountains it passes into the more silvery red-legfred var- 
arfjentata Wheeler, a form which also occurs even near sea level but 
very sporadically in the Atlantic States, 

9. Formica ftisca L. var. neorufibarhifi Emery. A few workers from 
two colonies; 15 (I, 1) and 20 (I, 1) H. A. G. in vials with specimens 
of Lasius neoniger and Cnmiponotus whympcri. Of the numerous 
varieties of F. fvsca this is the most boreal, being known only from 
Alaska and British America as far east as Labrador and Nova Scotia, 
and from higher altitudes in the Rocky Mountains (9,000 to 12,500 
feet). It forms rather small colonies under stones and logs in moist 
or shady places. 

10. Camponotus hercuUaitue L. var. ichymperi Forel. Workers 
from 10 colonies, with larvae and pupae; 15 (I, 1), 18 (I. 1), 22 
(I, 1). 30 fl. 1). 63 (I, 2), 140 (I, 3) H. A. G. and 105 (Y, 2), 126 
(V, 11), 148 (TIT, '04), 149 (III, '04) C. C. A. "Although an abund- 
ant species on the rock and gravel beaches (15, 18, 22 H. A. G.) where 
it forages for dead insects, its actual home appears to be the ridges. 
On the dry ridges it occurs singly, usually in soil under stones (62, 
H. A. G.), It was also collected |140 n. A. G.) in the dense balsam 
fir woods, where it forages over the surface. This variation in habit 
lends to the conclusion that it belongs properly to the rock ridges." 
Like the preceding variety of F. fusca, C. ivhympcri is a truly boreal 
ant. It is our North American representative of the typical paleo- 
boreal C. hcrculeanu8 and in the Tnited States is known to occur only 
at considerable elevations in the Rocky Mountains (above 8,000 feeti 
and on the summits of the Green Mountains of Vermont. The types 
of whtfinpori were taken in the mountains of Alberta, B. C, by the noted 
monntain climber, to whom the variety was dedicated. I have seen 
specimens from Nova Scotia (Russell) and Labrador (Henshaw). 

The foregoing series of Formicidae, though represented by only ten 
different forms, is of considerable interest on account of its pronounced 
boreal character. Only two of the forms {Formica subsericea and 
Ta/iinoina sritsilc) are abundant at ordinary elevations in the northern 
states, Myrmica canadensis, Leptothorax canadensin, Formica ascrva. 
F. ohscuriventris and Lasius neontger occur sparingly in the same 
region, but always in situations which indicate that they are not in 
their optimum environment or station, or where they seem to represent 
the laggai'ds of a wave of post-glacial migrants to more northern lati- 
tudes cr higher altitudes. F. adamsi. F. neorufiharhis and Camponotiis 
trhyviperi are exquisitely boreal ants of circumscribed alpine distribu- 
tion in the United States, but probably of extensive range in British 
America. 



ECOLOOY OF ISLE ROYALE. 



THE COLD-BLOODKI) VKKTRRKATES OF ISLE BOYALE. 

DR. ALEXANDER G. RfTIIVES. 

The collection wliitli Las wrved as the basis for this report was made 
by the T'nivereity of 5IkliTK(in Muapiini cxiiedition to Tale Royale, in 
the Mimmer of 1!)0j». The rei»ort fthouhl be considered as supplementary 
to the papers upoo the fish, ampliibians and reptiles of the island, pub- 
lished in 1905. (Kuthvpn, IflOy. pp. 107 112.) This, the second expedi- 
tion to iBle Royale, hns added a nunilier of species to the fauna, and 
has established the fact that most of the previoosly known forma ex- 
tend thi-oughont the entire length of tlie island, which was, of coarse, 
to l»e expected. 

The amount of data on this fauna accumnlated by the two expedi- 
tions is considerable, ^vhon ii is considered that np to 1904 practically 
nothinn; was known of the cold-blooded vertebrates of the Island. Our 
knowledf^, however, is still very incomplete. In the case of the 
fishes this is due to the fact that no systematic attempt was made by 
the field parties to secure these forms, and the specimens obtained are, 
in most instances, those that came most easily to band. The list ifi, 
therefore, nndnuhtedly very incomplete both as regards the number of 
species and their distribution. On the other band, particular attention 
was paid to the amphibians and reptiles, and. although there is still 
much to be discovered concerninfr the local distribution of the species, 
the complete list includes nearly all of the species which would be ex- 
jjected to occur on the island. 

\ature of the fauna. — The cold-blooded vertebrate fauna of Isle 
Royale, as at present known, consists of eighteen fish (exclusive of 
Trifffopsis thompsoni, whicli was taken in deeper waters of I^ke Su- 
perior), one toad, one tree toad, three frops, the mud puppy, and two 
snakes,* 

AffinitifH of thr fnmia. — Adams, on a previous pafce, has dwelt at length 
on the fact that Isle Royale has never been connected with the main land 
wince glacial times, a fact that is of first importance in discussing the 
origin of the fauna. Most of the fish obtained on the island occur both in 
the inland waters and in the bays and coves about the shores. Since they ' 
are, moreover, forms of general disti-ibufiou in the Great I^akes drainage 
system, occurring also in Lake Superior, their presence on Isle Royale 
is easily explained. To account for the presence of the inland, brook- 
dwelling forms, however, another explanation must be sought; for such 
species as the common stickleback, ninc-spined stickleback, black-head 
minnow and Lcudsciis itcotjaciis can hardly be conceived as able to 
cross the fifteen miles of open lake intervening between the island and 
the nearest mainland. At present we have no data that throw light 
on this problem. 

