iiiiiiliii I liiiiiii ill PREFACE INTRODUCTION By R. C. L. PERKINS. ^FAUNA HAWAIIENSIS OR THE ZOOLOGY OF THE SANDWICH (HAWAHAN) ISLES Being Results of the Explorations instituted by the Joint Committee appointed by THE ROYAL SOCIETY OF LONDON FOR PROMOTING NATURAL KNOWLEDGE AND THE BRITISH ASSOCIATION FOR THE ADVANCEMENT OF SCIENCE And carried on with the assistance of those Bodies and of the Trustees of THE BERNICE PAUAHI BISHOP MUSEUM AT HONOLULU. EDITED BY DAVID SHARP, M.B., M.A., F.R.S. 11* SECRETARY OF THE COMMITTEE. VOLUME 1. PART VI. PREFACE By the EDITOR. INTRODUCTORY ESSAY ON THE FAUNA By R. C. L. PERKINS. Pages i — ccxxviii and i6 Plates. CAMBRIDGE AT THE UNIVERSITY PRESS 1913 January 1 5/"/^, 1 9 1 3. CAMBRIDGE UNIVERSITY PRESS ILonBon: FETTER LANE, E.G. C. F. CLAY, Manager WUinlm-Bl) : loo, PRINCES STREET TLmion: WILLIAM WESLEY AND SON, 28, ESSEX STREET, STRAND Berlin: A. ASHER AND CO. » 1Ltivi)i5: F. A. BROCKHAUS flcDJ lovk: G. P. PUTNAM'S SONS iSomtiajj anU Calcutta: MACMILLAN AND CO., Ltd. JUN151S55 3S2070 All rig/i/s >Yseifed IX CONTENTS OF THE THREE VOLUMES, SYSTEMATICALLY ARRANGED. As the vicmoirs have necessarily been published in an irregular manner, the following table laill be found useful. Vertebrata. Vol. I, viz.: — Reptilia pp. 365-368, Aves pp. 36S-465, Mammalia p. 465 Hymenoptera Aculeata. Vol. I, pp. 1-122 and Introduction in Vol. I, pp. Ixxiii-cii Hymenoptera Parasitica. Vol. I, pp. 277-364 and Introduction pp. cii-cxi Hymenoptera, Supplement Vol. II, pp. 600-686 Coleoptera (part i, viz.: — Phytophaga, Rhynchophora, Proterhinidae, Heteromera and Cioidae) Vol. II, pp. 91-270: (part ii, viz.: — Caraboidea) Vol. Ill, pp. 175-292: (part iii, viz.: — Cleridae to Hydrophilidae) Vol. Ill, pp. 367-579: (part iv, viz.: — Anobiidae and Supplement to Ceram- b\-cidae, Curculionidae and Proterhinidae) Vol. Ill, pp. 581-666: Review of Coleoptera, Introduction pp. c.xii-cxliv Strepsiptera. Vol. ill, p. 667 Lepidoptera. (Macrolepidoptera) Vol. I, pp. 123-275 and Supplement Vol. Ill, pp. 345-366. (Microlepidoptera) Vol. I, pp. 469-759. Review cf Lepidoptera in Vol. I, Introduction pp. cxliv-clxx Neuroptera. Vol. II, pp. 31 -89, Supplement Vol. tl, pp. 691-696, and Introduction in Vol. I, pp. clxx-clxxx Diptera. Vol. Ill, pp. 1-92 (incl. Supplement), and (second supplement) Vol. II, pp. 697-700: Review in Introduction Vol. I, pp. clxx.x-clxxxix Hemiptera. Vol. Ill, pp. 95-174, and Vol. II, pp. 531-599: Review in Introduction Vol. I, pp. clx.xxix-ccx Thysanoptera. Vol. Ill, pp. 669-701 and Review in Introduction Vol. I, p. ccxi Psocoptera or Psocidae : (included in Neuroptera, q.v.) Embioptera or Embiidae : (included in Neuroptera, q.v.) Isoptera or Termitidae : (included in Neuroptera, q.v.) Mallophaga. Vol. iii, pp. 305-321 Odonata (included in Neuroptera, q.v.) Orthoptera. Vol. 11, pp. 1-30, Supplement Vol. li, pp. 687-690, and Review in Introduction pp. ccxi-ccxx Thysanura. Vol. Ill, pp. 293-297 and Introduction p. ccxx Collembola. V^ol. Ill, pp. 299-303, and Review in Introduction p. ccx.x; Myriopoda. Vol. Ill, pp. 323-338 and Introduction p. ccxxi Arachnida. Vol. Ii, pp. 443-519, Supplement Vol. in, pp. 339-344, Review in Introduction pp. ccxxi-cc.xxiii : Acarina Vol. Ill, pp. 702-704 Crustacea. Isopoda Vol. 11, pp. 521-526. Amphipoda pp. 527-530 MoUusca. Vol. il, pp. 271-41 1 and Review in Introduction pp. ccxxiii-ccxxviii Vermes, Oligochaeta. Vol. II, pp. 413-426 and Introduction p. cc.xxviii Vermes, Entozoa. Vol. Ii, pp. 427-441 and Introduction p. ccx.xviii F. H. I. b CONTENTS OF THE THREE VOLUMES ARRANGED ACCORDING TO THE AUTHORS' NAMES. ASHMEAD, W. H. Hymenoptera Parasitica Vol. I, pp. 277-364 Bagnall, R. S. Thysanoptera Vol. Ill, pp. 669-701 BeddaRD, F. E. Earthworms Vol. II, pp. 413-426 Chapman, B. L. vide Kellogg and Chapman DOLLFUS, Adrien. Crustacea, Isopoda Vol. Ii, pp. 521-526 FOREL, Aug. Heterogyna or Formicidae Vol. i, pp. 1 16-122 GODWiN-AuSTEN, Col. H. H. Anatomy of Mollusca (intercalations) Vol. II, pp. 277 etc Grimshaw, p. H. Diptera Vol. iii, pp. 1-92 Kellogg, V. L. and Chapman, B. L. Mallophaga Vol. iii, pp. 305-321 Kirkaldy, G. W. Hemiptera Vol. 11, pp. 531-599 and Vol. HI, pp. 95-174 Meyrick, E. Macrolepidoptera Vol. I, pp. 123-275, and Supplement Vol. Ill, pp. 345-366 Pearce, N. D. F. Acarina Vol. HI, pp. 702-704 Perkins, R. C. L. Essay on the Land-fauna Vol. I, pp. xv-ccxxviii Vertebrata Vol. i, pp. 365-466 ■ Hymenoptera Aculeata Vol. I, pp. 1-115, Supplement Vol. II, pp. 600-612, Review in Introduction, pp. Ixxiii-cxi Coleoptera [parts of] Vol. II, pp. 117-270, Vol. HI, pp. 581-644, and pp. 650-666, also Review in Introduction pp. cxii-cxliv Strepsiptera Vol. Ill, p. 667 Diptera (Pipunculidae and Idioniyia) Vol. II, pp. 697-700, Review of Diptera in Introduction pp. clxxx-clxxxix Orthoptera Vol. II, pp. 1-30, Supplement thereto pp. 687-690, Review in Introduction pp. ccxi-ccxx ScOTT, Hugh. Coleoptera [parts of] Vol. Ill, pp. 415-422, 431-434, 455-474, 502-505, 508-538 and p. 644 Sharp, D. Coleoptera [parts of] Vol. 11, pp. 91-116, Vol. Hi, pp. 175-292, 367-579 and pp. 645-650 Shipley, A. E. Entozoa Vol. 11, pp. 427-441 Silvestri, F. Thysanura Vol. Ill, pp. 293-297 and Myriopoda pp. 323-338 t.c. Simon, Eugene. Arachnida Vol. 11, pp. 443-519 and Supplement Vol. hi, pp. 339-344 Speiser, p. Diptera Pupipara Vol. HI, pp. 86-92 Stebbing, Rev. T. R. R. Crustacea Amphipoda Vol. II, pp. 527-530 Syke.s, E. R. Mollusca Vol. i, pp. 271-41 1 Walsingham, The Lord. Microlepidoptera Vol. i, pp. 469-759 XI PREFACE THE importance of the fauna of the Hawaiian islands has long been recognised, but as no adequate exploration of their zoology had been attempted, the British Association for the advancement of science appointed a Committee in the year 1890 "to report on the present state of our knowledge of the Sandwich Islands, and to take steps to investigate ascertained deficiencies in the fauna, with power to co-operate with the Committee appointed for the purpose by the Royal Society, and to avail themselves of such assistance as may be offered by the Hawaiian Government or the Trustees of the Museum at Honolulu." The Committee of the Royal Society just alluded to was appointed almost simultaneously, and the two Committees have continued to work together till the present time. The joint Committee has received the most valuable support and assistance from the Trustees of the Bernice P. Bishop Museum at Honolulu. Indeed one of the most important duties in writing this Preface is to make this acknowledgment, and to return thanks for this support. At an early meeting of the Committee it was decided to limit its investigation to the Land-fauna. Two reasons influenced this decision, viz. (i) that this was as much as the Committee could hope to accomplish, and (2) that while the Land-fauna was known to be undergoing great impoverishment, it was believed that the Marine-fauna was comparatively exempt from analogous changes. The Committee decided to undertake an exploration of the Islands, and was so fortunate as to secure for the purpose the services of Mr R. C. L. Perkins, then a young graduate of the University of Oxford. Dr Perkins continued his exploration for some years. As he has given an account thereof in the Introduction that follows this prefa- tory notice it is unnecessary to give particulars here, beyond saying that he underwent great dangers and fatigues, in his arduous and solitary task, with the most determined perseverance, the most unflinching courage ; camping out in the mountains, without a companion, for periods as long as he was able to carry food and equipment. As the result of his work the Committee found itself in possession of an enormous number of specimens, and in pursuance of its work decided on investigating this material and reporting thereon. It is not possible to state exactly the number of specimens that have resulted from Dr Perkins' labours, but it cannot be far short of 100,000, and not improbably exceeds 62 xii FAUNA HAWAIIENSIS that number. The Insects of the Archipelago were previous to this investigation supposed to be scanty in the number of species, and it was believed that individuals of each species were as a rule also very few. Both these conclusions have now been shown to be incorrect. Dr Perkins estimates the number of known species of this Class of animals to be upwards of 3,300 ; and he considers this number to be probably not much more than one half of the total Hawaiian Insect-fauna. The other Classes of Arthropoda are represented by a considerable number of species. Mollusca is specially rich, nearly 500 species or forms having been recorded. Aves has about 50 peculiar species. The other Classes of animals have been by no means satisfactorily investigated, so that no general zoological Census of the islands can yet be given. But it may be said that at the present time if an exhaustive list of the land and marine fauna could be compiled it might amount to 10,000 species, the great majority of them being peculiar to these precincts. And even this number is liable to be greatly increased if the classes of microscopic animals were included ; the Protozoa being, so far as is known to the writer, still untouched. These points are mentioned because it would be a matter for profound regret were it supposed that the work of this Committee — long as it may have lasted — has completed our knowledge of Hawaiian zoology. The Islands having now passed into the control of a State super- abundant in wealth and power we may hope that some real effort may be made, by means of local Associations or expeditions from the United States, to supplement our imperfect knowledge. At present there is so far as we know, only one Natural History Association in the Archipelago — the Hawaiian Entomological Society, established in 1905. Though the task of the Committee may be considered as still incomplete, various reasons render its dissolution advisable. The Committee has lost during; its existence two Chairmen, and Dr Perkins who has done most of its work is not able to continue to make for it the great exertions he did make for so many years. He has however drawn up for us a review of the subject of Hawaiian Zoology. This, in the form of an introduction, will follow this prefatory note. The fauna of the Archipelago has undergone great changes owing to human interference with the natural conditions. It has always been a difficult task to estimate what part of the Fauna is due to recent natural immigration, what to human introduction. The intimate acquaintance of Dr Perkins with the fauna for so many years gives a special value to his review, which, we have no doubt, will be of great assistance to future investigators. It will be seen from his Introduction that he considers the whole of the fauna is due to ancient and recent immigration, and is now augmented by human introductions. And the peculiarities of the fauna apparently fully justify this conclusion. On this subject we may refer to Wallace's chapter on dispersal and migration (Geographical PREFACE xiu distribution of animals, Chap, ii) which correctly foreshadows the way in which this fauna has been formed. The peculiar conditions of life in Hawaiia are to some extent illustrated by the maps and landscapes that accompany Dr Perkins' Essay. The Committees when first constituted consisted of Dr VV. T. Blanford, Colonel Godwin-Austen, Mr O. Salvin, Dr P. L. Sclater, Mr E. A. Smith, with Sir W. H. Flower and Professor Alfred Newton as Chairmen, Dr S. J. Hickson as Treasurer and D. Sharp as Secretary. Subsequently Dr F. D. Godman and C. V. Riley joined it. Sir W. H. Flower and Professor Newton, the early Chairmen having deceased, their duties were taken up by the present Chairman Dr F. D, Godman. Professor Hickson the Treasurer, and D. Sharp the Secretary, have filled these offices throughout the 2 2 years of the existence of the Committee. The Honourable C. R. Bishop, the founder of the Bernice P. Bishop Museum, has afforded by the instrumentality of the Trustees of that institution the most valuable assistance to the Committee; and as an acknowledgment thereof the Fauna Hawaiiensis has been dedicated to him. It appears from the Accounts of the Treasurer that the Committee has derived its funds from seven sources, viz.: — (i) the Trustees of the Bernice P. Bishop Museum; (2) the Council of the Royal Society; (3) the Government Grant Committee of the Royal Society; (4) the British Association for the advancement of science; (5) the British Museum (Natural History); (6) Sale of the Fauna Hawaiiensis; (7) Bank Interest. The Treasurer reports that the total from these various sources amounts at present to ^47 1 7. 6^-. 