CCARABID BEETLES (INSECTA: COLEOPTERA: CARABIDAE) OF THE QUEEN CHARLOTTE ISLANDS, BRITISH COLUMBIA By David H. Kavanaugh \ a ie Weval Published by The California Academy of Sciences pe - a Memoirs of the California Academy of Sciences Number 16 Carabid Beetles (Insecta: Coleoptera: Carabidae) of the Queen Charlotte Islands, British Columbia Carabid Beetles (Insecta: Coleoptera: Carabidae) of the Queen Charlotte Islands, British Columbia By David H. Kavanaugh Department of Entomology, California Academy of Sciences Golden Gate Park, San Francisco, California 94118 Published by the California Academy of Sciences Natural History Museum & Aquarium San Francisco January 24, 1992 Memoirs of the California Academy of Sciences Number 16 SCIENTIFIC PUBLICATIONS COMMITTEE: Thomas F. Daniel, Scientific Editor Terrance M. Gosliner Tomio Iwamoto Tom Moritz Wojciech J. Pulawski Ann Senuta, Managing Editor © 1992 by the California Academy of Sciences, Golden Gate Park, San Francisco, CA 94118 All rights reserved. No part of this publication may be reproduced or transmitted in any form or by any means, electronic or mechanical, including photocopying, recording, or any information storage or retrieval system, without permission in writing from the publisher. Library of Congress Catalog Card Number 91-058794 ISBN 0-940228-17-3 Cover illustration: Nebria charlottae. Mlustration by Mary Ann Tenorio. Table of Contents J NUS eae PE CER a ese RSDP I SR 1 RR SUNT ee a ae ee re ee ee ATE ee NE SOE RN EPS a ceel Doel or 1 VEU AeA Ye M1 YeR 0 CU) (ai ese ep te ee EID sR Sa ero En RE Eater eit ea SEU ee ee Brief History of Geologic and Biotic Studies in the Archipelago... ee ) Carabid Beetles and Their Usefulness in Biogeography and Palececolosys 6 History of Study of the Carabidae of the Archipelago: A Context for the Present Report. 6 Materials and Methods 0 ccccccccvccececne-neeeneseeeennnnnnnnnninnininnnnnnnnnsisssite nnn 7 Collection and Preservation of Specimens 7 Collecting Sites and Techniques... 8 Collecting Equipment 8 Preservation of Specimens 8 Logistics... 8 Cron and Habitat Venueiee. 9 Islands:Sampléd=. =... 9 Locality Information 22. 9 Coverage and Classification of Habitats 9 Formats for Taxonomic Accounts 29 Carabidae of the Queen Charlotte Islands... 31 A Key for Identification of Adult Carabidae from the nee Charlotte Islands . 31 Systematic Accounts for Tribes, Genera, and Species..... 36 Arrangement of Taxa 36 Tribe Cicindelini See 36 ae Gndea. freaaeus : 36 . Cicindela oregona Peconte Seneca 36 Tribe Ce POR ce NTE eee 40 eae Scaphinotus Latreille.. Per 40 . Scaphinotus Hanan (Misnaerheind) eee 41 3. SCAPNINOTUS MAL ZINALUS (FUSCHEL) occa socseeeeoeeneeeceeneeneteesessene 43 Genus Cychrus Fabricius. sates Paton 48 4. Cychrus tuberculatus Hone: : 48 Tribe Carabini... ae a a EA a 49 Genus Carabus Linnaeus... : 51 5. Carabus nemoralis Miller. ee 51 6. Carabus taedatus Fabricius... 52 Monibe NGDraM i 2. see ea cee ee eee 53 Genus: Leistus Frohlich)2.2.03.2echcassscacs: z BE} 7. Leistus ferruginosus Mannerheim 53 Genus Nebria Latreille : 55 8. Nebria charlottae Lindroth 55 9. Nebria louiseae Kavanaugh .0 7 55 10. Nebria haida Kavanaugh .......... ee eg ee Ie 56 11. Nebria sahlbergit sah ber git Fas Ver ieee eee cee sence neenetncennnn 56 12. Nebria rranmer Retr it Faschee reece a Bg oe ene ere es 56 13. GUNCDFI IVER SA: TC COM rasan cates epoe e Pare ree eater nes 57 Wribe No trp aya os a ccc rem : 57 Genus Notiophilus Dimer D1, 14. Notiophilus sylvaticus Eschscholtz 000... 57 bribe: Blairs see cence 57 Genus Elaphrus Fabricius 57 152. Elaphireisclatrviller Wat iy cn acc ee eee 57 16. Elaphrus americanus sylvanus Goulet. 58 Tribe Loricerini... 4’. Mandible with a setiferous puncture in scrobe (Fig. 34B) ... (tribe Nebriini) 9 5(4). | Metacoxae contiguous in ventral middle of body (Fig. 35A)... (tribe Carabini) 6 De Metacoxae separated in ventral middle of body (Fig. 35B) ... (tribe Cychrini) i 6(5). Upper surface of elytron and pronotum with faint or marked brassy, copperish, or blue metallic reflec- tion 5. Carabus nemoralis Miller 6'. Upper surface of elytron and pronotum without me- tallic reflection, black or rufous brown 6. Carabus taedatus Fabricius Labrum with four setae on anterior margin (Fig. 36A); elytron dull black, with contrasting large, shiny tu- bercles 4. Cychrus tuberculatus Harris The Labrum with six setae on anterior margin (Fig. 36B); elytron rufous-brown or black, without contrasting tubercles 8 7(5’'). 8(7'). 9(4'). 10(9’). 10’. 11(10’). 12(11’). 13(12'), 1S": 31 Elytron rufous-brown, without metallic reflection, with 18 regular, punctate striae . 2. Scaphinotus angusticollis (Mannerheim) Elytron black, with faint or marked metallic violet or green reflection, with 16 or fewer striae (irregular and difficult to count, especially laterally, in some specimens) 3. Scaphinotus marginatus (Fischer) Lateroventral margin of maxilla with several long, finger-like processes, each with a long, stout seta in- serted apically (Fig. 37A) 7. Leistus ferruginosus Mannerheim Lateroventral margin of maxilla without finger-like processes, setae inserted directly on ventral maxil- lary surface (Fig. 37B) 10 Pronotum (Fig. 38A) with basal angles obtuse (slight- ly denticulate in some individuals) AND lateral ex- planation narrow, at least at middle, basal sinuation of lateral margin shallow or absent; hindwings full- sized 12. Nebria mannerheimii Fischer Pronotum (Fig. 38 B—F) with basal angles rectangular or slightly acute, lateral explanation narrow or broad at middle, basal sinuation of lateral margin shallow or deep; hindwing full-sized or short (truncate distal to stigma or also narrowed) 11 Body pale yellow or tan; pronotum (Fig. 38B) very broad, markedly narrowed basally (posteriorly), lat- eral explanation narrow at middle; elytral silhouette (Fig. 39A) markedly ovoid, markedly narrowed ba- sally (anteriorly); hindwing short and narrowed (Fig. 40A) 13. Nebria diversa LeConte Body dark rufopiceous or black; pronotum (Fig. 38C— F) narrower, less markedly narrowed basally (pos- teriorly), lateral explanation broad, even at middle; elytral silhouette (Fig. 39B-E) rectangular, sub- rectangular, or subovoid, not or moderately nar- rowed basally (anteriorly); hindwing full-sized (Fig. 40C) or short (truncate distal to stigma) (Fig. 40B) 12 Head (Fig. 38C) moderate in relative size and width; pronotum (Fig. 38C) shorter, relatively narrow an- teriorly and broad basally; elytral silhouette (Fig. 39B) subrectangular, only