The same diSBculties arise in attempting to account for the origin of 

I. 1905. pp. 109-112) rtala irr»irolorin<l ThamaophiM taarUu* h«TB 
but the records cannot be verified. J | ij 



330 MICHIGAN SURVEY. 1908. 

the aniphibian and reptile faunas. As ia the caae of the fish, the npeciea 
are all of general distribution in northeaetero North America, but, 
with the exception of the mud puppy, none of the species recorded from 
the island are aquatic, and, as thej also belonf^ to groups which are 
very sensitive to cold, they could neither reach the island through the 
water in summer or over the ice in winter. The theory of involuntary 
transportation thus seems to he the only tenable one. At present the 
most plausible explanation for the presence of the reptiles and amphi- 
bians (with the exception of the mud puppL?, which might swim across) 
found on the island is that they have been transported on driftVood, 
Unlike several of the other groups of animals, and the flora, the 
amphibian-reptile fauna is not strongly boreal in its affinities. It ib 
true that the forms which are found on the island also range to the 
northward, but the principal range of the species is to the southward, 
and only one species {Rana septmtrirmalis') does not extend rather far 
south in eastern North America, The southern affinity of this fauna 
is undoubtedly due to the fact that the amphibians and reptiles are both 
pre-eminently warm climate groups, and the representatives in this 
region are those few (hat are able to endure the colder climate. 

ANNOTATED LIST. 

Pisces. 

1. CafOKtomiin mmmcrmmi fTJ^c^<p^de). Common Sucker. Taken 
in the sontheast coves of Rock Harbor fllT. 6). As this species was 
found in a similar habitat at the south end of the island in 1904, it is 
undoubtedly to be fonnd in all of the suitable hays and oovea along 
the shores, and probably also in the liirger inland lakes. 

2. Fimephdlen promelas Raflnesqtie. Blaok-head Minnow. Speci- 
mens of this fish were taken in Pumner Lake fllT. 5). This is the only 
locality known for the island. 

3. Lnwincus nmgapus (Cope). As in the case of the Rlaok-head 
Minnow, this species was only taken in Sumner Lake (TTT. 5). 

i. Coretjonuit qnadrUateralix Richardson. Menominee Whitefish. 
This species, a common food fish in I^ake Superior, was taken in Siskowit 
Lake fV). 

fi- Artfi/rosnmvs artcdi fljc Sueur), Lake Herring. Taken hy the 
1905 expedition in Rock Harbor and Lake Desor fVIT. '04). Like the 
Sucker this fish, which is a common Great Lakes species, probably occurs 
in most of the larger inland lakes. 

6. Ar0msomm nitrripinmis 0\\\. Bliic-fin; Black-fln. This flsh was 
only fonnd in Rock Harbor. 

7. Cristivomcr namaj/cuah fWalbaum). Mackinaw Trout; Lake 
Trout. Adult specimens were taken in Rock Harbor, and a single im- 
mature specimen (41 mm. in length) in Benson Brook fll. 1). 

8. Salvelinus fontinnlh (Mitchell). Brook Trout, The 1905 expedi- 
tion secured specimens of the Brook Trout only in Benson Brook (IL 11. 
As it was found on the southern end of the island, in Washington Harbor 
and river, in 1004. it may be considered as occurring throughout the 
length of the island, in suitable habitats. 

9. Liiriiin lucms (Linnaeus). Common Pike; Pickei-el. Taken in Sar- 
gent Lake. This is apparently the only Isle Boyale record. 



ECOLOGY OP* ISLE ROTALE. 33l 

10. Eitcalia inconstans (Kirkland). Brook Stickleback. This species 
was found iu the following localities: Tamarack awamps, giskowit Lake 
(V. 5) ; Spruce swamp, Siskowit Lake (Y. 11) ; Sumner Lake (III. B). 
It is pTobabl; to be found in most of the ponds and small streams on 
the island. 

11. Pygosteus pungittta (Linnaeus). Ninespined Stickleback. The 
Nine-spined Stickleback is represented in the collection by specimens 
from the "Bulrush and Delta zone at the western end of Rock Harbor" 
{III. 3), and from Tobin Harbor (IV). 

12. Percopsia guttatus Agaasiz. Trout Perch. This fish was taken 
about a small island in Tobin Harbor (lY. 6). 

13. Ferca flaveacens (Mitchell). Yellow Perch. Taken in Forbes 
Luke (II. 5). This sitecies is probably to be found in most of the larger 
inland lakes as well as in tbe coves and harbors about the island. It 
was taken in Washington Harbor in 1904. 

14. Cottiis ictalops (Bafinesque). Miller's Thumb. This cottid was 
found along the shores of Rock Harbor (III. 6) and the island iu this 
harbor (III. 2). As it was found in a similar habitat at the southern 
end of Isle Royaie in 1904, it may be coueidered to occur throughout 
the entire length of the Island in this habitat. 

15. Vranidea franklini (Agassiz). There are specimens of this form 
in the collections, labeled Rock Harbor and Benson Brook (II. 1), 

16. Triglopsia thompaoni Oirard. Three specimens of this rare 
species were taken from the stomachs of Lake Trout {Criativomer namay- 
cash) taken by flshermen off the east coast of Isle Royaie. Jordan and 
Erermann write of this form as follows; "Deep waters of the Great 
Lakesj not common; known from Lake Michigan and Lake Ontario; 
doubtless a relic of a former arctic marine fauna, and descended from a 
species of Onocotfus." BoUman (1890, p. 225) records a specimen from 
Torch Lake, Michigan, which was also found in the stomach of a Lake 
Trout. 

17. Lota maculosa (Le Sueur). Lake Lawyer; Burbot. Taken in 
Tobin Harbor (lY. 5) and Rock Harbor (III). 

AmpMhia. 

1. liecturus maculos'ua (Bafinesque). Three immature amphibians 
that are undoubtedly this species were taken in Benson Brook. They 
are verj' young and lack the dorsal fin and stripes. Dr. L. Stejneger, who 
has kindly examined these specimens for U8> states that tbe limbs and 
gills are proportionately shorter than the smallest in the U. S. National 
Museum. It should be noted here that Yarrow (1883, p. 144) has pre- 
Tiously recorded this species from the island. 

2. Bufo amtricanus (LeConte). Common Toad. The capture of a 
number of specimens of this species on the northern part {II) estab- 
lishes its occurrence throughout tbe length of tie island. 