3^. The first set of the specimens resulting from the work of the Committee has been placed in the British Museum (Natural History), which Institution possesses the types of all the new species described in Fauna Hawaiiensis (except possibly a very few described though the property of others). The second set of specimens has, as a rule, been given to the Naturalists who worked out the collections, and whose names will be found in the Table of Contents in Vol. I of the Fauna Hawaiiensis. The third set has been sent to the Bernice P. Bishop Museum, Honolulu. A considerable number of specimens has been given to Dr Perkins for his assistance in Honolulu, where he has been for some years a resident. In addition to these extensive parts of the collections, others have been given to the great Museums at Edinburgh, Dublin, Manchester, Cambridge, Christiania, Stockholm, Copenhagen, Paris, Berlin, Dresden, the Netherlands, Florence, Turin, and Boston U.S. The few specimens still remaining are promised to the Museum of Harvard University. D. SHARP (Hon. Secretary of the Committee and Editor of the Fauna Hawaiiensis). ERRATA The following corrections should be made in the text : Vol. I, p. 31, line 3, for 'first' read second. Vol. II, p. 13, line 16, for ' interno ' read externo. ,, p. 607, at bottom of page delete N. conifer, which is an Odynertis. „ PI. V, figs. 7 and 7 n, in description of Plate, for ' A. koelense ' read A. amaurodytum. Vol. Ill, p. 591, second line from bottom, for ' Anobiids ' read Xyletohius. ,, p. 616, line 7, twice, and lines 11, 12, 16 and 18, for 'I' read II. Introduction, p. Ixiii, 6 lines from bottom, for 'isolation' read specific endemicity. XV INTRODUCTION BEING A REVIEW OF THE LAND-FAUNA OF HAWAIIA By R. C. L. PERKINS Arrangement. — Prefatory, p. xv ; general features of the islands, p. xvi ; general aspect of the fauna and flora, p. xxiv ; time devoted to collecting, and methods, p. xxxii ; some previous misconceptions, p. xxxviii ; number of species of insects and causes of their extinction, p. xli ; introduced, immigrant, endemic, p. xlii ; distribution by natural agencies, p. xlvi ; flight- lessness, p. xlviii ; origin of the fauna, p. Hi ; species formation, p. Ixi ; summary of the general part of the introduction, p. Ixx. Special review of some of the divisions of the fauna', viz.: Hymenoptera, p. Ixxiii, Coleoptera, p. cxii, Lepidoptera, p. cxliv, Neuroptera, p. clxx, Embiidae, p. clxxiv, Termitidae, p. clxxiv, Psocidae, p. clxxv, Odonata, p. clxxv, Diptera, p. clxxx, Hemiptera, p. clxxxix, Orthoptera, p. ccxi, Thysanura and Collembola, p. ccxx, Myriapoda, p. ccxxi, Arachnida, p. ccxxi, Mollusca, p. ccxxiii. Vermes, p. ccxxviii. This portion of the 'Fauna Hawaiiensis ' may be termed an Introduction to the study of the land-fauna of the Hawaiian islands, and its chief use may be as a guide to further and more special research. It is rather in the form of a series of disconnected essays than a continuous account, this being due to the fact that it has been mostly written at wide intervals of time and in small portions, when the pressure of other work, which could not be neglected, was relaxed. Some repetition will consequently be found, and it has been necessary finally to cut short some questions, which it was intended should be fully discussed, and also to pass by many altogether. The rather full resume of the Fauna that is given will enable any one to find further illustrations of many of the points that are discussed and furnish hints for further investigation. During the twenty years that I have been working on this Fauna I have received much help in various ways from many people, all of whom I cannot mention individually. To Dr D. Sharp and the late Prof. Alfred Newton I have been particularly indebted for advice and encouragement and to the former also for much help in working out various groups, which otherwise I certainly could not have attempted. Of those, who have their home in the islands, I owe much to the kindness of Messrs Francis Gay and Aubrey Robinson on Kauai, to Mrs Greenwell and her sons on Hawaii, and also to the late Mr R. Meyer of Molokai ' For a similar review of Aves refer to Vol. i, pp. 368-465. xvi FAUNA HAWAIIENSIS and the late Mr Valdemar Knudsen of Kauai, both of whom took much interest in scientific work. To numerous others, many alas! now dead, I am also indebted. Mr G. C. Munro, himself an ornithologist, gave me much assistance on many occasions and once or twice spent a day collecting with me. My friend Mr Albert Koebele, the only entomologist in the islands, during most of the period of my active collecting, accompanied me on several collecting trips and gave me many valuable specimens. Unfortunately for the most part he was absent on economic work in other countries. To the Hon. C. R. Bishop, the founder of the Bernice P. Bishop Museum in Honolulu, to the Trustees of the same, and to Dr W. T. Brigham, the Director, under whose guidance it has become a model institution, I am also indebted for help in various ways. Of recent years my colleagues in economic entomology have furnished me with specimens and information, and I have particularly made use of the observations of Mr O. H. Swezey, who has so successfully studied the life-histories of many of the larger and smaller Lepidoptera. The untimely death of my two assistants Messrs G. W. Kirkaldy and F. VV. Terry, who were particularly interested in the Hemiptera and Diptera of the islands is much to be deplored. To Mr W. M. Giffard I am indebted for the gift of numerous specimens and the chance to examine many others, by which I have been enabled to gain additional knowledge on the variability of many species. From Bro. Matthias Newell I have received much help on many occasions, including the gift of valuable specimens. Mr Scott B. Wilson, who himself studied the Hawaiian Avifauna on various occasions, gave me much useful informa- tion, when I first left England for Honolulu, and other assistance. General features of the islands. The material collected by myself and others and described or enumerated in this work has been gathered almost entirely from the six largest and most lofty of the eight chief islands of the Hawaiian archipelago. The other two islands Niihau and Kahoolawe, each with an altitude considerably exceeding looo ft. above sea-level, have long since been denuded of their forest, having served as pastures for sheep and cattle. If any native fauna now exists on these, it must consist of very few species, and is not likely to be of any great importance. Kahoolawe, as seen from the island of Lanai, is often for days together, owing to the destruction of its vegetation, enveloped in clouds of red dust and may now be said to be blowing out to sea. It is no longer of any value even as a pasturage. Niihau, the most north-westerly of the main group, still supports large numbers of sheep. From Niihau to Hawaii, in a direction north- west to south-east, the main islands extend over a little more than five degrees of longitude. North-west of Niihau is a chain of small islands, reefs, and shoals, continuing the archipelago west through i8 more degrees of longitude, the whole Hawaiian group ■ INTRODUCTION . xvii covering some 1800 miles. These small islands have been to some considerable extent explored in search of birds, but very little is known of their other inhabitants. Laysan, a low island, is remarkable for its Drepanid birds and various Noctuid moths, both clearly allied to species inhabiting the main islands of the group, while Midway also has produced a few insects allied to those on the larger islands. Both these islands, however, have yielded species with no allies on the main group, e.g. Laysan the small bird Taiare faniiliaris and Midway a species of true Chrysopa in Neuroptera and a Noctuid moth of the genus Prodenia. Laysan has for a number of years been worked for its guano deposits, and, more recently, a cable station has been made on Midway, so it is probable that both these islands are now, or soon will be, overrun with imported insects. Niihoa or Bird island rising to an elevation of about 900 ft. is the most lofty of the smaller islands, and was visited by an excursion party some 25 years ago. Its geology was studied by Dr S. E. Bishop, but its zoology was neglected. No doubt it contained some much more interesting animals than the countless sea-birds that were noticed. The vegetation being dry was carelessly set on fire, and probably the most interesting part of the fauna was destroyed. It is known that there then existed a thick-billed Passerine bird, probably allied to Telespiza of Laysan. Oahu, Plate V. Of the forest-bearing islands the one whose fauna has been most studied is Oahu. Here more than half a century ago Deppe and Townsend made an extensive collection of the native birds, at a time when these were evidently abundant, more numerous, as they state, than on the more northern island of Kauai. In more recent times the relative abundance of birds on these two islands has entirely changed, no doubt owing to the chief port and settlement of the group having become established at Honolulu on Oahu, and the greater and earlier interference with its natural conditions, that were brought about thereby. Blackburn's well-known re- searches on the insect fauna were mainly carried out on Oahu during the six years of his residence at Honolulu. Similarly the great variety of its land molluscs and the beauty of many of these have induced many residents in Honolulu to become collectors, either temporarily or continuously. The area of Oahu is 598 square miles and for its size there is reason to suppose that it is or has been one of the richest of the group in its fauna. It is rather more than 70 miles from Kauai, at their nearest points, and 25 from Molokai. Like Maui and Molokai it is formed of a fusion of two islands. Of quadrangular shape the west side is occupied by the Waianae range, of much greater age than the longer Eastern or Koolau range, and, doubtless, existing as a distinct island long before the latter emerged from beneath the sea. The highest peak of the Waianae range is Kaala (4030 ft.). There are several large and deep valleys each containing a stream of water on its western side, and on the eastern side it is also much eroded into deep gulches and valleys. On this side its lower slopes are overlaid by the F. H. I. c XVIll FAUNA HAWAIIENSIS later lavas from the younger Koolau range, so that between the two ranges is an elevated plateau rising to nearly looo ft. above the sea. This plateau was once densely forested, but is now practically denuded, having served as pasturage for cattle, while more recently it has been partly put under cultivation for the production of pine-apples. The extent of forest on the Waianae range is now small, the intro- duced goats and cattle having been largely responsible for its destruction. The Koolau range is much more heavily forested, though it has been greatly denuded during the last half century. The backbone of these mountains is much serrated, and the highest point Konahuanui, rather more than 3000 ft., lies a few miles behind the town of Honolulu, while there are several other peaks between 2500 ft. and 3000 ft. On the windward side for a long extent the erosion has been very great, forming perpendicular or very steep cliffs ; on the leeward side the slopes are much more gradual, but are channeled by many valleys and gulches, which, in the more southern part especially, are often separated from one another by very narrow and sharp ridges at least towards their heads. More recent than the Koolau range proper ace some other craters, e.g. the Tantalus series, lying close behind Honolulu, and forming a favourite collecting ground, owing to their nearness to the city and easiness of access, while younger still are various isolated craters situated on or near the coast, e.g. Punchbowl, Diamond and Koko heads and others. Diamond head near Honolulu is interesting from the fact that it contains in a subfossil state great numbers of the shells of land molluscs, including several species of the Achatinellid genus Aniastra and Leptachatiria as well as Succinea, Pupa, Eudodonta and others. Achatinella itself does not appear to be found with these. A species of Succinea is still living there, but molluscs such as Leptachatina, Eudodonta and Aniastra could no longer exist under the present dry and barren conditions. Formerly vegetation and moisture must have been very different to support these creatures. Oahu, though the area of its forests is much diminished, is certainly rich in the number of species of the plants that compose this. The most extensive forest is found in the north-western area of the Koolau range. Parts of this area, however, appeared to me to afford much less variety in their botanical productions than the southern and more northern ends of the range. I have reason to suppose that the part of the range nearest to Honolulu (and containing, as has been said, the highest peak) has the richest fauna. Kauai, Plate V, the most northern of the forest-clad islands, is separated from Oahu by a channel of rather more than 70 miles, where narrowest, a much greater distance than divides any other island from its nearest neighbour. It is of 'slighdy less area than Oahu, and its outline is more nearly circular. The culminating peak is near the centre of the island, and is above 5000 ft. high. From this summit, Mt. Waialeale, and the adjoining plateau, valleys and gulches radiate to the coast. INTRODUCTION xix As usual, erosion is very great on the windward as compared with the western side of the mountain. There are a number of constantly flowing streams of considerable size, some almost worthy of the name of river. As on Oahu and in fact on all the islands, the forests have been much denuded, but the central portion of the island is well-wooded. In some parts the vegetation is very varied, but in others at no great distance it is very uniform, and it is quite clear that causes quite apart from age may determine the variety or otherwise of the plants on different islands or on different parts of an island. The elevated plateau of the interior is for a large part of a boggy nature, and forms the main watershed of the island. Niihau, the small island west of Kauai and now forestless (as has been already mentioned), was not visited by me. Shells of land molluscs of the genus Carelia have been found in a subfossil state there, indicating no doubt a once heavy forest and considerable moisture. It does not appear that the destruction of the forest has been recent, or that any considerable vegetation of larger growth existed at the time of Cook's arrival at the