slightly narrowed basally, elytral striae deeply punctate; hindwing full-sized (Fig. 40C) 11. Nebria sahlbergii sahlbergii Fischer Head (Fig. 38D) relatively large and wide; pronotum (Fig. 38D—F) longer, broader anteriorly and nar- rowed basally; elytral silhouette (Fig. 39C—E) sub- ovoid, distinctly narrowed basally, elytral striae faintly punctate; hindwing short, truncate distal to stigma (Fig. 40B) 13 Pronotum (Fig. 38F) short, wide, basal sinuation of lateral margin short, very deep; elytra (Fig. 39D) with silhouette relatively short and broad, intervals flat; legs relatively short 8. Nebria charlottae Lindroth Pronotum (Fig. 38D and 38E) longer, more slender, basal sinuation of lateral margin longer, moderately deep; elytra (Fig. 39C and 39E) with silhouette longer and more slender, intervals slightly or markedly con- vex; legs longer, more slender 14 CALIFORNIA ACADEMY OF SCIENCES Ipm2 \ sos : \ prbg prbf prbs prbm__Prba KAVANAUGH—CARABID BEETLES OF THE QUEEN CHARLOTTE ISLANDS 33 14(13’). Pronotum (Fig. 38D) with apical angles short, rela- in size; frons without distinct carinae between eyes tively broad and slightly rounded; elytral silhouette (or, if present, then carinae convergent anteriorly) (Fig. 39C) very narrow; femora and tibiae piceous —_ 20 or black; habitat above treeline in alpine zone 20(19’). Mandible with a setiferous puncture in scrobe (Fig. 10. Nebria haida Kavanaugh 45A) (OR if scrobal seta absent, then body with dis- 14’. Proncnim (Fie. 38E) with apical angles longer, rel- tinct metallic purple reflection); antennomeres 5-11 atively narrow and pointed; elytral silhouette (Fig. covered with short, dense setae, antennomere 3 (Fig. 39E) less narrowed; femora piceous, tibiae rufous or 46A) glabrous except for long setae at distal apex, rufopiceous; habitat upper sea beach area ...... oe antennomere 4 with short, dense setae restricted to ee er _......9. Nebria louiseae Kavanaugh distal one-half or absent; pronotum narrowed ba- 15(3’). Mandible with a setiferous puncture in scrobe (Fig. sally, body therefore slightly pedunculate in shape 41A); elytron with three rows of large, dull, ocellate . (tribe Broscini) ; 21 foveae ... (tribe Elaphrini) 16 20’. Mandible without a seliferous puneture in scrobe LS: Mandible without a setiferous puncture in “serobe (Fig. 47B) (and body without distinct metallic purple (Fig. 41 B); elytron without large, dull, ocellate foveae reflection); antennomeres 4—11 covered with short, (or if suggested, then foveae shiny) - 17 dense setae, antennomere 3 (Fig. 46B) also with short 16(15). Frons with a deep, finely and densely punctate pit dense setae, at least on distal one-third; pronotum in midline between eyes, pit surrounded by a shiny not or only slightly narrowed basally, body not pe- raised area with very sparse punctures (punctures dunculate in shape . . . (tribe Harpalini) 22 separated by gaps of more than twice their diameter) 21(20). Body length 12.0 mm or more; body with distinct . 15. Elaphrus clairvillei Kirby (brilliant in some individuals) metallic purple re- 16’. Frons without a pit in midline between eyes (or, if flection (some individuals with brassy and/or copper faintly present, then pit surrounded by a flat or only highlights) 20. Zacotus matthewsi LeConte slightly raised area with dense punctures (punctures, Diliss Body length less than 10.5 mm; body without me- on average, about one diameter apart) tallic reflection nen Nene eee Fee 16. Elaphrus americanus sylvanus Goulet 19. Broscodera insignis (Mannerheim) 17(15'). Body cylindrical, pedunculate (markedly constricted 22(20'). Pronotum (Fig. 47A) with basolateral setae present; between pronotum and elytra); antennomeres 24 elytron with numerous short, scattered setae laterally without long, erect setae (Fig. 42A). . . (tribe Scariti- (on eighth and ninth intervals) and apically ni) 18. Dyschirius pacificus Lindroth 61. Trichocellus cognatus (Gyllenhal) 1 Body dorsoventrally flattened, not pedunculate (not Di Pronotum (Fig. 47B) with basolateral setae absent; markedly constricted between pronotum and elytra), elytron without short, scattered setae laterally or api- antennomeres 2-4 with several very long, erect setae cally (eighth and ninth intervals bare except for um- (Fig. 42B) .. . (tribe Loricerini) bilicate series of setae in eighth stria and on ninth _7. Loricera decempunctata Eschscholtz interval) 23 18(2'). Head with one supraorbital setiferous puncture dor- 23(22'). Penultimate labial palpomere Bisex antesorly (Fig. somedial to each eye (Fig. 43A) 19 48A); scutellar striole absent or very short (Fig. 49A) 18’. Head with two supraorbital setiferous punctures dor- 62. Bradycellus nigrinus (Dejean) somedial to each eye (Fig. 43B) 24 19(18). Procoxal cavity open behind, not enclosed by pro- pleuron posteriorly (Fig. 44A); eyes very large; frons with several parallel, longitudinal carinae between clearly delineated (Fig. 49B) . eyes ... (tribe Notiophilini) 60. Harpalus somnulentus Dejean 14. Notiophilus sylvaticus Eschscholtz 24(18'). Terminal maxillary palpomere small, narrow and 19% Procoxal cavity (Fig. 44B) entirely enclosed by pro- less than one-half the length of the penultimate pal- pleuron posteriorly; eyes small, moderate, or large pomere (Fig. 50A) . . . (tribe Bembidiini) 25 23". Penultimate labial palpomere plurisetose anteriorly (Fig. 49B); scutellar striole present, relatively long, = Ficure 30. General structure of a carabid beetle (Coleoptera: Carabidae). A, general appearance, dorsal view; B, general appearance, ventral view; C, head, dorsal view; D, left mandible, dorsal view; E, left maxilla, ventral view; F, labium, ventral view; G, pronotum, dorsal view; H, left elytron, dorsal view; I, right metathoracic (hind) leg, dorsal view. Abbreviations: an = antenna; anl = scape (first antennomere); an2 = pedicel (second antennomere), an3—anl1 = third through eleventh antennomeres; bl = body length; cl = clypeus; el = elytron; elbm = basal margin of elytron; elds = discal setae of elytron; elep = epipleuron of elytron; elhu = humerus of elytron; elil, eli2, etc. = first and subsequent intervals of elytron; ellm = lateral margination of elytron; elmm = medial margin of elytron; elss = scutellar striole of elytron; els, els2, etc. = first and subsequent striae of elytron; elus = umbilicate setae of elytron; ey = compound eye; fe = femur; fr = frons; frf = frontal furrows; Im = labium; Imp = labial palpus; Imp2 = penultimate palpomere of labial palpus; Ir = labrum; mics = preferred area for comparisons of elytral microsculpture; mn = mandible; mns = scrobe of mandible; mnss = seta in scrobe of mandible; mscc = mesocoxal cavity (i.e., cavity into which mesocoxa is inserted), msem = mesepimeron; mses = mesepisternum; mss = mesosternum; mtc = metacoxa; mtes = metepisternum; mts = metasternum; mtsp = metasterr 4 - - - - - - - - - - - - - - - - I ae ‘s a = = = 2 eS 2 = a - _ _ = = _ = I - = x - - - - - - - - - - - - - - - - Z - = x - - - - x - - - = - - - - - - - ci - - x - - x - x x x % x x - - - x x x I = - &% = = i = = = = 7 - - = = = = - = I - - x - - - - - - - - - - - - - - - - WiayiouuRYy wunjDjoaaofiaponb uoipiquiag Ays[nyosi0op; Winjaoul UoIpiquag U0 DaT SudISapldl UOIpIqMag Aased Uin]SD) UOoIpIquiag aUu0Da] WnULIYISAp UoIpiquiag Aase) sdaa1ndo ajonbapul uoipiquiag [ey uaudydaz voipiquag uersfad snaqdjpy? snYyIad]. AY¥S[NYISIOPY smuuadiao snyIad | uosyouy ssnjgo snyoas uealag snudiiaip snojdiq ANUODI] WMayIOU SNJOIDZ wiayiouuRYy SIUSISUL DABPOISOL yloipury snoyiond sn1yostq Zyoyosyosy DIDIIUNAW IIAP DAINOT AQAryy aj/aaipjo snaydnp y JQINOH snupapas SNUDIAIUID SNAYAD] ZYOYsYyosy snopayds snjtydouon IUODY] VSMaalp SUGIN J9yosty MUMIYIIUUDUL DLUGON Joyost{4 NBLIG] YDS US1IG]YOS DUGAN ysneueary opipy Dugan ysneueary avasino] DUgIN YLOIPULT AP}O/ADYI DUGAN wiraysauuRy SNSOULSNALAL SAISIAT sniouge, smjopan} snqvav.) JOIINAL SjPaouiau snqoiv.) sLueH smypjno4aqn) Sn4YyIAD 4oyosi4 snjouisapud snjoulydnog wWtaysouUeyy syjoousnsun snjourydvog MUAY] Vuosaso0 Yjapuiiy d# OT VH YD GW HO IH MS OT AM TL NL AX AT WS YW SH VY NH NA OA OW NV LX z' {PURIST uOXE “SGNVIST JLLOTYWH) NddNO AHL NI SalOadS GIAVAVD 4O SNOLLNAITALSIG TWOIHd Va OOD) “p aTav 81 KAVANAUGH—CARABID BEETLES OF THE QUEEN CHARLOTTE ISLANDS Chet =ta Ba w w sSBUIM pda «P *P aa? nN «Co «?P aaD -— Ne | | * i | fon | | x | ' oz =x - x x x 7x - x - x d# OT VH YO AW HO = cs = = = s = = as = = - = FS = x = = = = = = = = = = = = = “ = = Ss x i = = = = = = = = = = = = = = = = x = = - - x - - x x x - - - - - - - x - - - - x x - x x x - - x x x x - x - - - - x x bd x x x x x x x x x x x x x - - x x x x - x - - - x x x - x - - - - x x - x x x - - - - - - - x - - = = = = x = z = = = = = = = = = = = a = = = = = 2 = - = = = = = a x = = - - - - - - x x - - - - x - - x - - = = = = = = = 2 S = # x = = ‘= x = = oo = = = = = i = a = = = = = = x = = = = = = = = = = = = = = 2 = = x = = = = = S = = = ey = = a = = = _ x = i) = = = = - = = = = = Se = = = = x = = e = = 2 = = = = x = = 2 = = = = x = = = = x = = as = = & es = = _ = a x - - = = = = a = = = = = = = = = = x = = = = x = = = = = = = = = = = = x = = Aased Sisdijja aD AQsed DSONUIS DAD uvalag ajjoomadq WnUuos a1U0D7] SUudISA]]DIAU WNUOS F yey lajjaq wnuosp uealag UNSOUISNALA, WUNUOS uealoq SNiUDdl SNYINSOLII Zyoyosyosq SNJIAJSPO SNYINSOSI ueroloq SNIUDISDI SHYIISOLA WioysiouURW, SNUITISAYJIUUD SNYINSOLI 21U0D2] SNPIB]D SNYINSOIA aoa SIJJOMUILI SNYIUNSOLI) SguIQUI DUD] SNYINSOLA uvalag snapusdd sniuoN wioaysiouuRey uinjnsuojgo uoipiquiag yloipury sapiojnsuojgo uoipiquiag wiaysiouuRE, ajiqvjioads uoipiquag Ays[nyosijow UiNjD14]SA}40f UOIpIquiag 31U0D 7] AOJOISAIA UOIPIQUeag uealaq WNJOUIISIPUL UOIpIquiag 310097 WN]DUAdIUI UOIpIqMiag uealaqd a] DS4aASUDA] UOIPIQuiag [ey Wnaunpiuas uinjounias uoIpIquag wioysiouuReW, wunjnosniunjd uoipiquiag wioysiouuRy] unjnuvjduoo uoipiquiag Aasea 40]D1A uoIpIquiag yey avipiip{ uoipiquiag IH MS OT AY TL NL AM AT VA UW SH VA NH NA OF OW NV IN zy PUeIsT "GANNILNOD “pf ITAV uOXe CALIFORNIA ACADEMY OF SCIENCES “odeadiysie ayi Ul Puno} sjenptatput snosajdAyoeig pure snosajdoioew y10q ‘stydioulIp = p ‘sdulm pamoueu JO/pue pausyoys ‘snoiajdAyoesq = q ‘s8UIM pazis-|[ny ‘snoja}doioewW = Wi :sBuLMpuTY ay] JO UOTIPUOD = sBUIAA ¢ ‘YSBOD PURTUTEUT JUDIE(Pe oY} 0} PalouIsas OFR[adryose ayi apisino sues = x, “SPuRIS] MOLeYD udsand ay) 1e adueI [eorydes30I3 UMOUY Jo ITT] wiayuiou = “BOWUTW YON 0} p2onposUy] = 9g ‘o1Mapuy = ¢ ‘[PISPOD 1SaA\ = p ‘URILIDUTY YUION WiajsaM = € ‘URDUAWISUBIY = 7 SONdse[OH = | :satads ay Joy Waned adues [eorydess0a3 = qu y ‘Ppapsodas U9aq sey satdads ay) YSIYM WO SpuUr]sI JO JaquINu = TH ¢ “odejadiyoie ay ut jounnxa mou Apqeqgold satsads 1nq ‘pues! Uo pajda][09 suswtdads Wo UMOUY = (x) {PUR]SI UO IN390 01 UMOUY JOU = — ‘puURIsT UO 1N990 07 umoUy = X :odejadryose ayi ul uonnquasiq ; AUeL = NY ‘uRMyUNTeL, = TL ‘suepays isaqq = HS Jood = dy ‘Aeswey = yy ‘wosiyoinw = YW ‘Aqsxa0w = OW ‘opneW = GW ‘[1947 = AT ‘asinoq = OF ‘euedueq = OT esuny = ny ‘uysuny = Lo ‘AexNH = NH “8uudsioy = SH ‘usqqrY = [H “‘uOsueH = YH “weyeIn = yD ‘Aepeiey = Wy ‘saddod 1seq OF ‘NeeyD = HO ‘Aqeuing = ng ‘Auoyluy = Ny ‘spurys] I * id 2S I P I i Oo 9 9 Ot 8 FI I I £ 9 iS 0. I Or I P Purjsr tod sje1o wi G - = x - = - - = - - = - - = - - ces - - fe x - = uealaq SNULSIU SN]JIIAPDAG uw I I - a x a = = a = = = = = = 7 7 ad 7 = Ss ™ = 7 y feyuay|AD SNIDUSOI SNJJIIOYIU J w G. OZ - - x - - - - - - - = - - = - - = - - = x - = urafoq SNJuanNUWOS SnyDdsv ET w Z € - = x - - - - - - - = - x - - - - - - - x - - WwIaysJouURYy S1]DI0}11] DAD S8uIM dd c# OT VH YO GW HO IH MS OT 4AM TL NI AN AT V4 UW SH VU AH AA OFT OW NV LM uoxel 7:1 PURIST “GANNILNOD “p AJTaV KAVANAUGH—CARABID BEETLES OF THE QUEEN CHARLOTTE ISLANDS 83 weevils (Anderson 1988), and carabids are illustrated in Figure 89. The vascular plant flora and dragonfly fauna include a large Holarctic component (38% and 23% of the flora and fauna, respectively), whereas the blackfly, weevil, and carabid faunas have only a minor Holarctic representation (only 5 or 6% each). Among dragonflies, the Transamerican element is dominant (70% of the fauna), the Western North American element small (only 7%), and more restricted coastal elements absent. Both blackfly and carabid faunas include diverse Transamerican el- ements (31% and 19% of the faunas, respectively) but vascular plants and weevils do not (9% and 5%, respectively). The dom- inant element (44% of the species) in the blackfly fauna is the Western North American component, which is also very well represented in the carabid and weevil faunas (29% and 21% of the species, respectively). The latter two faunas are also similar in having the West Coastal element dominant (58% of the weevil and 40% of the carabid faunas). Among the groups compared here, only the vascular plant flora and carabid fauna include an endemic element at the species level (one and two species, 0.02% of the flora and 3% of the fauna, respectively), although endemic subspecies of mammals, birds, and plants also have been de- scribed. The