3. Syla pickeringi {Storer). Pickering's Tree-fri^. This amphibian 
was taken in the woods on tbe northern end of the island (lY. 8), and 
in the woods {Y. 4) and Tamarack swamps (Y. 5) in the vicinity of 
Siskowit Lake. It probably occurs also on the southern end of the 
island, although it was not taken by the 1904 expedition. 

i. Rana septentrionalia Baird. Mink Frog; Northern Frog. A single 
specimen of R. scptentrionalis was secured at Sumner Lake (III. 5). 
This establishes the presence of the species on Isle Royaie, a point that 



MICHIGAN SURVEY, 1908. 



ha£ hitherto been in question, owiog to tlie unidentifiable condition of 
the specimens taken on the island bv Dr. A. K. Foote (see Rufhven 1904, 
110). Mies Dicltersou (1900, 225) writes of tlie habits of this frog as 
follows: "The Northern Frog is described as decidedly a river frog; 
it is never captured in lakes and ponds." Our observations are exactly 
the reverse, all of the specimens taken on both expeditions having been 
found about the shores of the inland lakes. 

5. Rana clamitans Dand. Green Fro};. As represented by the col- 
lections of the 190r> expedition, this is the common frog of the island. 
Numerous specimens were taken on the shores of Kock Harbor (I. 1), 
at Sumner Lake (III. 5), and Siskowit Lake (V). Although it was 
not found on the southern end of the island in 1004, it doubtless occurs 
there. 

6. Rana sj/Jmitica cantuhrigcnsia (Baird). Northern Wood Frog. This 
frog is now known from practically the entire length of the island. 
Specimens were taken by the 1905 expedition at Forbes I^ake (II. 5), 
the small island in Tobin Harbor (IV, 0), and at Siskowit Lake (V. 5). 

Berpentes. 
1. Storeria occipitomaculata (Storerl. Red-bellied Snake. This 
little snake is the characteristic reptile of Isle Royale. It was taken by 
the 1905 expedition at Rock Harbor (I. 7 and IV. 5) and Siskowit Lake 
(V. 5). No notes are available on the habits of the individual specimens 
obtained, bat they are doubtless similar to those noted in 1904. As the 
variability of the scutelJation of this snake has apparently never been 
determined, I add the scale formulas of the specimens examined. 



Huseum No. 


Dorsals. 


bb^s. 


,i'ilS., 


Tempo- 


Oculars. 


caudils. 


v,„,„u. 


Total 
length. 


,e». 


33475 

33476 

33478 

33463 

33104 

33408 

33410 

33411 


• IS 

15 

15 

15 
16 

15 

IS 

15 


I 



7-G 

7-e 

7-8 
7-T 


(I:? 

1-2 
1-2 

1-2 

1-3 
1-2 

ti 
{S 

i-i 

{l:f 

1-2 
1-2 


2-3 
2-2 

2-2 


48 

41 
43 

43 

48 


115 
124 

ISC 
133 

132 

127 

120 
120 




250 
263 
230 
200 
208 
310 
294 


GO 
59 
61 

58 


33413 

33414 

33436 

33416 




2-3 

3-2 

11 
2-2 


243 
235 


52 
00 













• In these apecliiiens there is tio derreuse hi the iiuiiilier or s<'ale rons on the posterior pirl of tbe 
body. (Compare Rulhven. 1908.) 

2. Thamnophis sirtaUs (Linnaeus). Garler-snake. Garter-anakes 
were taken in the following localities: Light-house clearing (I, 7), Etan- 



ECOLOGY OP ISLE ROYALE. 333 

soni Wearing (II, 1), Shoi-e of Siskowit Tjake (V, 1), Sumner Lake (III. 
o), Tamarack Swamp Rock Harbor (V, 5), and SUkowit Bay, Tin? 
specimens of this snake obtained vary greatly in color. A number, like 
the few obtained in 1904, would be referred unliesitatiugly to variety 
parietaliS; were it not for the fact that tliere are other specimens in the 
colle4-tion which have a smaller amount of red on the sides, as well as 
some in which it is entirely wanting. Wiien the series ia examined as a 
whole it is quite evident that the speoimeuB are intermediate between 
the typical fomi and variety pai-iclalis, which is not surprising since 
the island lies in the latitude of the kuowii "intermediate zone" to the 
southward (see Ruthven, 1908, p. 168). 

This, however, in no way vitiates the statement made in 1904 that 
the presence of this snake on the island is an evidence of a westem 
affinity in the fauna; for the frequent presence of a considerable amount 
of red pigment on the aides and the tendency toward tlie fusion of the 
upper row of spots, indicate that the Isle Koyale specimens are more 
closely related to the western parictalis than to the garter-snake of 
eastern Canada and Northern Michigan, which belongs to the typical 
form, in that it never has the interspaces of the first row of spots en- 
tirely suffused with red, nor the upi)er row of spots usually fused. 

A number of specimens from Kock Harbor, differ so much from 
the usual <rolor of Isle Royale specimens as to merit special 
mention. The ground color in the»;e individuals is black or blackish. 
The stripes are dark greenish, the dorsal being indistinct. The spots 
of the first row are distinct, the interspaces being of a light bluish color; 
the upper row of spots is usually fused except for short bars or spots 
of bluish. Kelly dark blue with a black band on the outer margin of 
each scute. Supralabials dark blue, bases of the second to the fifth, 
white. Iris black. These si)ecimens are identical with those from New 
Hampshire described by Allen (1899, 64) as pallidula; that they are 
only dark individuals of the* form wiiich inhabits the entire island is 
shown by the numerous "intermediate'' specimens in this collection. 

References. 
Allen, O. M. 
1899. Notes on the Reptiles and Amphibians of Intervale, New Hamp- 
shire. Proc. Host. Soc. Nat. Hist., XXIX, 0.1-75. 
Itollman. O. H. 

1890. A Report ui>on the Pishes of Kalamazoo, Calhoun, and Antrim 
Counties, Michigan. Bull. U. S. Fish Com., 1888, 219-225. 
Dickerson, Mary E. 