islands. Molokai, Plate VI. East of the south coast of Oahu, and separated from it by a channel of about 25 miles, is the narrow island of Molokai, with an area of 261 square miles. Like Oahu and Maui it is formed of two distinct components, the western mountain or Mauna Loa, with an elevation of nearly 1400 ft., and the larger and more important eastern division with its highest peaks, Olokui 4600 ft. and Kamakou 4958 ft. Mauna Loa has for years been used as a sheep-pasture and is now almost denuded of native trees. No doubt before the advent of the white man it was fairly well forested, and the empty shells of dead Achatinellids of the genus Amasira have been found there, sure evidence of a condition of moisture that has passed away with its forest. The eastern mountains have also been much denuded of their woods on the leeward slopes and for a large part not much dense forest remains except near the backbone of the mountain, where leeward and windward sides meet. On one large area fairly well covered with forest in 1893, but showing some signs of further deterioration in 1896, I reckoned that between the latter year and 1902 two-thirds or more of all the trees had died. This destruction was due to cattle and an introduced Japanese deer. The latter, indeed, became so plentiful that skilled hunters were brought from California to slaughter as many as possible. Both cattle and deer finally entered the wettest and densest parts of the forest, so that in 1902 the appearance of these was totally different from that which they presented six years previously. The Molokai woods have a fairly varied growth of trees and bushes in some localities, but there is an entire absence of Koa forest, and as the Koa [Acacia kod) is one of the most productive trees for the entomologist, a deficiency of species of insects is natural on this account. Evidently there was once a considerable growth of the allied Acacia koaia or of this and A. koa on the lower slopes, below 2000 ft., but these were at the c 2 XX FAUNA HAWAIIENSIS time of my earliest visit mostly dead, and what remained alive were scattered trees, outside the limits of true forest, and quite unproductive. On the north side of the island there has been much erosion, forming the deep valleys of Waikolu, Pelekunu and Wailau. Lanai, Plate VII, is the smallest of the forest-bearing islands, and its highest point is 3400 ft. above the sea. The forest occupies only a small area of the whole island, in the neighbourhood of its highest ridge, above the Palawai valley. Catde, sheep and wild goats have been responsible for this deforestation. It is quite obvious on examination that parts of the higher mountains of Lanai, though now much drier, were once of the boggy nature of the wettest parts of Molokai and other of the islands. A few of the insects peculiar to such boggy uplands still maintain their existence in spite of the drying up of the forest, but, doubtless, many have disappeared. I found a large variety of mostly small-sized trees in the remaining wooded parts of the island, but, as on Molokai, Koa forest is wanting. One feature in 1896 was the compara- tive abundance of the half-dozen species of native land-birds in this small patch of forest, their number being out of all proportion greater than that of the species on Oahu with its comparatively large forest area. The large Maunalei gulch on the north side contains the one permanent stream of water ; in 1894 the forest on its sides had already been destroyed by wild goats, so that it was difficult and dangerous to descend into the upper parts of the valley from above, since, in climbing down, the bare rocks and earth were easily dislodged both by the climber and by the numerous flocks of startled goats. Maui, Plate VI, a large island with an area of 728 square miles, lies east of Lanai and Molokai and is separated from these islands, at the nearest points, by channels of only about 8 to 10 miles in width. When standing in the mountains of Molokai above Kamalo and looking at West Maui it is quite easy, under favourable conditions of light, to distinguish several of the trees, which are chief components of the forest of the latter, the different colours of their foliage being obvious to the naked eye. Maui consists of two geologically distinct islands united into one, the connection being a sandy isthmus. The west division is much older than the larger eastern mountain, but the connection between them does not rise to any considerable height above the sea (about 150 ft.) as compared with the intermediate plateau of the Oahuan ranges. Probably it has never borne more than a scrubby vegetation, and this fact is of importance in considering the means of distribution of species from Haleakala to West Maui and vice versa. The highest altitude of West Maui is nearly 5800 ft., Eke with its boggy summit being about 1 200 ft. lower. These high altitudes are usually wrapped in mists and rain ; the ground, mosses and other vegetation are always saturated with moisture, so that, unless under exceptional circumstances, collecting in a general way is almost impossible. The streams issuing from these watersheds have eroded great numbers IN TROD UC TION xxi of gulches and larger valleys, the lao, Waihee, Olowalu and others being conspicuous. The vegetation of the west and east side of West Maui is very different. East Maui is formed of the great mountain Haleakala, the highest point being rather more than 10,000 ft. above the sea, and on the edge of the summit caldera. Owing to its altitude. East Maui differs essentially from any of the islands already mentioned in the fact that here for the first time we are dealing- with a mountain, in which there is an extensive area above the limits of forest growth, while the summit is nearly devoid of vegetation. The bottom of the great caldera is about 7000 ft. above the sea and its sides are in some parts precipices of 2000 — 3000 ft. high. It contains numerous cinder craters, which are themselves five to nine hundred feet in height. The northern slope of the mountain is seamed with gulches, the forest-belt on that side intercepting the heavy rain-fall, while the western slope is dry and comparatively little eroded. This northern forest-belt differs much in the variety of its plants in different parts of its extension from west to east. In some parts during recent years many trees have died from a disease of obscure origin, so that fears have arisen as to the permanency of the watersheds, the water supply being of the greatest value for purposes of irrigation. On the drier western side the trees have been much destroyed by cattle, but on the windward side the growth of forest is extensive and in parts forms a very dense belt. Owing to its elevation, the summit of Haleakala is sometimes capped with snow in the course of the winter months. Hawaii, Plate VIII, is by far the largest of the islands, its area being 4015 square miles, or much more than that of all the others together. Its northern extremity is 30 miles distant from Maui. It is of subtriangular outline and has been formed from five volcanic centres. Mauna Loa (13,650 ft.) and Kilauea (4040 ft.) are still active volcanoes, occupying with their slopes the south portion of the island, Kilauea lying east of the large mountain ; Hualalai (8269 ft.), near the middle of the west side of the island, was last in eruption in 1801. Mauna Kea (13,825 ft.) is the most lofty of Hawaiian mountains, and is regularly snow-capped during the winter months. In the Hilo district its slopes bear a wide and dense forest-belt, though this has been lessened on the lower side by the cultivation of sugar-cane and other causes. North of this, in the Hamakua district, the belt of forest is much thinner, and some years since was considerably ravaged by fire. However, although much has been destroyed by cattle or cultivation, it may be said that all the districts, into which Hawaii is divided, still have extensive forest areas. In the variety of their vegetation these forests differ much according to locality, nor is an older part of the island always everywhere more rich than a younger one. Thus some parts of the older Mauna Kea are comparatively uniform in their vegetation, some parts of Mauna Loa well varied ; but it is possible that the richest localities of Kea are richer than those of Loa, and the poorest of Loa poorer than the same of Kea. In the northern corner of the island lie the Kohala mountains, much the oldest xxii FAUNA HAWAIIENSIS portion of the island, and which no doubt existed as a separate island long prior to the emergence of the other volcanic centres. These mountains rise to an elevation of five and a half thousand feet, and on the windward side are covered from the coast-line to the boggy summit with a dense and very humid forest. On this side are the large, deep valleys of Waipio and Waimanu, due to erosion. There is no real break between the forests that clothe the different mountains of Hawaii, but all are or have been in some parts continuous, so as to allow of a ready passage for their denizens from one mountain to another. Doubtless, owing to the destruction wrought by man and his imported domestic animals during recent years, there are now large areas treeless or nearly so, where before continuous forest existed and made the connection still wider. Between the Kohala mountains and Mauna Kea are the uplands of Waimea, which have long been used as a pasturage for cattle to the detriment of their native vegetation. Between the three great mountains Loa, Kea and Hualalai is a vast arid tract reaching an elevation of over 6000 ft, of excessive roughness, being traversed by many lava flows from Mauna Loa. The maps of the islands, which we have been permitted to reproduce by the kindness of Mr Willis T. Pope, Superintendent of Public Instruction in the islands, give a good general idea of the rough and broken character of the ground over which most of the collector's work is done. A map showing the relative positions of the Islands was issued with Part i, Vol. I of this work. As a group the Hawaiian islands are amongst the most isolated portions of land surface. The main islands are placed a little within the tropic of Cancer or in the same latitude as a middle strip of Mexico. They are much nearer to the North American continent than any other, the coast of California being distant about 2100 miles, the portion of Mexico, above mentioned, being considerably more distant still. On the west Hong Kong is in the same latitude. Almost south of the group, the Samoan and Fijian groups, the only islands of considerable size, are 30 to 40 degrees distant. Between these and the Hawaiian group lie a number of small islands, but the nearest of these, Palmyra (5° 49' 04" N.), is very remote and separated by an enormous depth of ocean. The extreme outlying islands (Midway, Ocean, etc.) to the N.W. of the archipelago are equally separated from other lands. The general features of the Fauna are somewhat similar to those of Madeira and the Canary islands and to New Zealand, but in details they are very different from any of these. Tahiti and the Samoan group are not sufficiently well known to make any comparison of much value. Owing to the mountainous character of the islands there is great diversity of climate in different localities. The prevalence of the trade-winds through a great part of the year causes much heavier precipitation on the windward side of the islands, while on the lee side, though the mountains may have copious rains daily, the coastal INTRODUCTION xxiii region gets little or none of these. The region of heaviest precipitation, whether on windward or leeward side, is generally from 1 200 to 5000 ft. above sea level, or corre- sponds with the region of dense forest. The temperature at the United States Weather Bureau Station in Honolulu for the years 1905 — 1909 is recorded as follows : 1905 1906 1907 1908 J 909 Highest shade temp. (Fahr.) 84° 85° 86° 85° 84° Lowest temp. 57° 57° 62° 61° 56° On the lowlands a temperature of 90 or even more is occasionally but very rarely noted, and one as low as 52° rarely occurs. The decrease of temperature with elevation is reckoned as i°F. for each 320 ft. of ascent in the mountains. At the U.S. Weather Bureau Station the annual rainfall in Honolulu for the five years above quoted was i6'99, 2577, 30*13, I9"i7 and 20"8i inches, while in some coastal localities the rainfall would be considerably less. A few miles behind the town the rainfall is much heavier; thus on Mount Tantalus, in 1905, at an elevation of 1300 ft. it amounted to 99'68 inches, while on the ridges behind Tantalus it would have been much heavier still — probably twice as heavy — since they precipitate much of the rain before it reaches the Tantalus group. In a wet district, such as windward Hawaii, the rainfall for the same five years at Hakalau in the Hamakua district at an elevation of 200 ft. was i67"59, i22'8o, i82"6o, I47"i9 and 152-54 inches, and a few miles above, at 1200 ft. elevation, it was much heavier still, 26o"67 (1907), 2o6'33 (1908) and 23375 (1909). In 1907 (a year selected at random) at Waiakea (50 ft.) in the Hilo district of Hawaii it was rainy on 333 days ; at Waianae, a very dry district of Oahu, on 60 days of the same year. As by far the greater part of the Fauna has its home in the wet-belt or forest-belt it will be easily understood that any particular collecting trip can be, and often is, very seriously interfered with by climatic conditions. On the high mountains of the Kona side of Hawaii, there is (where it has not been destroyed) a very dense forest-belt with heavy rainfall and a much drier but well-forested region above