vascular plant flora also includes a diverse (about 20% of the flora) Introduced element. This element is also rep- resented, by two species each, in the weevil and carabid faunas. Habitat Distributions of Carabids Within the Archipelago Table 6 lists the known habitat distributions of carabid species in the Queen Charlotte Islands. Differences between data pre- sented by Kavanaugh (1989, Table 2) and those provided here reflect both additional sampling and slight changes in the habitat classification itself (see Methods and Materials section for dis- cussion of habitat types). Habitat distributions for two of the 62 carabid species in the fauna of the archipelago, Scaphinotus angusticollis and Bembidion dyschirinum, remain unknown. As noted above in the species account and elsewhere, S. angusti- collis has probably become extinct in the archipelago. Additional sampling in all habitats, particularly in the Masset area, may yet produce specimens of B. dyschirinum and help to identify the habitat requirements of members of that species in the ar- chipelago. Carabid species diversity in relation to habitat type is sum- marized in Table 7 and illustrated in Figure 90. The fauna of the Queen Charlotte Islands is notable for its overall balance in diversity among the 15 habitat types recognized. Habitats with the highest diversity of carabid species are the lowland decid- uous forest and lowland, small, shaded stream shore types (15 species each), followed closely by lowland marsh or bog, lowland coniferous forest, and lowland open/synanthropic habitats (with 14, 13, and 13 species each, respectively). The least diverse habitats are the upland lake or stream shore and subalpine forest types (with five species each), followed by the two upper sea beach habitats (with six species each). If the basic habitat types are grouped into beach, open ground, forest, wetland, and alpine habitat complexes (Table 7), then high species diversity in wet- land (a total of 35 species, 58.3% of the fauna) and forest (20 species, 33.3% of the fauna) habitats is more readily apparent. In the carabid beetle faunas of most areas studied to date, species diversity is highest in lowland and/or lower montane habitats and decreases steadily with increased elevation (e.g., see Armin Tas_e 5. GEOGRAPHICAL DISTRIBUTIONS OF QUEEN CHARLOTTE ISLANDS CAR- ABID SPECIES IN RELATION TO GENERALIZED RANGE PATTERNS. Range pattern No. spp. NL! Holarctic 3 0 Transamerican 12 4 Western North American 18 0 West Coastal 25 Endemic 2 n/a Introduced 2 ' NL = number of species that reach their northern distributional limit at the Queen Charlotte Islands. 1963; Darlington 1943, 1971; Greenslade 1968). This seems to be true also for the fauna of the Queen Charlotte Islands. If the basic habitat types are regrouped as either lowland or upland habitat complexes, then higher diversity of the lowland is ob- vious. Fifty-four species (90.0% of the fauna) occur in one or more lowland habitats, while 29 species (48.3% of the fauna) live in at least one upland habitat. Although species diversity is distributed broadly among the different habitat types, carabid species nonetheless exhibit marked habitat specificity in the archipelago (Fig. 91). Twenty- four species (40.0% of the fauna) are apparently restricted to a single habitat type, and 65% of the species occur in only one or two habitats. Less than 12% of the species occupy more than four of the 15 habitat types recognized. At present, much less is known about the habitat distributions of most of the 60 non- endemic carabid species outside of the archipelago than within it. This is indeed unfortunate because equivalent knowledge of the habitat distributions of these same species on the mainland could provide data needed for meaningful tests of such concepts as “competitive release” (MacArthur 1972) in island popula- tions, or conversely, competitive restriction in mainland pop- ulations. Diamond’s (1970) study of the habitat distributions of birds on Karkar Island in relation to their distributions on ‘mainland’ New Guinea provides a classic example of such a test. The only carabid species for which such data are now available for populations both on the adjacent mainland and in the archipelago are the four non-endemic Nebria species. For each of these four species (Nebria haida, N. sahlbergii sahlbergii, N. diversa, and N. mannerheimit) habitat distributions of main- land and islandic populations are identical, even though main- land populations coexist with populations of at least six other congeneric species (Nebria crassicornis Van Dyke, Nebria acuta Lindroth, Nebria gebleri Dejean, Nebria kincaidi Schwarz, Ne- bria meanyi Van Dyke, and Nebria piperi Van Dyke), as well as with other potential competitors. These data suggest that (1) “competitive release” has not occurred for Nebria species in the archipelago, (2) habitat distributions of these same species on the mainland are not restricted by competition, or, most likely, (3) competition plays little or no role in restricting habitat dis- tribution in either mainland or insular populations. To what extent competition affects the habitat distributions of other spe- cies in the archipelago remains unknown. Table 8 illustrates relationships between geographical and habitat distributions and carabid diversity. Several patterns are readily apparent. The only faunal element represented in all habitat types 1s the West Coastal element, which is also the most diverse element in all but the lowland large lake or stream shore 84 and upland lake or steam shore habitats. No one habitat type includes representatives of all six faunal elements. Holarctic and Introduced elements are restricted to lowland habitats, the latter to the immediate vicinity of human habitation or environmental disturbance. Transamerican elements are also restricted to low- land habitats, except for their occurrence (six species) in the upland marsh or bog habitat. The Endemic element is restricted to the cobble upper sea beach habitat type. Finally, the fauna of the sandy upper sea beach habitat type includes six species, all of which belong to the West Coastal faunal element. Hindwing Development and Carabid Distributions Table 4 records the degree of development of hindwings in adults of each of the 62 carabid species represented in the Queen Charlotte Islands, based on examination of all available speci- mens of each species. All adults of 38 species (61% of the fauna) have fully developed (i.e., macropterous) hindwings; 22 species (36%) have all adults with hindwings reduced in size (1.e., bra- chypterous), and 2 species (3%) are dimorphic in this regard (1.e., both macropterous and brachypterous adults occur in popula- tions in the archipelago). Without exception, condition of the hindwings for members of each species in the archipelago 1s typical for that species throughout its geographical range. There is no evidence, for example, of the development of brachyptery within the archipelago for any species in which adults elsewhere are all macropterous; nor is there any indication that, in di- morphic species, either macropterous or brachypterous indi- viduals are at any particular (selective) advantage in the archi- pelago. Intuitively, one would expect that, all other factors being equal, organisms that can walk and fly should be able to disperse more efficiently and extensively than those that can only walk. Also, they should be able to cross at least some barriers, areas of unsuitable habitat (such as bodies of water), that walking in- dividuals cannot cross. By extension, species that have individ- uals capable of flight should be able to establish and maintain larger geographical ranges, and perhaps also larger habitat rang- es, than species with only flightless members. To the extent that such relationships between flight capability and distribution can be expected, what is observed is the carabid fauna of the Queen Charlotte Islands is in some ways surprising. For example, Fig- ure 81 illustrates the relationship between hindwing develop- ment and the number of islands in the archipelago known to be occupied by a species. All members of the seven most wide- ranging species (i.e., all those that have been recorded from seven or more islands in the archipelago) are brachypterous, not macropterous. Hence there is no evidence to suggest brachyptery as a significant factor in limiting geographical distribution within the archipelago. However, the observed relationship between hindwing development and overall geographical distribution, as illustrated in Figure 92, is as predicted. Adults of those species with widest geographical distributions (i.e., all the Holarctic species, 92% of the Transamerican species, and 67% of the Western North American species) are macropterous, whereas 50% of the West Coastal and both of the Endemic species have brachypterous adults. The relationship between hindwing development and habitat in the fauna of the archipelago is generally similar to that found in other faunas studied to date (Erwin 1979). The highest in- cidence of brachyptery is in forest habitats, whether lowland CALIFORNIA ACADEMY OF SCIENCES (67-85% of species represented in the habitat) or upland (88— 91%), deciduous (67-88%) or coniferous (85-91%). The lowest incidence of brachyptery (and highest of macroptery) is in wa- terside habitats (0-27%), whether marsh or bog (7—18%) or lake or stream shore (0O—27%). Whereas 33% of species in the sandy upper sea beach habitat have brachypterous members, 67% of those in the cobble sea beach habitat have short-winged adults. The relationship between habitat range and hindwing devel- opment, as seen in Figure 91, 1s similar to that for geographical range within the archipelago. The four species that occupy the broadest habitat ranges (more than seven habitats each) all have brachypterous adults. Hence, there is no evidence to suggest brachyptery as a factor limiting the habitat ranges of species in the archipelago. It is clear that carabid species with only brachypterous adults nonetheless may be highly successful in establishing and, pre- sumably, maintaining broad geographical and habitat ranges within the archipelago. In fact, some of these species actually appear to be more successful in this regard than those with macropterous adults. How can this be? What, if anything, is wrong with the initial predictions concerning relationships be- tween hindwing development and distribution? Nelemans (1987) has shown that wing size alone may not be a reliable indicator of flight capability. He found that, at least in Nebria brevicollis Fabricius, all adults of which are macropterous, functional wing muscles develop only in adults that were well fed as larvae. While sufficient larval food could be provided under laboratory conditions to induce subsequent flight muscle development in most reared adults, few larvae appear to receive even the min- imum requisite food in the wild to trigger development of mus- cles in the adult stage. The incidence of functional flight muscles among adults in the wild is very low. Darlington (1936, 1943) and Kavanaugh (1985) have discussed the suite of other struc- tural adaptations, such as shortening of the metathorax and narrowing of the elytral base, that frequently accompany bra- chyptery per se among carabids. As a result of these associated structural modifications, brachypterous beetles are often far more efficient walking and running organisms than were their flying ancestors. If, in fact, flight muscles rarely if ever develop among macropterous carabids in the Queen Charlotte Islands, then these beetles may actually be poorer dispersers than their bra- chypterous equivalents. Clearly, studies of flight muscle devel- opment among the macropterous adults of species in the ar- chipelago are needed to ascertain their true flight capability. If these beetles are functionally flightless, then their relatively re- stricted distributions do not, in fact, negate the original predic- tions. Affinities of the Carabid Fauna Inventory of the carabid species of the Pacific Northwest coastal region and of the distributions of these species within the region is far from complete. The only areas along the coast that have been intensively studied to date are Kodiak Island (Lindroth 1969), the Lituya Bay region of southeastern Alaska (D. H. Mann and Kavanaugh unpubl.), and the Queen Charlotte Is- lands (Kavanaugh 1989 and present report). The known faunas of other coastal areas from northwestern Washington to south- western Alaska represent unknown fractions of the total number of species actually represented in each area. Nonetheless, enough KAVANAUGH—CARABID BEETLES OF THE QUEEN CHARLOTTE ISLANDS Q A) iN Ss d) Ls ROKK SORE EX OK Oe y, Y Elaphrus clairvillei Kirb p y NJ Amara littoralis Mannerheim Ficure 83. Geographical distributions. Transamerican Range Pattern. 