1906. The Prog Book. New York. 
Jordon, T>. S. and Evermann, B. W. 
1898. The Fishes of North and Middle America. Bull., 47, U. S. 
Nat. Mus., Vol. II. 
Ruthven, A. G. 

1905. The Cold blooded Vertebrates of the Porcupine Mo,untainH and 
Isle Royale, Michigan. Ann. Kept. Geol. Surv. Mich., 1905, 
107-112. 
1908. Variations and Genetic Relationships of the Garter-snakes. 
Bull. CI, TJ. S. Nat. Mus. 
Yarrow, H. C. w\c 

1883. Olieck List of North American Reptilia and Batrachia.^ Mil. 
No. 24, U. S. Nat. Mus. 



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ECOLOGY OF ISLE ROTALE. 



ANNOTATED LI«T OF THE BIRDS OF ISLE BOYALE, 
MICHIGAN. 

BY MAX MINOE PEET. 

1. Introduction. 

Our observatioDB on the birds of IbIc Boyale extended over the period 
between July 5 and September 22, 1905. Three parts of the island were 
stadied, namely Kock Harbor, Siskowit Bay (especially near the outlet 
of Siskowit Lake), and Washington Harbor. The party remained at 
Bock Harbor from July 5 to August 1; at Siskowit Bay from August 1 
to August IT; and at Washington Harbor from August 17 to September 
22. At Rock Harbor the ol«ervation9 were made by O. M'Creary, N. 
A. Wood, and Dr. B. A, Brown. At Siskowit Bay the work for the first 
week was carried on by M'Creary and Wood, as Brown had left the 
island; on August 8 they were joined by the writer. On our arrival 
at Washington Harbor M'Creary left the island, and the work was con- 
tinued by Wood and the writer until September 1, when the former 
was called home. However, he was forced by severe storms to remain 
on Washington Island at the mouth of the liarbor until September 5, 
and while there made a number of observations which are included under 
their respective heads. The observations during the remaining period 
(September 1 to September 22) were made by the writer. Before 
joining the party at Siskowit Bay, he had spent three days, August 5 
to 8, at Washington Harbor, the observations giving some idea of the 
bird life at that place before migration had set in. 

Practically all the birds observed at Rock Harbor were nesting, those 
observed within a few days after our arrival at Siskowit Bay may also 
be considered as breeding, but after about the first of August it is not 
safe to say whether the bird nested there or was an early migrant. As 
an example of this we may cite the case of the Tennessee Warbler, which 
probably did not nest on the island, and yet was first observed there 
August 2. Unless the nest was found or young unable to fly, we did 
not consider them as breeding in that vicinity, if seen after August i. 
In the case of the waders,- the earliest migration date must be placed in 
the latter part of July. 

Under the head of stations, the particular habitats in which the birds 
were actually found are given with their numbers, so that a fuller 
description of the conditions existing there can be easily obtained by re- 
ferring to that number under the "Description of Stations." It must not 
be supposed that the birds were limited to the station in which they are 
recorded. In all probability the birds noted in one tamarack swamp 
would be found in nearly every similar habitat on the island. But owing 
44 



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838 MICHIGAN SURVEY, 1908. 

to the limited time Bpeot here it was of course impossibie to examine 
every locality, and so the records simply indicate the particular habitatB 
in which the species under disciipsiou were actually found. If observa- 
tions were conducted for a sufficiently long period, the majority, of the 
birds on the island would probably be recorded for nearly every habitat 
even if tliey did not breed in them; especially would this be true during 
migrations. It is not my intention, therefore, to give every habitat 
in which a particular species viight be found, but rather to give the 
habitats which are preferred by that bird, — conditions which can be said 
to be characteristic of that species. 

When a species was seen before the opening of migration, and yet 
no other signs of its breeding were found, it was considered simply 
as a resident, and the first and last dates when it was noted are given. 
The migration records of the resident birds are prohably nearly all later 
than they ehonld be, but the dates are given when they were first seen in 
actual migration. Many of the birds were still migrating jit the time I 
left the island (September 21), and in snjch instances this is the last date 
given, and signifies that the migration of the bird wns still under way. 
More ext«ided observations on this interesting movement of the birds can 
be found in the paper "The Fall Migration of Birds," which is included 
in this volume. The paper on "The Ecological Distribution of Birds" 
should also be consulted for a discussion of that phase of the work. 

From July 5 to September 22 we recorded 63 summer residents, 3 
winter residents, 31 migrants, and 14 permanent residents, making a 
total of 111 Biiecies. In 1904 we obsen'ed eight birds which were not 
recorded the second year; these were: Sora, American Coot, Ijeast Sand- 
piper, Rhort-enred Owl, Bronzed Crackle, American Goldfinch, Clay-col- 
ored Sparrow, and White-breasted Tiuthatch. Besides these, the Club- 
■ house i)eo])le described three other forms, the Snowy Owl, Snow Bunting, 
and Ijapland Ix)ngspur, niaking a total of 122 species known to occur on 
the island. Many ducks come to the island, but the descriptions given by 
the fishermen were of no help' in their determination. A complete list 
of the birds observed during the summer and fall of 1905, arranged as 
"Summer Residents," "Migrants," '-Winter Residents," and "Permanent 
Residents," is included in this paper. Forty-two species were found 
breeding, 

T have attempted to make this more than a simple annotated list — 
a list giving nothing but the occurrence, relative abundance, and dates 
of migration. Besides this usual data, I have given as complete a life 
history of each species as I could, using nothing but the original records 
secured by the expedition. All habitat records are also included so that 
the characteristic environment of the birds may be understood. 

I wish to acknowledge my Indebtedness to Mr. Chas. C. Adams for the 
opportunity of accompanying the expedition, and for his kindness and 
assistance in the preparation of this paper. 

Specimens representing nearly every species found on the island were 
secured and are now in the collection of the University of Michigan 
Museum. For the determination of certain specimens we are indebted 
to Mr. n. C. Oberholser of the Smithsonian Institution. 