this. Something like this is also found on the windward side, but there is this great difference, that, on the latter, the region below the forest enjoys a large rainfall, while on the Kona side the lower slopes and coastal region are for the greater part of the year extremely dry and parched up. In the winter the highest mountains of Hawaii and less frequently Haleakala on Maui are snow-capped, the snow sometimes extending down into the forest-belt. Frost is rarely observed below 4000ft. even in the winter months. In Feb. 1902, in some more or less open forest land on Molokai at 3000 ft. the ground was white with hoar-frost on several consecutive days, and all the tender pink-coloured terminal leaves xxiv FAUNA HAVVAIIENSIS of Myrsine were blackened and killed by the cold. In the denser forest at a higher elevation no such effect was visible, and no signs of frost were seen there on these days. When one is camping in the rain-belt and in search of creatures, for successfully collecting which a fair amount of fine weather is essential, it is very tantalizing to see the lowlands blazing in the sunshine, while the forest-belt day after day is wrapped in fog and rain. General aspect of the Flora and Fauna. The fauna of the islands is necessarily so intimately connected with the peculiarities of the flora, while the peculiarities of the latter are in like manner sometimes evidently connected with the remarkable characters of the fauna, that some consideration of the vegetation of the islands is necessary. Every student of the natural history of the islands is much indebted to the late Dr W. F. Hillebrand, whose "Flora of the Hawaiian Islands" has proved a most valuable work. In some introductory remarks, which he did not live to complete, he cites 860 species of Phanerogams and Vascular Cryptogams as natural immigrants or as peculiar to the islands, after deducting those introduced by man since and before the discovery of the islands by foreigners. Of the 860 species 653 are endemic (or 75"9 per cent.) and 250 of the species belong to 40 endemic genera. Of Phanerogamous plants 81 "4 per cent, are endemic, of Dicotyledinous 85'6. Since Hillebrand's work, as is natural, additional species have been discovered from time to time and some changes have been made in the number of the genera that are endemic, but until the completion of the study of the flora by botanists of to-day the above figures may be accepted. From my own observations I have reckoned that the endemic plants, given as 653 by Hillebrand, are probably not more than two-thirds of those that exist. This reckoning is made from the considerable number of species, not included in Hillebrand, that I have casually observed, when investigating the fauna, and on the consideration that the systematic botanist would generally speaking adopt Hillebrand's views as to specific characters. It is clear that the additions to the list of endemic plants will be mainly in an increase of species allied to those already known, rather than in the discovery of striking new forms. In other words the general features of the flora are adequately known. Owing to the mountainous character of the islands, with great differences in rainfall and temperature at different altitudes, the flora varies very greatly at different heights on the mountains. Littoral plants are very poorly represented, mostly by natural immigrants of wide distribution elsewhere. Many of these have fruits or seeds that float for long periods, and have no doubt reached the islands on ocean currents. A few only are endemic, e.g. the parasitic dodder i^Heliotropiuni anonialuvi) and species of the Composite genus Lipochaela. The littoral insect fauna (excluding foreign forms) is likewise poor, and INTRODUCTION xxv most of the species that are attached to the coasts also range inland over the plains or ascend the dry and bare lower slopes of the mountains. They are xerophilous rather than strictly littoral, and some of them even reappear in the dry open country above the forest-line and rain-belt, even at elevations above 7000 ft. Some, however, if not absolutely confined to the region of littoral plants, yet never wander far inland. These include a number of endemic bees and wasps, which visit the flowers of such plants as Vitcx trifolia, Scaevola kocnigii, Tribuhts, Waltheria etc., the wasps finding their prey amongst these and other sublittoral vegetation, such as species of Sida, Lipochaeta and foreign weeds. The Otiorhynchine beetle Rhyncogomis vestitus is attached to Vitex, the endemic moth, Pyrausta iitorea, to Scaevola, while the caterpillars of a few Noctuid moths and some Hemiptera (both Heteropterous and Homopterous) feed on grasses and low flowering plants that grow in the sand. On the whole the littoral fauna appears to be at any rate less poor than its flora. On the leeward side of the islands, owing to the comparatively small amount of rainfall and (usually) the absence of any heavy rains during the greater part of the year, the vegetation of the coastal region and lower mountain slopes is for the most part dried up, and only after heavy rains does the verdure reappear. At the present time, however, numbers of xerophilous plants, introduced by man, do much to lessen the arid appearance of the lowlands. Chief of these is the algaroba or mesquite {Prosopis), the most valuable tree ever imported into the islands. In some of the driest coastal districts it now forms quite a forest-belt, affording food and shelter to cattle and other animals, and an abundance of excellent firewood. An Opuntia of very early introduction, a mimosa [Acacia farnesiana) and Lantana camara are very conspicuous and to the stranger might have the appearance of being really native. Most of this region and the mountain slopes that lie below the forest, where they are not under irrigation or cultivated, have for years been grazed over by cattle. This region is also overrun by foreign weeds, imported by man, and little is left of the native vegetation. Where there is a covering of grasses, these are generally foreign. How- ever, in the most arid districts of the islands, where foreign weeds have failed to establish themselves to any great extent, or at least do not thrive, and on some of the rough lava flows, avoided by the cattle, a considerable number of native shrubs and trees may still be found. In a few places on these dry mountain slopes there yet remain a few choice endemic plants, now mostly extremely rare, such as the red-flowered cotton tree [Gossypium drynarioides) and the shrubby \\o\^X. (Viola helioscopia). Though so greatly denuded, here and there, either singly or in clumps, we meet with various endemic or immigrant species of interest, e.g. Maba, Sanfahtm, Nothocestruni, Gardenia, Nototrichitim, Dodonaea and Erythrina. Pandanus odoratissimits grows scattered or in thickets in many places, and groves of bread-fruit trees occur rarely, as evidence of former cultivation. A number of plants of aboriginal introduction belong to this region, but all of F. H. I. d XXVI FAUNA HAWAIIENSIS them more or less frequently extend upwards till they form part of the regular forest. The Hau tree i^Paritium iiliaceum) thnve?, equally near the sea-coast and above loooft. elevation in the mountains, where it sometimes forms impenetrable thickets. Akiiritis triloba, the candle-nut tree, and Eugenia malaccensis, the so-called 'native apple,' are most partial to the valleys, the former in some localities extending upwards to an elevation of 3000 ft. or more, the latter attaining only about half that height in the mountains. Both, but the former more especially, may form an important element in the true forest-belt, so as to have the appearance of indigenous plants. Their foreign nature, apart from other considerations, is evidenced by the absence of any special fauna in connection with them. Neither of them are unsuited as pabulum for indigenous insects, since some of the polyphagous species feed upon them and especially on the Aleurites, but they have no special fauna. With these the 'Ki' or ' Ti ' {Cordyline) is often very conspicuous. Apart from these aboriginal introductions, as one ascends the lower slopes or valleys and there is an increase in the rainfall, the introduced guava may occupy the whole soil to the exclusion of all other trees or shrubs. Where the moisture is sufficient, the foreign grass, Paspaliiin conjugahim, commonly known as ' Hilo grass,' from the fact that it first appeared in that district of Hawaii, covers the whole ground ; or in drier localities other equally indifferent grasses are in occupation. There is no doubt that what one now sees on the dry lower slopes of the leeward side of the islands is but a remnant of a once more or less open forest land, which has had a very special fauna of its own. In addition to the trees already mentioned some of the more common components of the present continuous forest-belt extended down into this region, Acacia koa and Metrosiderus for instance. Only in special localities can one now get a glimpse of this fauna, for excepting a few wide ranging or hardy species, or such as are specially protected, all the native insects of this region have been destroyed either by imported predaceous insects or by the destruction of the forest itself and the changes of conditions that have ensued. In a few spots there yet remain in Lepidoptera species of Scotoryfhra, Talis, Thyrcopa, Hodegia etc. ; in Coleoptera PlagitJimysus and special species of Rhyncogotms, Proterhinus and Labrocertis, together with some Hemipterous insects, Jassids and Fulgorids, quite peculiar to it. The loss of all but a remnant of this, no doubt, once rich and extensive fauna is much to be regretted, for it may be suspected that here some of the finest native species once occurred. Passerine birds regularly descended into the clumps of trees of this open country, as in fact we have ourselves noticed in special localities in past years. From these, in Cook's time, they passed still downwards, since we are told that several species were found frequenting the coconut palms on the beach. Achatinelline shells as well as other land Mollusca were no doubt numerous, in the upper part at least, of this region. It is certain that in prehistoric, but comparatively recent times, the climate and vestiture of the lowlands of some of the dry districts of the islands must have INTRODUCTION xxvii been very different from the present condition. Thus at the base of the cone of Diamond Head on the coast a few miles S.E. of Honolulu are found extensive deposits of many species of land Mollusca {Amastra, Leptacliatina, Endodonta, Succinea etc.), the species differing little, if at all, from those now living in the forest-clad parts of the Oahuan mountains. Similar deposits are found on the leeward side of the Waianae range at no great elevation above the sea and in various other localities. Such places are amongst the most arid to be found on the island at the present time. While Succinea can be found living near the coast in the driest localities, most of the other genera can thrive only where there is a liberal supply of dampness, and where the vegetation is dense enough to retain this. It is very important to remember that such different conditions have existed in past time, when one considers the possibility of the survival and establishment of any chance immigrants that may have reached the shores. Where the immigrant would now find a barren and arid region on its arrival, very different conditions may have existed in other times. Above the lower dry mountain slopes, at elevations varying usually from 1200 — 3000 ft., one reaches the belt of continuous forest. Where undisturbed by man or beast, this is often so dense as to be impenetrable without cutting a pathway, or at least breaking through by force. Luxuriant growths of leie {Freycinetia) in many localities and in others wiry-stemmed ferns [Gleichenid) make progress difficult. The width of this forest-belt varies very gready according to the locality and according to the destruction that has been the work of man. In some places no such forest-belt exists, in others it has been reduced to an open timbered country, covered with foreign grasses. This is the last stage preceding destruction, when the remaining trees produce their seed in vain, for the thick growth of grass prevents any young trees springing up to replace the old. We have known a forest so den.se that it could be traversed only along a narrow made path, generally knee-deep in mud, to be reduced to open wood- land by the ravages of cattle within a period of about fifteen years. On Oahu (and elsewhere) the lower part of this forest consists or in past time has consisted to a large extent of the fine and valuable Acacia, A. koa, which altogether fails or becomes sparing in the highest forests of this island. On some of the islands (Molokai and Lanai) there is now no Acacia forest, though evidences of its former existence still remain. On the lofty mountains of the other islands the Koa may extend throughout the whole forest-belt, but it is wanting over extensive areas from natural causes. The Ohia, Metrosiderus, is the commonest of all forest trees extending both above and below the region of dense forest, and is the main constituent of the forests throughout the whole group of islands. It is extraordinarily variable, and can adapt itself to the most varied conditions of climate and station, to an extent indeed of which the Koa seems quite incapable. From one of the largest of forest trees it may become, on boggy mountain tops, a trailing shrub, or a small erect plant with simple stem no thicker than a pencil and bearing a single