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Smithsonian Miscellaneous Collections SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 103. NUMBER 1 DISTRIBUTION AND VARIATION OF THE HAWAIIAN TREE SNAIL ACHATINELLA APEXFULVA DIXON IN THE KOOLAU RANGE, OAHU (With 12 Plates) BY D'ALTE A. WELCH (Publication 3684) CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION DECEMBER 16, 1942 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 103, NUMBER 1 DISTRIBUTION AND VARIATION OF THE HAWAIIAN TREE SNAIL ACHATINELLA APEXFULVA DIXON IN THE KOOLAU RANGE, OAHU (With 12 Plates) BY D'ALTE A. WELCH (Publication 3684) CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION DECEMBER 16, 1942 BALTIMORE, UD., 0. 8. A. DISTRIBUTION AND VARIATION OF THE HAWAIIAN TREE SNAIL ACHATINELLA APEXFULVA DIXON IN THE KOOLAU RANGE, OAHU By d'ALTE a. welch (With 12 Plates) CONTENTS Page Introduction i Scope of work i Place names of the Koolau Range 2 Climatological data and physiography g Table i. Annual precipitation records in the Koolau Range 10 Material and methods 12 Table 2. Data on material 15 Species concept 21 Taxonomy 23 Synopsis of the subspecies of Achatinclla apcxfulva Dixon 23 Group of A. a. ccstits Newcomb 28 Group of A. a. vittata Reeve 42 Group of A. a. pilshryi new subspecies 52 Group of A. a. turgida Newcomb 57 Group of A. a. polymorpha Gulick 91 Group of A. a. leiicorraphe Gulick loi Group of A. a. invini Pilsbry and Cooke 115 Group of A. a. coniformis Gulick 126 Group of A. a. lilacea Gulick 133 Group of A. a. apicata Newcomb , 146 Group of A. a. aloha Pilsbry and Cooke 160 Group of A. a. apexjulva Dixon 176 Conclusions 188 Summary 206 References 208 Explanation of plates 208 Plates 1-12 224 Index to sections, genera, species, and subspecies 225 Index to place names 229 INTRODUCTION SCOPE OF WORK In a previous paper (Welch, 1938) the species Achatinella mustelina is studied. The present paper continues the work on Hawaiian tree snails and deals with the species Achatinclla apcxfulva, also of the island of Oahu. Does the same variation or series of variations occur Smithsonian Miscellaneous Collections, Vol. 103, No. 1 2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. IO3 in A. apcxfulva of the Koolau Range as in A. uiusielina of the Waianae Mountains? During the 7 years I spent in Hawaii studying Achatinella I was greatly helped in acquiring a background for the present work by Dr. C. M. Cooke, Jr. Working under Dr. Cooke, I also learned many methods of studying and preparing material which greatly facilitated my work. I am also indebted to Dr. E. A. Andrews, Dr. Henry A. Pilsbry, Dr. H. B. Baker, and Dr. Ernst Mayr for criticisms and advice in the preparation of the manuscript. Dr. F. Ray- mond Fosberg was most helpful in identifying all genera of Hawaiian plant names given by various collectors of shells. I also wish to express appreciation to the following for various courtesies and favors : Dr. Paul Bartsch, W. J. Clench, W. T. Caiman, Col. A. J. Peile, J. R. le B. Tomlin, G. S. Robson, Dr. Paul Galstoff, Dr. Gilbert D. Harris,. Dr. C. E. McClung, Dr. Peter H. Buck, Dr. Herbert Gregory, the Trustees of the Bernice P. Bishop Museum, and the au- thorities of the United States National Museum. The gr®und work of this paper was done at the Bernice P. Bishop Museum in Honolulu, the writing of the paper was carried on at the United States Bureau of Fisheries at Woods Hole, the Zoological Laboratory of the University of Pennsylvania, the Zoological De- partment of the Johns Hopkins University, and the Zoological Department of Barat College, Lake Forest, 111. I wish to thank these institutions for granting me facilities for work. PLACE NAMES OF THE KOOLAU RANGE Most of the names of the valleys or gulches of the Koolau Range are well known, but the names of some have never appeared on a Government map. Other gulch names have not been known, and still other gulches, although their names are well known, contain large subgulches that need to be designated by some name in order that localities may be described exactly. Pilsbry and Cooke (1912-1914, p. 277) published a rough dia- grammatic sketch map of the northwestern half of the Koolau Range in which they included the new names Kalaikoa, Ahonui, Main Poamoho, Central Poamoho, West Poamoho, Kawaihalona, and Luapo. This map was compiled from field notes made by Irwin Spalding and is the first published map showing the approximate regions occupied by colonies of AcJiatinella. Many of these names do not exist on the advance sheets of the United States Geological Survey topographic sheets of Oahu, Hawaiian Islands, 1/20,000, which form the standard map I used for plotting all localities of NO. I ACHATINELLA APEXFULVA DIXON WELCH Achatiiiella and from which all maps published in this paper have been compiled. Therefore a considerable amount of work has been done to check the place names on the Pilsbry-Cooke map and to ascertain the place names of all valleys not yet published. I am indebted to the following people for help in this work: The Bishop Estate for access to maps, Miss Jane L. Winne for place ISLAND OF OAHU KaneoheT Luakaha Upper* Nuuanu Pali Maunawili PLANT ZONES AVWaimanolo A' MARITIME Luakaha Lower, A HAOLE KOA B GUAVA C KOA D OHIA E CLOUD [G. I.