3y Google 



ECOLOGY OF ISLE ROYALE. 
2. Classified List of Birds Obacrvid hi lO'l.'}. 



1. Summer Residents, 

ried-bitled Grebe. "Xt. 

Loon. 'Si. 

American Herring Gull. *.'!"(. 

American Merganser. ".30. 

Hooded Merganser. ".'^7. 

American Bittern. *^S. 

Spotted Sandpiper. 39. 

Marsh Hawk. 40. 

Sharp-shinned Hawk. •41. 

Cooi)er8 Hawk. •■12. 

American Goshawk. •4.3. 

Red-tailed Hawk. 44. 

Red -shouldered Hawk. *i5. 

Pigeon Hawk. •4({. 

American Sparrow Hawk. 47. 

American Osprey. •48, 
Saw-whet Owl. 

Blackbilled Cuckoo. "40. 

Belted Kingfisher. 50. 

Yellow-bellied Sapsucker. •51, 

Flicker. *!>2. 

Whip-poor-will. Ki. 

Night Hawk. .^4. 

Chimney Swift. ,55. 
Kubv-throated Huiiimingbird.*5G. 

OliTe-sided Flycatcher. "57. 

Yellow-bellied Flycatcher. •58, 

Alder Flycatcher. ".50, 

American Crow. •fiO. 

Vesper Sparrow, •61, 

Savannah Sparrow. *«2. 

Slate-colored Junco. •C3, 



• = Breeding. 

White-throated Sparrow. 

Chipping Sparrow. 

Song Sparrow. 

Swamp Sparrow. 

Cliff Swallow, 

Bam Swallow. 

Tree Swallow. 

Bank Swallow. 

Cedar Waxwing. 

Red-eyed Vireo. 

^'ashville Warbler, 

Black-throated Blue Warbler, 

Myrtle Warbler. 

Magnolia Warbler. 

Bay-breasted Warbler. 

Black-throated Green War!)- 

ler. 
Oven Bird. 

(Jrinnell's Wafetthmsh. 
Mourning Warbler. 
Canadian Warbler. 
American Redstart. 
"VM liter Wren. 
Brown Creeper. 
Ked-breasted Nuthatch. 
Chickadee. 

Golden-crowned Kinglet, 
Wilson's Thrnsh. 
Olive-backed Thrush, 
llermit Thrush. 
American Robin. 
Bine Bird. 



Baldpate. 

Green-winged Teal. 
American Scanp Duck. 
Canada Goose. 
Wilson's Suii*. 
Yellow Irf'gs. 
Greater Yellow Legs. 
Solitary Sandpiper. 
Killdeer. 

Broad-winged Hawk. 
Kingbird, 
Phoebe. 

I>ea8t Flycatcher. 
Thick-billed Redwinged 
Blackbird, 
Busty Blackbird, 



Migrants. 

Ifi. White-crowned Sparrow. 

17. Lincoln Sparrow. 

IS. Migrant Shrike. 

1!J. I'hiiiidelphia Vireo. 

20. Blue-headed Vireo. 

21. Black and White Warbler. 

22. Tennessee \A'arbler. 

23. Cape Jlav Warbler, 

24. Black-poll Warbler, 

25. Palm Warbler. 

20. Connecticut Warbler. 

27. Wilson Warbler. 

2S. American Pipit. 

20. Catbird. C'.^t^nln 

30. Ruby-crowned Klti^h'^'a'^ 

31. Gray-cheeked Thrush. 



340 MICHIGAN SURVEY. 1908. 

3. WiDter Kesidents (migrants from tUe north). 

1. Hoi-ned Lark. 3. Northern Shrike. 

2. Pine Grosbeak. 

4. Permanent Residents, 

•1. I'rnirie Sharp-tailed Grouse. 8. Northern Pileated Wood- 

•2. Bald Eagle. pecker. 

"3. Great-horned Owl. 0. Blue Jay. 

"4. American Hawk Owl, 'lO. Canada Jay, 

5. Hairy Woodpecker. 'll. Northern Raven, 

fi. Downy Woodpecker. 12. Purple Finch, 

7. Arctic Three-toed Woodpecker 13, White-winged Crossbill, 

14. Pine Siskin. 

3. Annotated List. 

1, Podilymhus podiceps (B). Pied-Wlled Grebe, 

Range: British Provinces southward to Brazil, Ai^entine Republic, 
and Chili, including West Indies and Bermuda, breeding nearly through- 
out its range. 

Stations : Wa8hing:ton Harbor, X '04 ; Washington River, II '04. 

Breeding:, Brood of 5 young, Aug. 18. 

The Pied-billed Grebe was not found either at Bock Harbor or Sis- 
kowit Bay, but was a common summer resident at Washington Harbor, 
frequenting the river and upper end of the Harbor, 

Breeding Notes: A family consisting of t^'o old birds and five young 
were seen almost daily at that place. They seldom came oat into the 
harbor. Although very shy when approached from land I succeeded in 
getting quite close when in a rowboat. They were never seen to take 
wing, generally diving or swimming rapidly away upon the approach of 
danger. Sometimes when badly frightened, instead of diving, they would 
rise upon their small wings so that their feet just touched the sur- 
face and in this way half ran, half flew across the water. The young 
were still unable to fly by the middle of September, and I doubt if the 
parents had completed their moult auCBciently to use their wings much 
either. During the rainy days when the creek was swollen and v'ery 
rapid the grebes generally stayed out in the harbor near the river's 
mouth. A shallow spot covered with water plants and grasses near the 
bend in the river was their usual feeding place, 

2, Qai'ia imhcr (7). Loon. 

Range: Northern part of northern hemisphere. In North America 
breeds from the northern tier of states northward; ranges in winter 
south to the Gulf of Mexico and Lower California. 

Stations: Rock Harbor, III, 2; Snmner Lake, III, 5. 
Siskowit Bay, V, 1 ; Siskowit Lake, V, 6, 
Washington Harbor, X, '04. 