85 CALIFORNIA ACADEMY OF SCIENCES 86 Loricera decempunctata Eschscholtz . Bembidion viator Casey ZG Scaphinotus marginatus Fischer dd @ 2 Cc oO @® @ P| w” > @ << = = oOo £ wn a 2 fo] oO oO NI 87 KAVANAUGH—CARABID BEETLES OF THE QUEEN CHARLOTTE ISLANDS SO ONAN MH HMB AIONMNHMNAANA THI PFHtentOnadt(nrHeonrnnrnnnaa sieiqey ‘ON cad | | | “~ « 6 e oe io i 7:p9dAl 1e11qeH “SAGNVIS]T JLLOTAYVH) NEEISTO) AHL NI SATLAIA GIAVAYVO AO SNOILNAIMLSIG LVLIGVHE 9 ATAV WlaysJauURY] WnjnuDnjdiuos uoipiquiag Aase_ 40jD1A uoIpiquiag yey aviv.v{ uoipiquag WwiaysJauuRy] lunjpjoaaofiuponb uoipiquiag AYS|NYISIOW WNjWaIUI UOIpIquag MUODF] Suasapul uoipiquiag Aase) Unis) uoIpiquag MUODY] Winuiiyosdp uoipiquag Aased Ssdaaiavdo ajonbavul uoipiquiag lB Wrnudydaz uoipiquag uealagd snagdjoy? snyIa4 I, AYS[NYyOSIO|P S1UUadiAo snyIad uosysuq snsnjgo snyra4 I uvolag snuiiusaj]p snojdiq MUODIT] mays snjoovz WwioysJouuRYy S1usisul Diaporsosg Wloipury snoyiond sniu1yosdq ZYOYISYISA DIDJIUNdWUAIAP DAIIUOT AQu0ry aypiaarppo snaydoy sy JQINOH snupajds snuvduawuy snaydoj, yz ZYOYOsyIsq snoalds snprydon0N NUODY] Vsuadlp DUGAN JOYOSL] MM 1aYysIUUDL DUGAN JOYISLA NSAIG]YDS 1349G]YDS DUGAN ysneueaey ppipy Dugan ysneueary avasino] V1UgaN YIOIPUTT ADIJOpADYI SUGAN WIdYJOUURYY S7SOUISNALUAf SNISIAT sninuqey SniDpap] snqpiv.D JIN SDs4Owau snqoivy sue] Snipjnosaqnyi snayod Dd JOYyISI{ SNIDUISIDU SNjoUulydDIg wiaysauuRY] S1//OI1Jsnsuv snjoulydvIg MUODF] DUuosas0 Djapulr1y uOoxe FiGcures 84-85. Geographical distributions. FiGuReE 84. Western North American Range Pattern. FiGuRe 85. West Coastal Range Pattern. CALIFORNIA ACADEMY OF SCIENCES 88 FH NANN OTH KH HBA THT TFHHAntnerawnannnm nN Ol S sieqey ON S ol 8 at x ‘UMOUYUN UONNQUIsIpP 1eWqeYy = { ‘WeUQeY Ul PuNoy JOU = — “Je1IGeY UI INd90 OF UMOUY = X :sUOTINqLISIP 1eUqeH 7 “Quoz SuId;y = ¢] ‘1Ssas0j auIdjeqns = pf] ‘a10YS Weds 10 ayR| pueld”) = E] ‘IS910J sNOsayIUOD pueldy = TI ‘1sasoy snonptoap purjdy = [| {30q Jo yssew pueldy = QO] ‘OYs Was papeys ‘]]PWS PUR[MOT = 6 ‘JOYS WRIIIS ITIL] 10 aye] UBdO PUR|MOT = g ‘IS2IOJ SNOIJIUOD PUR|MOT = / ‘182104 snonptoap purmo7 = 9 ‘30q 10 ysieW puR|MOT = ¢ ‘dIdosylueUAs/uado puR|MOT = p ‘MOpeaw [epn-eidng = ¢ ‘yoeaq Bas Jaddn Apueg = Z ‘yoraq vas Jaddn ajqqo.y = | ‘sadA1 ie1qeH | Sl x Il +1 SI rl el xX x x x ol x KK OK 7°, adA1 1e1qeH “GaNNILNOD) 9 a 9 qeiiqey jad sjrio] ueslod snulsiu snjjaodpoig [EYUITJAD snjpusoI snjjaroyu | uvalag snjuajnuwmos snjpdivy] WayiauURY] $1/D40]]1] DDL Kasey SISdyja DAD Aased Dsonuls Dap ueraled ajjooiiaig UiNUuos F NUON] SUIosaj]DjaU WNUOs P yey lajjaq wnuosp uralag wWnsoulsnisaf Wnuos F uealoq snupdi snyoisosa ZYOYISYISA SNJIUISPY SNYINSOLII uvalad SnaUDISVI SNYINSOIII WIIYIIUURY] SUTISAYJIWUD SNYINISOIA MUODIT SNpisjd SNYyINSOsI) MUODF] S1JOMUIII SNYINSOMA SOLIDUIP] DUD] SNYINISO4II)] uealad snapwsdd snnuony WIdysJouURY, Winjnsu0/go UOIpIquiag yloIpury] Saplojnsuojqo uoipiquag wraylouuRYP ajiqDjoads uoipiquiag AYS[NYISIOW WiNJDLUJSaOf UOIpIquiag NUODA] 40/0IIS49A UOIpIqueag uvalag winjoulsipul uoipiquiag MUODI] wnjouasul uoipiquag uvalag a/psaaasuvs] WoIpiquiag [4 Winainviiuas Winjounlas uoIpiIquiag wlaysiouuRy Winynosniunjd uoipiquiag uoxe 9 a1av KAVANAUGH—CARABID BEETLES OF THE QUEEN CHARLOTTE ISLANDS 89 Vv Nebria charlottae Lindroth we Nebria louiseae Kavanaugh FiGure 86. Geographical distributions. Endemic Range Pattern. is known about the fauna of the region as a whole to permit recognition of general trends in species diversity and patterns in the regional distributions of species. Kavanaugh (1989) compared the carabid beetle fauna of the Queen Charlotte archipelago with the known faunas of eight other areas along the Pacific Northwest coast, using both Simp- son (Simpson 1960) and Jaccard (Braun-Blanquet 1932) coef- ficients of faunal similarity. Areas compared included (1) the Aleutian Islands, (2) the Alaskan Peninsula, (3) Kodiak Island, (4) the Kenai Peninsula, (5) the Lituya Bay region, the (6) Prince Rupert and (7) Vancouver areas of mainland British Columbia, and (8) Vancouver Island. Although those comparisons were based only on data available by early 1984, inclusion of the many additional distributional records that have accumulated since that time does not significantly alter the original findings, which can be summarized as follows: (a) The fauna of the Queen Charlotte Islands is unexpectedly diverse in relation to the adjacent mainland, when compared with other island areas worldwide (Darlington 1971; MacArthur and Wilson 1967). In general, insular faunas are either depau- perate versions of adjacent mainland faunas (e.g., Iceland or the Taste 7. Diversity OF QUEEN CHARLOTTE ISLANDS CARABID SPECIES IN RE- LATION TO HABITAT. No % of Habitat type spp. fauna! 1. Cobble upper sea beach 6 10.0 2. Sandy upper sea beach 6 10.0 3. Supra-tidal meadow 10 16.7 4. Lowland open/synanthropic site 13 2157 5. Lowland marsh or bog 14 23:3 6. Lowland deciduous forest 15 25.0 7. Lowland coniferous forest 13 21 8. Lowland open lake or large stream shore 11 18.3 9. Lowland small shaded stream shore 15 25.0 10. Upland marsh or bog 11 18.3 11. Upland deciduous forest 8 13.3 12. Upland coniferous forest 10 16.7 13. Upland lake or stream shore > 8.3 14. Subalpine forest 5 8.3 15. Alpine zone 10 16.7 Grouped habitat complexes” Beaches (1 + 2) it 18.3 Open areas (3 + 4) 14 23.3 Forests (6 + 7 + 11 + 12 + 14) 20 33:3 Wetlands (5 + 8 + 9 + 10 + 13) 35 58.3 Alpine zone (15) 10 16.7 Altitudinal habitat complexes? Lowland habitats (1 through 9) 54 90.0 Upland habitats (10 through 15) 29 48.3 ' In this calculation, the total number of species in the fauna is 60, rather than 62, excluding the two species for which habitat distribution is still unknown. 