—Island of Oahu, showing the plant zones in Kipapa Gulch (after Hosaka) and the location of rain-gage stations in the Koolau Range. Ridges outlined in solid lines. names obtained from an old Hawaiian native to the district of Maunalua; Manuel Baptista, a former cattle ranger native to the districts of Kalauao and Waiawa; G. D. Robinson; William Holt, a former cattle ranger in the regions of Wahiawa and Paala; Benny Pulaa, a Hawaiian native to the districts of Kawailoa, Waimea, and the northern section of Oahu. The Koolau Range is divided into six large land grants or districts which are numbered A to F on figure 2, p. 4. Within these large SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. IO3 NO. I ACHATINELLA APEXFULVA DIXON WELCH 5 districts are smaller districts or subdistricts indicated by Roman numerals, which may include a single valley or gulch with the same name or several valleys with different names. The valleys or gulches within a subdistrict are numbered in Arabic. In the discussion of place names, all names not published on the United States Geological Survey topographic advance sheets of the island of Oahu will have in parentheses the names of the authorities after them. Main valleys or gulches are often subdivided by the author for convenience into north, south, east, west, or central. These main valley subdivisions may be broken up again into north or south branches. Figures i and 3-6 were drawn by Gordon Brett Littlepage from maps compiled by me. Figures 2, 7, 8 were drawn by me. All maps are compiled from the United States Geological Survey topographic advance sheets of the island of Oahu. Ridges on all maps are repre- sented by lines. Place names of the Koolau Range A. Honolulu District. I. Maunalua. I, ; Kealakipapa Valley 2, Mauuwaii Valley ; 3, Kalama Valley ; 4, ; Kamiloiki Valley ; 5, Kamilonui Valley 6, Hahaione Valley ; 7, Kaalakei Valley (Bishop Estate maps: J. Winne). II. Kuliouou. I, Kuliouou Gulch; la, East Branch (Welch); ib, West Branch (Welch) ; 2, Puu O Kona. III. Niu. I, East Niu Gulch ; 2, West Niu Gulch. (1-2 Welch.) IV. Wailupe. I, Kului Gulch; 2, Laulaupoe Gulch; 3, Wailupe Gulch. V. Waialae Iki. I, Kapakahi Gulch (in this paper Waialae Iki Gulch will be used in preference to Kapakahi Gulch). VI. Waialae Nui. I, Waialae Nui Gulch (spelled Waialaenui on U. S. Geol. Surv. topogr. advance sheets). VII. Palolo. I, Waiomao Stream ; 2, Pukele Stream. VIII. Manoa. I, Manoa Stream; 2, East Manoa Stream (Welch). IX. Makiki. (This name is not on the U. S. Geol. Surv. topogr. advance sheets, but it is a well-known name and has been in use by shell collectors for many years.) I, Maunalaha Stream; 2, Moleka Stream; 3, Kanealole Stream; 4, Kanaha Stream; 5, Puu Ohia (Tantalus). X. Pauoa. I, Pauoa Valley. 6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. IO3 XI. Nuuanu. I, Glen Ada; 2, Glen Elm; 3, Lulumahu Valley; 4, Nuuanu Stream; Glen ; Maole Stream Waio- 4a, Nuuanu Pali ; 5, Hillerbrand's 6, ; 7, lani Stream (Rook's Valley, Cooke) ; 8, Puu Konahuanui ; 9, Puu Lanihuli. (1-3, 5, place names given by Dr. Cooke to certain un- named places where shells were collected.) XII. Kapalama. I, Niuhelewai Stream ; 2, Kapalama Stream. XIII. Kalihi. I, Kamanaiki Stream ; 2, Kalihi Stream. XIV. Kahauiki. I, Kahauiki Stream. XV. Moanalua. I, Manaiki Stream; 2, Moanalua Stream. B. Ewa District. I. Halawa. I, South Halawa Stream; la, South Branch (Welch); ib, North Branch (Welch) ; 2, Central Halawa Stream (Welch) ; 3, North Halawa Stream. II. Aiea. 1, Aiea Stream. III. Kalauao. I, Kalauao Stream. IV. Waimalu. I, Hanaiki Stream (Bishop Estate maps) ; 2, Waimalu Stream; 2a, ; ; South Branch 2b, Central Branch 2c, North Branch ; 3, Punanani Gulch. (2a-2c Welch.) V. Waiau. I, South Waiau Gulch ; 2, North Waiau Gulch. (1-2 Welch.) VI. Waimano. I, Waimano Stream; la, South Waimano Stream; ib, South Central Waimano Stream; ic, Central Waimano Stream; id, North Central Waimano Stream; le, North Waimano Stream. (la-ie Welch.) la-ic are in the district of Waiau but for convenience have been considered part of the Waimano Stream system. VII. Manana. I, Manana Stream. VIII. Waiawa. I, Waiawa Stream; la, South Waiawa Stream (Welch); ib, North Waiawa Stream (Welch) ; ic, North Waiawa Stream, South Branch (Welch) ; id.
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