Breeding: Ti^'o voung, two or three davs old, were taken on August 
10. 

Common summer resident throughout the island, as shown by such 
records as these: '"Seveu loons seen in the west end of Rock Harbor. 
July 13" and "eight adults seen at Siskowit Bay. August Ist," These 



ECOLOGY OF ISLE ROYALE. 311 

birds had not left the iBlaud September 21st, aa the fishermen reported 
them at this time. 

They appeared to be more common at the northern end of the island 
and at Siskowit than at Waahin{»ton Harbor. It is doubtful whether 
any nested in the immediate' vicinity of the latter pla**. 

Breeding notes: Prom their actions a pair were supposed to be breed- 
ing on Sumner Lake (III, 5) during July, but no nest was found. 
As long as any one was in sight the pair remained together, calling 
and diving continuously, often coming up many rods from their diving 
point. 

On August 10th. a pair of adult birds were found with their two 
young on Siskowit Lake, The birds were swimming together with their 
young close beside them, .\lthough apparently not more than two or 
three days old, they »-ere expert divers and could swim uuder water 
much faster than the boat could be rowed when pursuing them. When 
approached, the parents swam rapidly away, leaving the young to take 
care of themselves, which they seemed perfectly capable of doing, and 
would have, had it not been for the use of a shot gun. As the young 
were approached, they swam rapidly away at right angles to each other. 
Upon being closer pressed th^ dived, swimming under water for twenty 
or twenty-five feet. This was kept up until they were procured. The 
young at this early age were capable of performing that remarkable feat 
for which the adults are so noted — the act of swimming at different depths 
with the head still above the surface. This is not done by diving, but 
simply by linking the body lower down as a fish might lower itself; 
no special motion is noticeable, the- sinking being gradual, and seemingly 
without effort. Towards the last the young swam with only the head 
out of water. One which was only wounded we kept alive twenty-fonr 
hours, after which it was killed as there was no food which we could 
easily procure for it. When placed in a basin of water it swam briskly 
about, seldom using the whole leg, but simply the foot, bending at the 
upper end of the tarsus, which was moved hack and forth with a fanning 
motion, the toes folding back on the forward stroke. It showed little 
fear, even when taken in the hand. Occasionally it uttered a call or 
cry, much resembling that of a young turkey. In the stomach of the 
other was found a dragon-fly nymph and four small fish about an inch 
and a half to two inches long, which shows that the fish diet is acquired 
early in this siiecies. Low marshy land suitable for nesting sites oc- 
curred in some part of nearly every lake or bay on the island, and 
probably many breed here every year. 

Miscellaneous notes: Many loons are caught each year on "set lines" 
and also in the gill nets on the shores of the island. One fine male 
was brought to the party by a fisherman, which was caught this way. 
The line had been sunk w'here the water was about 100 fathoms deep and 
about ten miles out in the lake from the Rock Harbor light-house. The 
line was down twenty fathoms, and this loon is supposed to have swam 
down this distance and taken the small herring used as a bait. The 
fisherman reported that this was a very common occurrence, the birds 
sometimes being found at the great depth of fifty and sixty fathoms; 
but this seems improbable. These birds roamed about much in the 
evening and during the night, their loud peculiar cry being heard at all 



S42 MICHIGAN SURVEY, 1908. 

lionrs as tliey paaweij over the camp. Sevei'e etormH like those during 
the first few days of September drove the birds into the seclueion of 
WaHhingtOQ Harbor, where tliey were usually in pairs, and very sliy. 

3, Laiiis argcntatus smithsonianvs (51a). American Herring <iull. 

Kange: Korth America generally, bi-eeding from Maine, northern Xew 
York, the Great Laltea and Minnesota northward; in winter, south to 
Cutwi and Lower (California. 

Wtations: Rock Harbor, I, 1; III, 2. 

Siskowit Lake and vicinity, V, YIII, '04. 
Long and Menagerie Islands, V, 10. 
Washington Harbor, X 'ft4, River. II '04. 

Breeding : Young seen Auj^st 1 and 0. 

Very abundant, their numbers at places being counted in thousands. 
This was the only gull seen on the trip. Gatherin_g on the rocky islands 
in such numbei-e as to make them look like one solid mass of white, 
their cries wci-e almost deafening. In the evenings they visited the 
various places where the fishermen had thrown away the fish cleanings. 
At these feasts the water would be fairly covered with the birds, which 
would remain feeding long after dark. So matter on what part of the 
island, within a mile or so of the water, these gulls were nearly always 
in sight. 

Breeding notes: On August 1st, near Chippewa Harbor, a brood of 
these young were seen swimming with their parents, the former being 
unable to fly. On August 6th a visit was made to the breeding grounds 
of these Bi)eciea at Riskowit, V, 10. Fig. ^5. Here a chain of small 
islands runs nearly parallel to the shore and about three miles distant 
from it. The largest of these is nearly two miles long, and from a few 
rods to a fourth of a mile wide. It is composed of red sandstone, which 
rises out of the lake at an angle of about 20^, The rocks are almost 
bare of vegetation, but above the wave swept zone there is a, narrow 
belt of shrubs and small trees. 

As the island was approached, the rocks could be seen covered with 
the adnlt birds, which, however, soon took wing and circled about our 
heads, making a great noise. As we landed, many of the young birds 
jumped into the water and swam hastily away to the gathering flock of 
adults which was forming just out of range. Others ran and hid in 
the small bushes, while some, particularly the youngest, merely squatted 
down between the rocks. Fig. 58, their mottled, downy plumage fur- 
nishing such excellent protective coloring that many were actually pass- 
ed over in the first search. The great mass of young remained about a 
quarter of a mile out in the lake, while overhead the parents flew and 
screamed. Tiie uesfs were built in crevices, Fig. 59, and nooks in the 
rocks from near the water's edge to the top of the ridge which in some 
places was from ten to twenty feet high. Most of them were placed 
on the southern exposure and were composed of grass, sticks and such 
rubbish as was easily accessible. No fresh eggs were found, and the 
young birds appeared to be a mouth or more old. 