2 Number(s) listed after each habitat group correspond to the basic habitat types listed in the first part of the table. Faeroe Islands [Downes 1988] in relation to the northwestern coast of Europe) or highly distinctive faunas of moderate to low diversity in relation to comparable areas on mainlands (e.g., the Madeiran or Canary Islands in relations to coastal Europe or northern Africa). In fact, the fauna of the Queen Charlotte Is- lands (60 species, excluding the two species thought to have become extinct) is, at present, more diverse than the known fauna of the adjacent mainland coast (50 species). Although the latter area has not yet been intensively sampled and significant additions to its faunal list can be expected from future collecting efforts, the balance in diversity 1s, nonetheless, exceptional. (b) The fauna of the Queen Charlotte archipelago is much more similar to faunas of the Prince Rupert area and all the included areas to the south than to those of any areas to the north, based on comparisons of both percentages of Queen Char- lotte species shared and the coefficients of similarity examined (Kavanaugh 1989). ORIGINS AND DEVELOPMENT OF THE FAUNA Much of the evidence needed to infer the origins of the extant carabid fauna of the Queen Charlotte archipelago is provided by the data and analyses presented above, including the geo- graphical and habitat distributions of individual species, pat- terns observed among these geographical and habitat ranges, and the relative affinities of the fauna with carabid faunas of other selected areas along the Pacific Northwest coast. This evidence must also be viewed in the context of the known en- 90 CALIFORNIA ACADEMY OF SCIENCES | IX) Sa ORK Pa eeisicinarine Bex eocictes Ceasers g Ox ey IRIE NG iY, Carabus nemoralis Muller Trechus obtusus Erichson Ficure 87. Geographical distributions. Introduced Range Pattern. KAVANAUGH—CARABID BEETLES OF THE QUEEN CHARLOTTE ISLANDS 91 Transamerican 20 Holarctic & te ntroduce s ; \ enaemic Xs S \ x \ ESS > ‘ West Coastal 4( Ficure 88. Diagram illustrating the relative representation of different faunal elements in the carabid beetle fauna of the Queen Charlotte Islands. Values = percentage of species (” = 62 species). vironmental history of the region. Although many details of this history for the Queen Charlotte Islands and for the coastal main- land are yet unknown, most of its major features are clear (Mat- thews 1979b). Most of the Pacific Northwest coastal region was heavily and repeatedly glaciated during Pleistocene time, last as recently as 10,000—13,000 years ago (Sutherland Brown and Nasmith 1962; Clague 1989). However, several areas have been identified as likely sources (refugia) from which post-glacial dispersal into and within the region may have occurred (Heusser 1989; Ka- vanaugh 1989). Included among these sources are both extrinsic (peripheral) and intrinsic (within the region) refugia (Fig. 93). The existence of extensive glacial refugia, both northwest and south of the Cordilleran and Laurentide icesheets, is supported by abundant geologic and paleoenvironmental data (Hopkins 1967; Scudder 1979; Morgan and Morgan 1980). Biotic data suggest at least two distinct refugia south of the icesheets during glacial periods: one east and one west of the Sierra Nevada/ Cascade crest (Kavanaugh 1988, 1989). In the northwest, the Yukon River Valley/Alaskan interior remained largely ice-free B Vascular plants (584 spp.) Odonata (13 spp.) oO Diptera: Simullidae (16 pp.) Z Coleoptera: Curculionidae (19 spp.) i Coleoptera: Carabidae (62 spp.) 6 of fauna Crd AAARRRRARRRRRRR RR HOL TRA WNA WwCO END INT geographical range pattern Ficure 89. Graph illustrating the relative representation of different faunal elements (groups of species sharing a common geographical range pattern) for several different groups of organisms in the Queen Charlotte Islands. Geographical range patterns: HOL = Holarctic; TRA = Transamerican; WNA = Western North American; WCO = West Coastal; END = Endemic; and INT = Introduced. and continuous with northeastern Eurasia across the broadly exposed, unglaciated Bering Land Bridge (Hopkins 1967; Lind- roth 1979a; Matthews 1979b). The Aleutian Islands remained separate from the land bridge during glacial periods and ap- parently served as another extrinsic refugial area (Lindroth 1963; Tasce 8. Diversity OF Que EN CHARLOTTE ISLANDS CARABID SPECIES IN RELATION TO GEOGRAPHICAL RANGE PATTERN AND HABITAT TY pe, ! Habitat type Holarctic Cobble upper sea beach 1 - Sandy upper sea beach - - Supra-tidal meadow 2 3 Lowland open/synanthropic site 2 3 Lowland marsh or bog 1 4 Lowland deciduous forest - a Lowland coniferous forest 1 - Lowland open lake or large stream shore 1 1 Lowland small shaded stream shore - 1 Upland marsh or bog - 6 Upland deciduous forest - - Upland coniferous forest - - Upland lake or stream shore - - Subalpine forest - - Alpine zone - - Transamerican Geographical range pattern W. No. Amer. West Coastal Endemic Introduced i} Ww N i} 7S an | nen ot Nod Www woonwn 1 N= Re PNNK AND -h—- OD OO i i} ' Values refer to number of species for each combination of habitat type and geographical range pattern. [] macropterous [] dimorphic | | brachypterous ® ® a = 10 o vr) —E 5 = 2 3 4 5 6 ca 8 9 10 Ww 12 1c 14 1S habitat type FiGure 90. Graph illustrating the number of carabid species, and the condition of flight wings for members of each species, in different habitat types in the Queen Charlotte Islands. Habitat types: | = Cobble upper sea beach; 2 = Sandy upper sea beach; 3 = Supra-tidal meadow, 4 = Lowland open/synanthropic; 5 = Lowland marsh or bog; 6 = Lowland deciduous forest; 7 = Lowland coniferous forest, 8 = Lowland open lake or large stream shore; 9 = Lowland small, shaded stream shore, 10 = Upland marsh or bog; 11 = Upland deciduous forest; 12 = Upland coniferous forest; 13 = Upland lake or stream shore; 14 = Subalpine forest; 15 = Alpine zone Karlstrom 1969). Geologic, paleoenvironmental, and biotic ev- idence suggest that several areas within the Pacific Northwest coastal region, including parts of Kodiak Island (Lindroth 1969), southeastern coastal Alaska (Heusser 1960, 1989), the Queen Charlotte Islands (Foster 1965; Heusser 1989; Kavanaugh 1989), and Vancouver Island (Heusser 1960, 1989; Ogilvie 1989) served as intrinsic refugia, which were potential source areas for the present carabid fauna of the Queen Charlotte Islands. Post-glacial Immigration From the South Species that are restricted to the Pacific Coast region make up 40% of the carabid fauna of the Queen Charlotte Islands, the largest single component (25 species) in the fauna. Of these, 10 species reach the northern limit of their present distribution in the archipelago. Clear affinities with coastal faunas to the south, but not to the north, suggest a south coastal origin for most if not all of the species in this component of the fauna, probably through range extension northward along the coast (Fig. 94) following deglaciation. In fact, fossil remains of two of the West Coastal species (Scaphinotus angusticollis and S. marginatus) have been recovered from late glacial deposits in a coastal site just south of the Cordilleran ice margin (Morgan and Morgan 1980). Together, species with Western North American (18 species), Transamerican (12 species), and Holarctic (three species) dis- tributions represent 54% of