large terminal flower. Its d 2 xxviii FAUNA HAWAIIENSIS variations under more normal conditions are very great, and it is presumably the parent of the other species of its genus that are considered distinct and endemic. Both the Koa and Ohia are of great importance to the fauna, since hosts of creatures are dependent on them. The former probably supports the larger number of peculiar forms, of which, no doubt, many have been exterminated by its extensive destruction. Throughout this forest-belt ferns, both epiphytic and terrestrial, are most numerous and varied, often covering the whole surface of the ground beneath the trees or clothing the trunks of the latter. Here the tree-ferns acquire their greatest size and beauty. The fauna connected with ferns is quite rich, not only tree-ferns but the lesser species also yielding their own peculiar insects. In the Rhyncophorous beetles a number of species of Proterhiniis are restricted each to a particular species of fern, and tree-ferns support Heteramplms and Dryophthoriis. Small crickets of the genus Pai'atrigonidiuin affect each their own special ferns, as do some Hemiptera (Heteropterous and Homo- pterous) and larvae of various Macro- and Micro- Lepidoptera feed only on these. Tree-ferns are a favourite resort of numerous Carabid beetles, and the dead fronds are the home of Machilis and many other creatures. It seems probable that in the islands the fauna attached to ferns is proportionately much more extensive than one usually finds to be the case elsewhere. For this reason some of the endemic birds are constantly to be found seeking for food on the stems and amongst the fronds, and some of the land Molluscs are also very partial to them. It is not necessary to give a list of the trees and shrubs that form this compact forest-belt, for herein are found species of nearly all the important genera of plants, while many are altogether confined to it. Its components are very different in different localities, the difference depending in many cases on the nature of the soil or of the rainfall. In drier localities one will sometimes find a less dense and more varied forest than in a closely adjoining and wetter district, or a wetter part of the belt may be more varied in its vegetation than that at a higher elevation and with a lesser rainfall. Apart from the Ohia and Koa already mentioned the following are very important trees or shrubs to the student of the fauna : the arborescent Lobeliaceae, the Rutaceous Pelea and Zanthoxyluin ; Cheirodendron and other Araliaceae ; the Rubiaceae generally ; Pipturiis and other Urticaceae ; Myrsme (Myrsinaceae), Broussaisia (Saxifragaceae), Elaeocarpus (Tiliaceae), Perrotetia (Celastraceae), the Leguminous Sopliora chrysophylla and an endemic Rubus. However, most trees and shrubs of this belt are more or less productive, and one may add to the above Eugenia sandwzcensis, the numerous species of Pittosporum, Santalum, Myoporuin, Nothocestrum, Charpentiera, Pisonia and the arborescent or woody Compositae. On the highest mountains towards the upper limits of the forest there is often a very uniform flora, rich nevertheless in species of birds and insects. Acacia koa, Sophora chrysophylla, and Myoporiun sandivichense sometimes form its chief com- ponents. Lower down Metrosiderus interspersed with lesser trees such as Myrsine, INTRODUCTION xxix Goiildia, Straussia, Pelea and arborescent lobelias may be dominant. Naturally two such localities though adjoining one another have a vastly different fauna. Of the lesser plants found within the forest-belt, violets with woody stems, a giant Gtmnera, the pink-flowered begonia (Hillebrandia), the liliaceous Asielia, and some of the endemic Labiates are the most likely to attract the attention of the non-botanist. On the summits of the lesser mountains (those not attaining 8000 ft.) there is sometimes developed a bog-flora, characterized by species peculiar to itself. Bog moss, endemic sedges, grasses, violets, Dfvsera and creeping and woolly forms of plants elsewhere erect and glabrous are characteristic. Such bogs are more or less evident on all the forest-clad islands, excepting Lanai, and even there one can see signs that such a condition formerly existed. On the highest mountains of Maui and Hawaii the forest fails long before their summits, often snow-clad, are reached. Above its upper limit dwarf trees and bushes are found with ferns and grasses to which a drier climate is congenial. Sophora, Santaluvi, Myoporiivi, arborescent or shrubby Composites are common constituents, while the Epacridaceous Cyathodes and Vaccinium reticulatum cover much ground. Some of the endemic species of Geranium attract the eye by their silver foliage, while the silver-sword Argyroxiphium, a handsome and peculiar Composite, is found as high as 12,000 ft. A somewhat similar vegetation is found in some localities within the limits of the forest-belt, where extensive open lava fields occur and there is a lack of permanent moisture. On the windward side the forest-belt on some of the islands has suffered less destruction than on the lee side and, if quite undisturbed, it still extends downwards to the cliffs that fringe the shore. Where the coast is not bounded by cliffs the lower slopes are usually denuded, but owing to the rainfall are green throughout the year and well covered with vegetation. The native fauna, however, as on the dry side of the islands has been mostly destroyed by change of conditions and the predominance of imported predaceous insects. The fauna of the windward side is, I think, contrary to what one might expect from the condition of the vegetation, poorer than that of the dry side, although the forest-belt (as on the slopes of Mauna Kea above Hilo) here reaches its greatest development. Wherever I have investigated, this is certainly the case so far as Coleoptera are concerned. However in the rain-soaked woods of the windward forests Lepidoptera are particularly abundant, as also are some Diptera, and the beautiful condition in which one finds the most delicate and minute moths, during the frequent torrential downpours is frequently a matter for wonderment. In such places more hardy insects, such as Pyrantels tammeamea and P. atalanta, Deilephila ivilsoni and others, pay little attention to the heavy showers and may be seen active under such conditions during the day time, while moths in general are equally so after dark. It is not necessary to give any special list of the fauna of the true forest-belt. XXX FAUNA HAWAIIENSIS Herein probably nineteen-twentieths of the endemic species now existing have their home. Here will be found all the genera and species of Passerine birds and nearly all of the land Mollusca, especially the Achatinellidae. Very rarely do these latter pass above the 4000 ft. line above sea-level and then only in the case of a few species of one or two genera or subgenera. Other creatures appear similarly to keep within certain bounds as to elevation, the crickets of the genus Paratrigonidmin and Banza [Brachy- metopa) rarely passing the line just mentioned in spite of the continuity of the forest upwards. Unless the woods have been opened up by man or cattle, wasps fail to establish themselves in the wettest and densest of these. In some localities xerophilous insects and plants pass through dense and wet forests by means of the recent bare and open lava flows, which cannot retain moisture. Above the forest-line on the highest mountains in the open country there is an insect fauna of species peculiar to the region, but it is very meagre as compared with that of the forest immediately below. This could scarcely be otherwise considering the limited character of the flora and the fact that so many of the insects are strictly attached to arboreal plants. Nevertheless, it is now a much more productive country than most of the open region below the forest, for the reason that its fauna has not been destroyed by introduced predators. In particular its species of Carabid beetles remain abundant up to an elevation of nearly 10,000 ft. Wherever the soil and other conditions are suitable a great deal of the lowlands and lower mountain slopes are given up to the cultivation of sugar cane, which on the dry side of the islands is grown under irrigation, but in wet districts usually finds sufficient moisture in the rainfall. For the sugar plantations of dry districts water is procured either from the mountain streams directly, or by tunneling in the mountains, or by pumping from artesian wells on the lowlands or at low elevations. Where cane is not grown either for lack of water or other reasons, the land is often overrun with cattle both above and below the forest and in many cases the forest itself is invaded. Fortunately now that it is realized how cattle inevitably destroy all Hawaiian forests and cause the drying up of the watersheds, on which the entire prosperity of the islands depends, large areas of woodland have been reserved and all cattle excluded. Though this exclusion has not been carried out nearly as rigidly as one might wish yet there is reason to believe that what has been done and may still be done in this matter will preserve a great part of the endemic fauna and flora for future generations. Only the small islands of Kahoolawe and Niihau are beyond redemption so far as native vege- tation is concerned. Leaving out of consideration all foreign plants, whether imported by man or naturally immigrant, the lack of conspicuous flowering plants is a most striking feature not only of the forest but of the open country. Were it not for the natural immigrant, Metrosiderus polytnorpha, the chief constituent of the forests, the latter would be sombre indeed. This tree with its abundant red flowers, when, as is often the case, it IN TROD UC TION xxxi comes into blossom over large areas at the same time, presents a really bright appear- ance, and this effect is heightened by the large numbers of the common scarlet and crimson Drepanid birds that resort to the flowers for the sake of their abundant nectar. It is true that there are some other flowers, such as Gardenia, that are conspicuous and fragrant, or showy like the native species of Hibiscus and the begonia, Hillebrandia, but these are too local or infrequent to change the general impression. In open places the yellow flowers of some of the gregarious Compositae attract the eye, but in other countries would hardly be noticed as being more than a rather superior weed. Frey- cinetia in flower or fruit has for a short season a bright appearance, though in many districts nearly every one is devoured or befouled by foreign rats. Perhaps after Metrosiderus, the Mamani [Sophora chrysophylld) which grows gregariously and also attracts the red birds, makes the best show. Of the arborescent lobelias the flowers, though often large and numerous, are rather strange than beautiful; in many species they are hidden beneath or amongst the foliage and often they appear to decay almost before they are fully mature. On the other hand the ferns with a large variety of species, often of considerable beauty, are the more attractive from the absence of con- spicuous flowering plants. Nowhere in the islands will one find any display of the latter comparable with that which is so usual in the meadows or open woodlands of countries outside the tropics. Having spoken of the general inconspicuousness of the flowers and the bright colours of some of the commonest birds, we may notice the general aspect of the fauna. The greater number of species of the native birds have green, yellow or oliva- ceous plumage and are not conspicuous, though some of them are of strange or even grotesque form. Here belong the Drepanididae, but five genera (at least in some of their species) have red plumage either in both sexes or at least in the male. The Meliphagine Acrulocerais have mainly black or dark plumage, as do several of the Drepanididae, but none are entirely black, since red, white or yellow forms part of the colour pattern. The thrushes {Phadornis) are of sombre colours, but the males of the fly-catchers {Offl.yz^»?