The young when frightened squatted flat on the Itai-e rocks, or squeezed 
theniPelves up in Muall nooks and crannies, trusting to their protective 
coloration for safety. 

It is a common practice among the fishermen at Wa^ingttm Harbor 

■ XiOCH^lc 



ECOLOGY OF ISLE ROYALE. 343 

to take tbe newly hatched young from the nest and raise thein at their 
homes. Some wiio have a few chickens take the eggs and place them 
under a setting hen. These young soon become an tame as chicliens. and 
feed upon any form of table refuse. The writer procured five of these 
young from some fishermen. They ranged in size from one about two 
mouths old and nearly able to fly to a little downy fellow scarcely three 
weeks old. These had all been obtained on some rocks just outside of 
Washington Harbor, They came into the writer's possession on August 
8, and in a few days were turned loose on the beach in front of our camp 
on Siskowit Ray. 

They ate everything offered them, vegetable as well as animal matter. 
Fish seemed to be particularly relished. Whatever they ate was 
swallowed entire if it could possibly be gotten into the mouth. One 
such instance was especially ludicrous. A northern red squirrel with 
simply the skin and head removed was fed to the youngest. Because of 
the weight it was with considerable difficulty that the squirrel was 
started in the right direction. The body was too long, so that when 
swallowed, the tail still protruded from the gull's mouth. By stretching 
its neck us high as possible, most of the tail disappeared, only to appear 
again as soon as the gult dropped its head down on its siioulders, and 
closed its eyes in the satisfied manner evinced by all the young gulls 
after a good meal. 

Often when food was given to them they picked it up and walked to 
the water, swashing it around several times before eating. The downy 
young never ventured into the water where they would have to swim, 
preferring to stand on the beach where the waves would just lap their 
feet. The older ones often swam out a considerable distance from land 
and made the acquaintance of a young wild gull able to fly. After a 
few days this latter bird became tame enough to come on the beach to 
be fed. One of the most characteristic habits of the young gulls was to 
walk to the water's edge where the waves would just wash their legs, 
and dive the head down into the water, raise it quickly and throw the 
water over their backs, at the same time giving the tail a few jerks 
sidewise. This performance was repeated many times each day, often 
not five minutes apart. The young which had acquired their full plum- 
age, but were not able to fly, could usually be told from those which 
were able to do so, by their manner of holding the head. The former 
rarely held the head erect, either when on the water or land, usually 
holding it well forward and often on a level with the back, while those 
able to fly held the head erect and nearly straight above the breast. The 
change appeared to take place immediately after the first flight. For 
many days before this occurred the young gull would be seen jumping 
«p and down on the beach, often to a height of two or three feet, flapping 
its wings rapidly at the same time. The first flight of our largest gull 
occurred one afternoon after one of these performances. Making a 
short run down the sloping beach it rose on its wings with a few rather 
uncertain strokes and sailed out over the harbor. The flight must have 
covered half a mile when it returned and alighted on the water near 
camp. Its alighting was anything but graceful, for not being used to 
this new method of locomotion, it raised its wings straight over its back 
and dropped heavily into the water, nearly submerging itself. When 



344 MICHIGAN SURVEY. 1908. 

it rwe to the surface and had completely arranged its feathers, it held 
its head upright, like the adult gulls, as if proud of its performance. 

General Notes; The Herring Gull's manner of feeding was interestiag. 
.When small bits of fish were thrown on the water, the birds would fly 
down and just pat the water with both feet, at the same time lowering 
the head and picking up the morsel with the bill, not even stopping in 
their flight. When the piece was too large to pick up, the bird alighted 
near it, and either picked it to pieces or swallowed it whole. They were 
often seen dropping into the water from a considemble height, appar- 
ently catching small fleh. The Herring Gull is sometimes taken on set 
lines like the loon, only in this case the book must be near the surface. 
^Tiile on "Ijong Island" (V, 10) the dried body of an adult bird was 
found with a large flsb hook attached to a short line in its throat. 

The majority of these birds go south with the freezing of the lake; 
a few, however, remain throughout the winter around the fishermen's 
huts. Whererer a cut is made in the ice at this time, many of these gulls 
may be found. 

4. Merganser aiiiericanus (129"). American Merganser. 

Range: North America generally, breeding south in the United 
States, to Pennsylvania and to the mountains of Colorado and Cali- 
fornia, 

Stations: Lake Superior (Rock Harbor), I, 1. Ill, 2. Ill, 3. II, 4. 
Siskowit Bay, V, 1. Siskowit Lake, V, 6. 
Washington Harbor, X, '04. 

Breeding: July 13, young; also July 27. 

The American Mei^anser is a ratlier common species on the island, 
breeding in suitable localities. 

Breeding Notes: The fisherman re]>orted several families of adults 
and young at McCargoe Cove on July 11, and on the 13th a female with 
several young was seen in the west end of Rock Harbor (III, 3). A 
much larger Hock of young was seen with the female at this same place 
on July 27th. 

At Siskowit Lake (V, 1), on August 1, a large flock of young not yet 
able to fly were found, and on August 8 another flock barely able to 
use their wings were met near the Siskowit Islands. As many as twenty- 
two young were counted with one female. No young were found this 
year on Washington Biver, but several were seen out in the Harbor, 
which might have been raised here. 

5. Lophodytes ciicullatus (131). Hooded Merganser. 

Range: North America generally, south to Mexico and Cuba, breeding 
nearly throughout its range. 

Stations: I^ake Superior (Rock Harbor), I, 1. 
Sumner Lake, III, 5. 

Breeding: Young observed July 27. 

This species was quite rare, being observed only three times on Sum- 
ner Lake (III, 5) July 26, 27, 20, and at the Caribou Islands on several 
occasions. 

Breeding Notes : From the actions of the single females seen at Sum- 
ner Lake they were thought to have young in the vicinity, but none were 
found. On July 27 a female and six young were seen on the Caribou 
Islands. The young were very small, not more than two we^s old, and 



ECOLOGY OF ISLE ROYALB. 345 

by rowing fast they were overtaken, but escaped by diving. This same 
tlock was seen in this vicinity several times afterwards. 