/w) are often prettily variegate. The absence of gaily coloured native butterflies or diurnal moths prevents the insects from making any conspicuous showing. It is doubtful whether to ordinary observers there is, excepting the fine Pyravieis tammeamea, a single insect that would be considered beautiful. At least that species only can be said to represent brightly coloured insects in the way that Vestiaria, Himatione and others do the birds. The vast majority of the insects are utterly inconspicuous and unnoticed unless specially sought for. In most places, whether in forest land or in the open, one will observe the endemic bees or wasps, insects of small or medium size, mostly appearing black in life, and the species very uniform in general appearance. Their great abundance in many places, the dark, blue-shining wings of many of them and their activity by day attract our attention. In many woodlands and by the side of mountain streams some of the xxxii FAUNA HAWAIIENSIS larger red-bodied dragon flies of the family Agrionidae may even be called conspicuous, while others like A. nigrohamatum have a remarkable appearance as they fly towards one, owing to the bright yellow colour of the face and the striking turquoise-blue of the eyes. Apart, from these it is quite possible for the casual observer to walk through miles of forest and see not a single even moderately conspicuous insect. At the most, moths of insignificant size and appearance are frequently disturbed from tree-trunks or undergrowth, as he walks along. These remarks, it should be understood, will apply to localities, where there is in reality a rich endemic insect fauna, and serve to show its hidden nature. The arboreal species of Achatinellidae, though not large, are many of them of very bright colours and even beautiful, but in the woods they are far from conspicuous. It is easy to imagine that any one not specially searching for these, or for other creatures in the same habitat, might pass them by daily without being aware of their existence. Even in the wettest weather they do not move around by day with the freedom of many land Molluscs of other countries. The terrestrial species are of sombre colour and well concealed, as also are many of the species which habitually frequent the trunks and branches of trees rather than the foliage. The endemic spiders are mainly inconspicuous and many of the forms that frequent the trunks and branches of trees match their environment so well as to be nearly in- visible. Others remain hidden beneath bark or between leaves spun together. Web- making species are often numerous in shallow caves or hollows in the sides of mountain ravines, or in the cavities of old tree-trunks, or they spin their webs near to the ground. Being so favourite a food of the Passerine birds, the arboreal forms especially, were they not well concealed by day, would have had little chance of survival. The great decrease in the number of native birds may well lead to an increase in the number of spiders, even if it has not already done so. Certainly, in 1892, individuals of the species living exposed on the limbs of trees on Hawaii, where birds were very numerous, were most uncommon. Account of time spent in collecting and methods employed. Before beginning a more special account of the Hawaiian Fauna it seems advisable to refer to the time that has been spent on the various islands of the group in collecting material. I arrived in Honolulu early in 1892 and during the spring months collected chiefly in the Waianae range of Oahu, partly on the eastern and partly on the west side of these mountains, but more successfully on the latter. Except for an odd day or two I did not attempt the Koolau range, which was the scene of most of Mr Blackburn's entomological field-work. Throughout the summer I was stationed on the west side of Hawaii, working chiefly on Mauna Loa, then a paradise for native birds, and to some extent on Hualalai. Collecting was done at all elevations from the dry coast, through IN TROD UCTION """'' "^ | ^ xxxiii the wet-belt, where heavy rain fell almost every day, to the higher and drier forest of 4000 ft. and upwards, and to the summit of Hualalai. The winter months of 1892 — 1893 were spent almost entirely on the western slopes of the Northern end of the Koolau range of Oahu, where the forest-belt was more extensive than elsewhere. This part of the range appears to me to be decidedly inferior for general collecting as com- pared with the Honolulu end, at any rate during so wet a winter as was this one, but here, if anywhere, there seemed to me to be a chance of meeting with certain birds, of which it was specially desired that specimens should be obtained. However, neither the 'Oo,' Acmlocei'ais apicalis, nor the species of Heniignathus and Heterorhyncluis peculiar to the island and obtained by early collectors in the first half of last century, were to be found, either by myself or other ornithologists, and it may be that they had already become extinct there. This, with perhaps one or two exceptions, was as unsuc- cessful a collecting e.xperience, as I ever met with. In May 1893 ^ crossed over to Molokai and, the west half of the island being practically forestless, camped on the southern slope of the forest-bearing end, near the middle of the island. All animal life seemed abundant after my experience on Oahu. Common species of birds were plentiful, though not in the extraordinary numbers observed on Hawaii, and of course all the Mollusca and most of the insects were new to me. This was a very wet summer in the mountains, and for the first six weeks there was hardly a day without long and heavy rains in the woods a few miles behind my camp, and when it was not actually raining, these were mostly enveloped in thick fog", through which objects were visible only for a short distance. In spite of this, collecting was good. After a time I pitched a tent in the midst of the highest boggy forest near the back ridge of the mountains. From my different camps I was able to get to Kalae on the west, where I also made a short stay, to the valley of Waikolu, and down into the deep valley of Pelekunu, where I stayed for a time in the then native village on the windward coast. It was not till the autumn that I was able to leave Molokai, owing to the large amount of time spent in securing the rarer birds. And here I should state, that not only on Molokai, but elsewhere the scarce birds are without doubt difficult to get even in moderate series. I should consider that, both on Molokai and other islands, at least half of my whole collecting time was taken up in acquiring those that I obtained. For some of these birds it was necessary to be continually on the watch, and even then, and in the best localities, it was quite possible to spend weeks without seeing, or even hearing, a single individual. For this reason, until I had secured my specimens, I never dared to go about unhampered with gun and other necessaries for bird-collecting, even though I was specially in search of insects. When it is considered how small and well-hidden are most of the latter, it will be easily understood that the many days spent specially in search of rare birds, for which both sight and hearing need to be kept constantly strained, did not result in a great accumulation of entomological specimens. F. H. I. e xxxiv FAUNA HAWAIIENSIS After my experience of the previous winter I concluded that it would be better to spend this season on the small and comparatively dry island of Lanai, the forest area being very small. Nearly all my extensive series of Lanai birds, shells and insects were obtained during these months. In March 1894 I proceeded to Maui, working with not too much success in the broken West Maui mountains, where I have invariably had bad luck from weather or other causes, and with satisfactory results on Haleakala. As I was returning to England for the winter I was unable to make a stay of more than two months, after which I went to Kauai and collected over the high plateau of the middle of the island and also to some extent on the lower and drier parts of the moun- tains. During this summer I revisited Lanai for a short period of camping and finally proceeded to windward Hawaii for a brief stay. At the end of the summer I left for England and for four months was engaged in working on the specimens, so far collected, and described the bees and wasps that had been obtained, this latter work being revised subsequently and published much later. In the spring of 1895 on my return to the islands I first made a lengthy stay on Kauai, visiting various parts of the islands and afterwards went to Hawaii, collecting in the Hilo, Puna and Kau districts. In the winter months several camping expeditions were made on Mauna Kea, birds being the chief object. On two of these expeditions there was not a single day without heavy rain and the collecting was very trying. On account of the density of the forest, I had a gang of natives with me, in order to cut trails through the forest. They all suffered much from the wet and cold, it being impossible to dry clothes for sleeping in, and one of them met with a bad accident. On my last attempt, however, I had exceptional weather for two weeks and secured the bird I chiefly wanted, and some rather nice insects. In February 1896 I spent some time in a tent on the Waianae mountains, at a higher elevation than had been possible in 1892, and afterwards revisited Kauai (more than once), Molokai, Lanai, Hawaii and Maui. One of the Kauai expeditions, when I camped on the wet side of the island, was very poor in results, but other localities yielded well. In January 1897 some time was spent in collecting on the west side of the West Maui mountains, but this collecting was done under many difficulties. Leaving Maui, I camped out for some weeks on Kauai. Shortly afterwards I left for England by way of Arizona and Mexico. After working on the collections and completing the descriptions of Hymenoptera already referred to, as well as the Neuroptera, Orthoptera and some families of Coleoptera, I returned to the islands early in igoo. In this and the following year my work was almost entirely confined to Oahu. I rarely carried a gun, and paid little attention to anything but insects and mostly limited my observations to certain groups of these. Very little indiscriminate collecting was attempted. Beetles such as are easily obtained by indis- criminate beating of trees or by sweeping these, were almost all collected by special search, as I wished to obtain a more certain knowledge of their habits and particulariy of the restriction of species to one food-plant, the variation of individuals found in INTRODUCTION xxxv company and that of different colonies. Practically all the specimens obtained were mounted, when freshly killed, and to a large extent examined at the time. Observa- tions were made in widely separated localities and in both mountain ranges. During this period I had facilities for working at the specimens in the office of my friend Mr A. Koebele, the Government entomologist of the islands, who accompanied me on a number of my trips. Although, as has been said, my latter work was of a more special nature, yet more new species than might have been expected were obtained. The time spent by me in general collecting (i.e. excluding my last mentioned visit to the islands) did not differ much in the case of the four islands Kauai, Maui, Molokai and Oahu, averaging about eight months to each. On Lanai the small size of the island and its few species of birds only necessitated about four months work, but the great island of Hawaii consumed about twenty months, largely on account of its richness in species of birds, as compared with any other of the group. All my collections were specially packed and shipped in Honolulu, and although practically all the Lepidoptera and many of the other insects were pinned, and must have received much rough handling in their several changes from steamer to train and vice versa, I think hardly a specimen was damaged or even shifted. A single lot unwisely sent by mail, for a special reason, was considerably damaged. In 1902 and 1903, having ceased to collect for the Committee and become engaged in local economic entomology, I was able to visit several of the islands and collected a great deal of material, some of it still being unworked or undescribed, while some of the species are included in the systematic portion of this work. In 1902 I made an exten- sive investigation, by breeding them, of many of the parasitic Hymenoptera and in 1903 I paid a good deal of attention to the native leaf-hoppers (Hornoptera). My account of the birds was published at the end of 1903 (Vol. i, p. 365). After this date except for occasional single days or for short periods I have done little field-work, and when working have nearly always had some special object in view, rather than to collect indiscriminately. Though greatly occupied with economic entomological w.ork I com- pleted the Anobiidae (published in 1910), a group of beetles on which I had been occupied, as occasion offered, for about ten years, and wrote supplementary papers on other Coleoptera and several of the other Orders. As to the groups of insects, which have been worked out for the Committee by well-known specialists, in most of these I have had occasion to determine large numbers of species, or even considerable collec- tions, the material having been collected both by myself and others. I have thought it advisable to refer at this length to such work as I have personally done in connection with the Hawaiian fauna, both in the field and study, because on this are based the conclusions that I have arrived at in this Introduction. From observations of a less wide nature or limited to a single group I might have arrived at other conclusions. It has already been said how much time was taken up in the search for some of the rarer birds and in the acquisition of a good series of specimens of these. As to the e 2 xxxvi FAUNA HAWAIIENSIS insects far more time was given to a special search for Coleoptera than any other Order. After the beetles the Microlepidoptera probably occupied the most time. Many of these were collected in very wet and sometimes windy localities and as they could not be carried alive in so rough a country without damaging themselves, I found it neces- sary to kill and pin them, as soon as they were caught. Liquid ammonia was always used in killing, except for stray specimens picked up, when I was specially in search of other insects or of birds. It was often necessary to carry an umbrella in order to obtain shelter from the wind and rain, when pinning the specimens in the field. On compara- tively few days the Hymenoptera claimed my whole attention, these being (at least the Aculeata) conspicuous and easily collected, and this also applies to the Odonata. The Neuroptera, Orthoptera, and Hemiptera, as represented in our fauna, are naturally met with in collecting beetles and now and then I spent a day strictly in catching the larger Lepidoptera. To the Mollusca on Molokai and Lanai I paid a good deal of special attention and very special attention to certain species. As to the modes of collecting the most successful are exactly those which would be employed by a successful collector in any other country, the difference between the islands and most countries being that many situations, which might be expected to produce a good harvest, will be found comparatively barren. Very few species of Coleoptera will be swept from low herbage, even in the finest localities. Sweeping, when employed, should be rather done amongst the branches of trees and shrubs. This is very natural, seeing that nearly all the beetles are attached to, or bred in wood, dead or alive, but chiefly in dead or diseased trees. A good many of the species sun them- selves on the foliage of living trees and shrubs, but rarely descend to low plants. Comparatively speaking, very rarely is any beetle seen upon the wing. These remarks of course apply solely to the endemic fauna. Light is very attractive to Lepidoptera, both large and small, and especially in wet localities, no doubt for the reason that in these, dark cloudy nights are most frequent. Clear cold nights are almost useless, and worse still if there is a strong wind. Even the smallest moths will sometimes come freely during very heavy rain, and a black windless night with torrential downpours, which cease at intervals, are the very best, during those intervals. Even the butterflies, Pyrameis tammeamea and P. atalanta sometimes are not infrequent visitors under such conditions. Flowers are attractive to many moths both by day and night, especially the Ohia {Metrosiderus) from which at night some kinds may be shaken to the ground, either unable or unwilling to fly. The usual bait of sugar, tried at various times, in good localities for Agrotis, failed to attract anything except the little Hypenodes, so similar to the European one, which, as is well known, is itself very freely attracted. Fern-fronds in a certain stage of decay are very attractive to Agrotis, and when feeding on these they make no attempt to fly, but fall to the ground, when disturbed. Aculeate Hymenoptera and Odonata do not need looking for, but it needs a practised eye to distinguish the very similar species of the former, so as to be able to pick out the INTRODUCTION xxxvii rarities without a wholesale slaughter. Flowers, except a few fleshy kinds that easily decay, are not attractive to beetles, excepting some species of Nitidulidae and the Elaterid genus Eopentlies. The minute and obscure Diptera, the endemic species largely consisting of small Dolichopodidae, which shrink and distort on drying, and of infinite numbers of Drosophilidae, many of these also becoming distorted, have been little collected. It is quite certain that only a fraction of the fauna has yet been collected, and that the groups are very unequally known. The small group of native dragon flies may not be greatly added to and the aculeate Hymenoptera are fairly collected, both of these being as one might almost say obtrusively present as a whole, as compared with other insects, of which but few species are both numerous and conspicuous at the same time. On the other hand three hundred species of DrosopJiila would be a moderate estimate of those existing. It is possible that half the number of existing species of insects have been collected, but this is by no means certain. During the last ten years, since the annexation of the islands, it has gradually become much more easy to get about than was the case during the chief period of my collecting, and many mountain localities have been opened up, which before were impossible for the collector or at least very difficult of access. At the same time the existing forest is being much more strictly preserved from destruction. I did much of my collecting from a small tent, which, usually, I had packed for me on horse or mule over the lower slopes to the lower edge of the forest and from that point packed it myself with the necessary supplies and collecting apparatus into the heart of the forest. Skinning birds and pinning insects while either sitting on the ground or lying down was at first troublesome, but after a time seemed natural enough. In some very wet forests, where I had to spend much time for the birds, I was obliged to carry a limited supply of oil and a small oil-stove on account of the difficulty or impossibility of making a fire on my arrival at night after a long day's collecting, during the continuous and heavy rains. But such cases were exceptional, and otherwise, after a little practice, it was always possible to build a fire in the open (even in very wet weather) for the purpose of cooking rice, — practically the only cooking that the habitual camper-out need attempt. This with coffee and sugar and one or two kinds of tinned meats (in addition to a tent, clothing and apparatus) will be found as much as the collector will care to pack on his back in so rough a country, and I found the stove and oil a sore burden, very reluctantly assumed. Except for very special purposes I never took a native with me on these camping expeditions, partly because it was almost impossible to get one man to go without companions and partly because the impedi- menta would be proportionately increased as the number to be fed and sheltered was added to. In some of these untrodden forests many of the birds (excluding the most wide-ranging species which seem generally to have more or less fear or shyness of man and the little fly-catchers, which are everywhere very tame or inquisitive) were absolutely xxxviii FAUNA HAWAIIENSIS fearless, and it was only just not possible to touch them with the hand. Sometimes it was quite impossible to shoot a rare species, since it would keep following up the collector, as he moved away, perching overhead at only a gun-length's distance. Then, its curiosity satisfied, it would fly off altogether and be lost in the density of the brush. Excepting in the more open forest, in the neighbourhood of ranches, the woods are quite uninhabited, and sometimes for weeks together I never saw a human being. Occasionally a native would come up and leave letters for me or take them down from my tent, but usually I was away collecting, when he arrived. Hawaiian islands much richer in species than has been supposed, and the scarcity of individuals of species exaggerated. Oceanic islands are generally said to produce but few species of native insects, but their unproductiveness has certainly been exaggerated, owing to the hidden life and small size of so many of these inhabitants. It has also been supposed that individuals of the species are as a rule and comparatively speaking rare. In the case of the Hawaiian islands, and probably of most remote oceanic islands, the insects and other animals are limited to a small variety of types compared with other lands, but the types represented are often extremely rich in species, which, one might almost say, do their best to fill up the vacancies caused by the absence of many forms of life almost ubiquitous elsewhere, but which have never reached these remote islands. When talking to the late Dr Ashmead, who visited the islands in 1900, he insisted that certain large groups of parasitic Hymenoptera, of which he had a specialist's knowledge, must, being of universal distribution elsewhere, be really well represented in the islands, though no species (excepting a few evidently imported) had ever been collected. But in reality it is not at all surprising that these do not occur. Firstly the chances of any vagrant insect, carried across the ocean either by wind or wave, happening to land on a few small spots in mid ocean must be very small, and then the possibility of its acclimatisation has to be considered. We know from practical experience that many insects, which it has been desired to introduce, fail to establish themselves in this climate. If a plant-eating species, it may find no fit vegetable food, if parasitic, no fit animal food. But in the case of such specialized creatures, as most parasitic insects, unless either a number of the host, they feed upon, has arrived with them, or previously, and become established, or unless there happens to be some other insect present, which may serve as a new host, the chance of a successful occupation by a parasitic insect is very small. The paucity of individuals of a species is also, I think, much less than has been supposed, this supposition being due to the small size, hidden life and special INTRODUCTION xxxix habits of many of the creatures. I do not feel sure whether the number of really rare species is greater than is the case elsewhere over an eqiial area of land. On large land areas a species may occur in very small numbers, but persistently, over a limited portion of the area, while in other parts removed from this, it may be continually abundant, and small islands do not afford these conditions. On a continental land if one picks out a continuous area, equal in size to one of our islands, there will be found large numbers of rare insects, even in countries much more collected over than the Hawaiian group. It is natural that a collector in a country of great roughness and with extensive virgin forests, collecting in a limited time not only all kinds of insects, but all other land animals, will obtain many unique specimens and also species represented by only one or two examples, both from lack of time to make special search for each particular species and because many insects even though they have no regular season, become more abundant at one period than another and the best time may be missed in a particular locality. Also in pathless forests, where a species is very local, and particular as to the exact condition of its surroundings, it is often not possible to find again the exact place where it happened to be met with. In restricted localities, that have been often visited and collected over, most of the species have at some time or another been found in some numbers and many in abundance. Again the personal equation comes in. The collector particularly fancies certain species and devotes much time to these, while others he is liable to neglect, in so far that he picks up only a few specimens. For this reason the number of specimens cited as captured, in the systematic portion of this work, should not necessarily be regarded as showing the rarity or otherwise of a species. For instance, the conspicuous dark-coloured, day- flying moth, Dasyuris holombra, of which two specimens are recorded, I observed in hundreds, if not thousands, on one occasion, flying in the forest on Maui, but being always hampered with a gun and much occupied with birds, I neglected to catch a series and did not again have the chance. A number of species of beetles, of which I captured long series, are undoubtedly amongst the rarest Hawaiian insects. My experience in this matter is different from Mr Blackburn's, who says " the very common insects" are "few indeed and the rather common ones almost none at all." We should rather say (speaking of course only of the endemic fauna) that the rather common species are numerous, the excessively common ones, less so, though there are a considerable number of these. He also remarks that " It is by no means an unusual thing to pass a morning collecting on the mountains (at any rate on those under 3000 ft. high) and to return home with perhaps two or three specimens secured, and having seen literally nothing else except the few most abundant insects." I am quite sure that this must be due to his having neglected to get beyond the range of the small ant Pheidole mcgacephala, before beginning to collect. A coleopterist of some experience in island work (and perhaps even without this) in the mountains close to Honolulu may take about forty species of beetles in a fair day's work, xl FAUNA HAWAIIENSIS between 1500 and 2000 ft. elevation, though he has a considerable walk to and from the collecting ground. Almost the last day I collected in that locality, I took exactly that number of beetles (Cis 6 species, Proterhinus 10, Carabidae 5, Curcu- lionidae 7, Histeridae i, Cerambycidae 2, Staphylinidae 1, Nitidulidae 7, Anobiidae i) in addition to numerous other insects, and had it been a different season, stray species of Elateridae would certainly have occurred. I do not doubt I must often have exceeded this number of species, especially as on this occasion 1 was by no means well and really unfit to attempt collecting in the mountains. It has been suggested that close interbreeding may result in diminished fertility and in a paucity of individuals of island species, and that such interbreeding is very common there is no doubt. But whether in insects this inbreeding has any such result is extremely doubtful. We can feel quite sure that some of the introduced insects in the islands have originated from one or two, or very few individuals, accidentally imported by man, and of others imported for economic reasons we know this to be the case. Yet though some of the former were introduced at least half a century ago and have produced probably at least several broods a year ever since, and some of the latter complete a generation each month or even more quickly, these all remain as common as ever and show no sign whatever of degeneration or impaired fertility. Of imported insect pests it is sometimes said in a careless manner that these naturally disappear or become insignificant in time from being "played out," but 1 do not doubt that careful investigation will show that in such cases the real cause of decrease is that they become attacked, after a while, by other insects or diseases, and especially by parasites and predators, which transfer their attack by degrees from native species to the foreign immigrant. Deterioration