6. Mareca americana (137). Baldpate. 

Range: North America from the Arctic Ocean south in winter, to 
Guatemala and Cuba. Bi-eeds chiefly north of United States, 

Station: Washington Harbor, II, '04. 

Migration : Sept. T>. 

On September 5 a single iadividual was found on Washington River, 
II, '04. It was poorly colored; perhaps an iminature specimen, 

7. Nettion corolinciisis (13!)). Green-winged Teal, 

Range : North America. Breeding chiefly north of the United States 
and migrating south to Honduras and Cuba. 

Btation : Washington Harbor, II, '04. 

Migration : September 4. 

A flock of five api)eared In the Harbor on the morning of September 
4 but soon disappeared. 

.S, Aythya marila (148), American Scaup Duck. 

Range: North Ameriea, breeding far north. South in the winter to 
Guatemala. 

Stations: Siskowit Bay, V. 

Washington Harbor, X, '04. 

Migration : Aug, 4 to Sept. 1. 

This duck was very rare here. One pair was found on Siskowit Bay 
near Wright's Island on August 4. Just before the hard storm which 
commenced September 1, a flock of these birds came into the harbor and 
were seen near Washington Island (X, '04). 

9. Branta canadensis (172). Canada Goose. 

Range: Temperate North America, breeding in the northern United 
States and British Provinces ; soutli in winter to Mexico. 

Stations: Washington Harbor, I, '04. 

Migration: September 16, 

On the afternoon of September IG, a solitary Canada Goose was ob- 
sen'cd flying over the island in a southerly direction. The residents on 
the island reported that in laie October great flocks of geeae pass over, 
sometimes stopping for a few hours, but never remainiug for any length 
of time. 

10. Boiaurus Icntiginosua (190). American Bittern. 

Range: Temperate North America. South to Guatemala, Cuba, Ja- 
maica and Bermuda. 

Stations : Shore of Sumner Lake, III, 3. 

Breeding : Dead young found on July 18. 

Only one of this species was seen during both years, although there are 
many haunts which seem suitable for it. On July 18 and again on the 
25th a single individnal was flushed from the grassy bog along the edge 
of Sumner Lake. 

Breeding Notes: An old nest was found July 18. It was on a grassy 
tuft in the bog, and contained an addled egg and two dead young. The 
nest consisted simply of a depression in the mat of green and dry grass. 

11. QaUinago delicata (230). Wilson's SniiMj. 

Range: North and Middle America. Breeding from the northern 
United States northward ; south in winter to the West Indies. 



346 MICHIGAN SURVEY, 1908. 

StationB: II, 'W, Washington Harbor. 

Migratiou : August 27 to September 21, 

On August 27, one of these snijie was found in the marshy spot near 
the mouth of the river, and again on September 20 another was found 
in the same place. They were very wary and were only found by walk- 
ing through the grass from which they were flushed. On several occa- 
sions, birds which might have belonged to this species were met after 
dark along the road to Wendigo, but owing to their very rapid flight, 
identification was uncertain. 

12. Totanus fiavipes (255). Yellow-legs. 

Range: America in general, breeding in the cold temperate and sub- 
arctic districts, and migrating south in winter to southern South 
America. 

Stations: Bulrush Zone and Delta, III. 3, Kock Harbor. 

Siskowit Bay, Beach, V, 1. Wasliingtou Harbor, I, '01. 

Migration: July 26 to September 15, 

On July 2t) a bird of this species was seen at close range at III, 3. 
Three others stopped on the bluff above the river September 15, They 
showed little fear, and appeared curious as I approached. 

13. Helodromas solitariua (256), Solitary Sandpiper. 

Range: Korth America. Breeding occasionally in the northern 
United States, more commonly northward and migrating southward as 
far as the Argentine Republic and Peru, 

Stations: Siskowit Bay beaches, V, 1. 
^Va8hington Harbor, I '04, 

Migration: August 6 to September 15, 

This sandpiper was rather common throughout August at Siskowit 
Bay, being found mostly on the bare wave swept rocks. True to its 
name, it was rarely seen when not atone. The pure white underparts 
and olive fuscous head and back made it quite conspicuous as it teetered 
bacli and forth on the rocks. They were seldom seen at Washington 
Harbor, although on September 5 small flocks were seen all along the 
road to Wendigo. They were picking up food and paid little attention 
to the writer, simply running ahead a few feet when approached too 
closely, 

14. Actitin nwcalaria (263), Spotted Sandpiper, 

Range: North and South America from Alaska, south to Southern 
Brazil. Breeds throughout temperate North America. 

Stations: Rock Harbor, I, 1. Siskowit Lake, V, 6. 

Siskowit Bav, V, 1. Menagerie Island, V, 10, 
Washington Harbor, I, '04; X, '04, 

Resident and Migrants: July 20 to September 16. 

The Spotted Sandpiper appeared to he rare at the northern end of the 
island, hut was rather common at the other two localities where ob- 
servations were made. It was seen almost daily at Siskowit and was 
by far the most common wader seen on the trip. At Washington Har- 
bor they often came around the dock and were also met with along the 
river and the road parallel to it. At this latter station the birds were 
probably migrants as they were not seen regularly, being present one 
day and absent the next with perhaps a day or two between thpir 
Tiaits. 



,.byCOOgIC 



ECOLOGY OP ISLE EOYALE. 347 

13. Oxi/c(hu« vocifenis (273). Killdeer. 

Range: Temperate North America, breeding north to Kewfoundland 
and Manitoba, migrating to the West Indies and Central America and 
northern South America. 

Stafionfi: itock Harljor region, II, 2. Washington Harbor, I '04. 

Migration : July 13 to August 5. 

Only one of these birds was seen at the northern end of the island; 
this was on July 13 when one was seen llj'ing over the tamarack swamp 
(II, 2), Xo others were found until August 5 when a flock of three 
we