in insects, supposed to be due to inbreeding, is much more probably due to the ill-effects of the unnatural conditions of captivity. The extreme sluggishness of so many of the insects and Mollusca leading to great restriction of locality, which we observe also in the birds, must lead to extreme inbreedino-. We have observed colonies of some of the fliohtless beetles to persist for years in a single tree and where these colonies are isolated from others, no doubt all the inhabitants have resulted from a few stray examples, probably often from one. While the food remains in suitable condition these isolated colonies thrive and become very numerous, but they are, doubtless, often totally exterminated when it becomes no longer suitable, unless, by chance, individuals can reach some other tree fit to supply a breeding ground. Obviously this sluggishness and restriction of range must itself diminish the numbers of individuals of a species, and the tendency of island creatures to limit their range and to specialize their habits is a striking- feature of the fauna. Loss of the powers of ilight is a very common phenomenon in this connection. INTRODUCTION xli Number of species of insects and causes of extinction. The total number of Hawaiian insects known to me at the present time is about 3325, but of these only about 2740 can be considered as belonging to the natural fauna. Owing to man's interference and his introduction of foreign creatures, I should consider that about 300 species have been exterminated, without being collected. Mr Blackburn's and my own collecting, together with some aid received from others, has in my opinion resulted in the collection of about half of the e.xisting species of native insects, so that a perfect collection of this section of the fauna, made before man's interference, might have yielded about 5780 species. I may have somewhat underestimated the number exterminated. It is known that several species of Passerine birds, formerly abundant, had become either totally or nearly extinct before I came to the islands, twenty years ago, and there is reason to fear that several others may since have disappeared. At any rate, we know certainly that some, which were abundant, have become much less so, and have totally disappeared from localities where they were common enough. In my paper on the birds (Vol. i, p. 393) I have referred to the causes which have brought about this extinction or diminution in numbers. As with the birds, destruction of forest has, doubtless, caused the disappearance of many local insects, but even of greater importance has been the introduction of foreign carnivorous species, especially of the dominant ant, Pheidole megacephala. There is no record of the time when this destruc- tive creature was imported, but even during the last twenty years it has occupied some considerable areas previously free from it. It may be said that no native Hawaiian Coleopterous insect can resist this predator, and it is practically useless to attempt to collect where it is well established. Just on the limits of its range one may occasionally meet with a few active beetles, e.g. species of Plagithmysus, often with these ants attached to their legs or bodies, but sooner or later they are quite ex- terminated from such localities. It is quite certain that native beetles and many other insects are absent from the localities occupied by Pheidole, solely on account of its presence. In several instances, as the ant has been observed to occupy a new area, this area having been collected over before it was present and yielding many native beetles, the latter have entirely disappeared. In a few low-lying localities, even close to the coast, there are some places, which from excessive dryness and other causes, the Pheidole is unable to occupy, at any rate permanently, and yet unfavourable, as these are, for insects of any kind, here only will native Coleoptera be found. On one occasion I came across an instructive instance of the effect of these ants on the native fauna. A more or less open piece of forest at an elevation of 1500 ft. above sea-level, with a large variety of trees scattered in it, appeared at first sight an excellent spot for collecting native insects. A number of native Hymenoptera F. H. I. / xlii FAUNA HAWAIIENSIS were seen flying round the foliage, hardy insects which the ants cannot exterminate, though they are often seen attached to them by the mandibles. Every tree trunk was invaded by Pheidole, and beating the boughs dislodged them in thousands. Not a single beetle nor any native insect was obtained from these trees. One solitary tree, however, for some reason was quite free from ants. It was a large Bobea, with hanging masses of ' Maile ' [Alyxia) dependent from the boughs. From the dead stems of this were shaken hundreds, if not thousands, of one species of Proterhinus, others also being present, as well as the large weevils, Rliyncogonus, and other kinds of beetles. I visited this spot on many occasions for the sake of a rare species of wasp, but never obtained a beetle except from this one tree, and a year later it too was occupied by Pheidole and barren of native insects. Fortunately Pheidole is not universal in its distribution. It can in some localities just attain 4000 ft. in the mountains, under certain climatic conditions. Below twelve or thirteen hundred feet it often occupies most of the islands, excepting some extremely arid localities. Though not so utterly destructive to other insects as to the beetles, yet many of them are destroyed by it, and generally speaking, collecting is very poor, where it abounds. Most of the native species taken in such places are vagrant, like Lepidoptera, and have bred in some adjoining area, either free from this ant, or where it is comparatively sparse. Miles of attractive forest in some parts of the islands are almost devoid of native insects, through its destructiveness. A very few endemic insects seem able to breed in its haunts, even where it is quite abundant, but many of the foreign or imported insects flourish in spite of it. It is not probable that it will spread to any great extent beyond the limits now occupied, for it has long since filled all suitable localities. Here and there the opening up of limited areas of forest may by change of conditions allow it to colonize these, but the great bulk of the forest is now reserved and not likely to be opened up. There is no reason to suppose that the endemic insect fauna will suffer any considerable further diminution, and it may, so far as one can see, remain as it is for ages to come. The chief danger would be in the introduction of some predaceous creature like Pheidole, which would be able to occupy the great area of forest land and the country above this, where Pheidole does not now exist. As no such insect has been imported in the course of the last century, it is on the whole improbable that it ever will be. Comparison of 'introduced,' 'immigrant' and 'endemic' insects etc. In a memoir on the Hawaiian Coleoptera, published in 1885, by Blackburn and Sharp, the beetles are divided into three classes : (i) 'introduced,' i.e. species imported by man, (2) ' immigrant,' i.e. species occurring elsewhere, but which have reached the islands by natural means, (3) 'autochthonous,' i.e. species peculiar to the islands, for which I have used the older and more often employed term, 'endemic' INTRODUCTION xliii Owing to our very incomplete knowledge of the insect fauna of the world, some of the introduced species and also some considered to be immigrant are not yet known outside the islands. The ant Prenolepis sharpii for instance, which was imported years ago in boxes of plants by the Commissioner of Agriculture of the islands, and soon became established and very common, was described from descendants of these im- ported examples. In judging whether an insect has been introduced by man, when the actual fact of its importation is unknown, I consider the following points. (i) If solely attached to foreign vegetation or parasitic or preying on imported creatures only, it is certainly ' introduced,' or (less probably) ' immigrant.' (2) If, being a beetle or Heteropterous bug, it is found within the range of certain foreign predaceous insects, it may in the case of the former be considered certainly and in the case of the latter most probably 'introduced' or possibly 'immigrant' In explanation of this I may state, that in the case of the truly endemic beetles, without exception, and usually of the Heteroptera, the area of distribution is absolutely fixed by the non-occurrence of imported predators. Owing to interference by man these predators may come to occupy new areas, and then from these too the above-mentioned endemic forms disappear. At first, in such cases, many of the endemic beetles will be found actually being attacked by these enemies, but subsequently they will disappear. Some insects, however, e.g. bees and wasps and some Lepidoptera, more or less successfully resist such predators, and though endemic, can occupy the same area, as also can others that rarely or never are attacked, e.g. certain Homoptera that excrete honey dew. (3) Doubtfully endemic insects may occur outside the range of foreign predators and mixed with many truly endemic ones, but usually they also range into the territory occupied by the former. If not, their habits should be considered and their food, and the probability of their being imported by man, but if they do so they are most likely ' imported ' or ' immigrant,' and surely so if Coleopterous. (4) Actual affinity to, or remoteness from outside forms must always be considered. (5) Whether the species is isolated from other members of the fauna, or whether one or more closely allied forms occur is most important. When several closely related species are found in the islands the probability of their being endemic is very great. Importations by man usually consist of very different insects and not of closely allied species, and the same would be likely to be the case with natural immi- grants. A careful consideration of such points in the case of any insect that is fairly well studied, will usually leave little doubt as to whether it is endemic or introduced. It is perhaps more difficult to distinguish the immigrants from either the endemic or the imported species. We must, however, except from this statement a number of species of known migratory powers, well-known species elsewhere, with great powers of /2 xliv FAUNA HAWAIIENSIS flight, and clearly natural immigrants. Excluding these, the number of immigrants is probably very small, for reasons hereafter given. Whereas the imported species are in most respects of little interest as compared with the immigrant and endemic, in one particular they are of considerable importance, especially such species as have been established in the islands for many years. House- hold pests, however, and some others, that we know have been imported again and again, are of no interest. It is probable that such introduced species as the Longicorn beetles Laoocheirus obsolehts and Astrvnus hirtus were established from a single importation of these species. They are evidently not frequently carried by man, for until recent years the former had not spread to the other islands and the latter has not, as far as is known, yet reached them. Yet the former is known to have become so plentiful, as to have been an injurious insect in Honolulu forty or fifty years ago, and the latter has been established at least for thirty years. Though for all these years inter-island traffic has been very heavy, and plants continually taken from one island, to another, and especially from Honolulu, where these beetles are common, yet the spread o{ Lagocheirus has been slow, and that oi Astrimus has not, so far as is known, occurred at all, as stated above. It is, therefore, very unlikely that either of these beetles has frequently or even occasionally been brought to the islands from outside countries, seeing that the traffic between these and Honolulu has been, until recently, compara- tively very infrequent. I suspect, too, that these beetles are likely to be more abundant with us than in their native countries. Owing to the equability of the climate, on account of which so many insects breed the year round, and often produce many broods in a year, many species have passed through very many generations since their intro- duction. Some of the imported creatures we know from observation complete their life-cycle in three weeks, and brood follows brood without cessation. Yet, when we examine examples of these species, we may well feel some astonishment at the fact that they quite resemble those found in their original home, even though this differs greatly in climate and otherwise, and they exhibit no particular variability. Some (e.g. certain Coccinellidae) notably variable in their native homes, appear to exhibit even less variation here than in their own country. Looking at the imported species as a whole, we must allow that these show no tendency to special variation, and there is not the least reason to suppose that the variability seen is greater than, or different from that, which they exhibit in their native homes. One is struck with the stability of specific characters, and a study of these imported insects as yet throws no light on the question as to whether variation arises suddenly or by slow degrees in a species. One would infer that much time is as a rule required for the appreciable modification of a stable species. The immigrant species are of two classes like the introduced ones: (i) those that are, doubtless, more or less frequent \m^m\