the carabid fauna of the archipelago. Although they differ in the extent of their present distributions, these species all have one feature in common —they presently inhabit at least some area(s) south of the region that was covered by Cordilleran and/or Laurentide icesheet(s) in full-glacial times. CALIFORNIA ACADEMY OF SCIENCES EJ macropterous o @ 3 el dimorphic a a B brachypterous o = © Da 10 iS 5 = 1 2 3 4 5 6 7 8 9 number of habitats occupied Figure 91. Graph illustrating the frequency distribution for number of hab- & y itats known to be occupied by carabid species and the condition of flight wings for members of each species, in the Queen Charlotte Islands. Fossil remains of seven of these species (Carabus taedatus, Ela- phrus clairville, E. americanus, Bembidion incrematum, B. ver- sicolor, and B. fortestriatum) have been recovered from late glacial deposits in sites just south of the Laurentide Icesheet in a ® L] macropterous a s J dimorphic o a 1 & brachypterous HOL TRA WNA WwCo END INT geographical range pattern FiGure 92. Graph illustrating the relationship between geographical range pattern and condition of the flight wings for members of carabid species, in the Queen Charlotte Islands. Geographical range patterns: HOL = Holarctic, TRA = Transamerican, WNA = Western North American; WCO = West Coastal; END = Endemic; and INT = Introduced KAVANAUGH—CARABID BEETLES OF THE QUEEN CHARLOTTE ISLANDS 93 Ficure 93. Map illustrating approximate extent of glacial icesheets, proportions of the Bering Land Bridge, and location of proposed refugia during the last glacial maximum (ca. 17,000 years ago) (adapted from Lafontaine and Wood [1988], Schwert and Ashworth [1988], and Kavanaugh [1989]): ticked solid line = boundaries of icesheets or major glaciers; dashed line = southern margin of the Bering Land Bridge; AO = Arctic Ocean (frozen, with polar icecap grounded on the Bering Land Bridge and broadly, but not completely, coalesced with the continental icesheet; CI = continental icesheet, including both Cordilleran and Laurentide Icesheets, a = Yukon/Beringian refugium; b = Aleutian Islands refugium; c = Kodiak Island refugium; d = southeastern Alaska coastal refugia; e = Queen Charlotte Islands refugia; f = Vancouver Island refugium; g = southern coastal refugium(-a); h = southern interior refugium(-a) southeastern and southcentral Canada and the northcentral and northeastern United States (Morgan and Morgan 1980). Based on both present distributions and the fossil record (Ashworth 1979: Morgan and Morgan 1980; Schwert and Ashworth 1988), it seems likely that all of these species survived (or could have) in the continental refugium(-a) south of the ice. There is no evidence to suggest post-glacial immigration of elements from the Yukon/Beringian refugium or any other proposed refugium north of the archipelago, except perhaps from the Aleutian area (Kavanaugh 1989). At least one of the carabid species in the extant fauna of the archipelago, Diplous aterrimus, is broadly distributed in western North America, but not in the southern coastal region. That such a southern interior form, rather than its closely related southern coastal counterpart, Diplous filicornis Casey, occurs in the archipelago suggests a separate, southern interior origin for at least part of the fauna. Just how many of the Western North American, Transamerican, and Holarctic species colonized the archipelago from the southern coastal refugium, from the south- ern interior refugium, or from both remains unclear at present. Why a southern interior form such as Diplous aterrimus should have colonized the northern coast while its southern coastal 94 CALIFORNIA ACADEMY OF SCIENCES Ficure 94. Schematic representation of proposed post-glacial immigrations into and emigration from the Queen Charlotte archipelago: a = immigration from southern coastal refugium(-a); b = immigration from southern interior refugium(-a), c = emigration from one or more refugia in the archipelago to the adjacent mainland counterpart apparently did not is an intriguing question. One possible explanation is that early post-glacial climate of at least parts of the archipelago and adjacent mainland coast was drier and colder than at present, conditions that would favor forms adapted to the southern interior rather than the southern coastal region. This interpretation is supported by some paleobotanical evidence (R. W. Mathewes 1989 and pers. comm.) and by the discovery of a fossil specimen of the carabid subspecies Nebria gyllenhali castanipes Kirby (by R. W. Mathewes and B. G. War- ner) in deposits from Cape Ball, in eastern Graham Island, dated at 11,100 + 90 years before present. This form, widely distrib- uted over most of boreal North America, is not known to occur at present in the archipelago or anywhere else on the Pacific Coast west of interior British Columbia. In the Rocky Mountain region, this species 1s often represented in upper montane and subalpine habitats in association with aspen (Populus tremu- loides Michx.), subalpine fir Abies lasiocarpa [Hook.] Nutt.), and the carabid species Carabus taedatus. As noted above, C. taedatus appears to have become extinct in the archipelago rel- atively recently. Clearly, the climate of the northwest coastal region did not become as maritime as it is today until after 11,000 years ago. Survival in Refugia Within the Archipelago The occurrence of plant and animal species that are apparently endemic to the Queen Charlotte Islands has been cited as evi- dence in support of the existence of one or more glacial refugia KAVANAUGH—CARABID BEETLES OF THE QUEEN CHARLOTTE ISLANDS 95 in the archipelago (Foster 1965; Calder and Taylor 1968; Ka- vanaugh 1989; Ogilvie 1989; Schofield 1989), which served as an additional, in situ source for the present fauna. However, as noted by Anderson (1988) and others, alternative hypotheses can account for the presence of these endemics. Each of these alternatives is considered here briefly. First, species thought to be endemic to the archipelago may actually be more widely distributed. In fact, this has proven to be true for all but one of the 11 vascular plant taxa previously considered endemic (Calder and Taylor 1968). Ten of them are now known to occur also on northern Vancouver Island (Ogilvie 1989). The freshwater amphipod, Paramoera carlottensis Bous- field, described as an endemic, is now known to be widely dis- tributed along the coast; and the supposed endemic carabid, Nebria haida (Kavanaugh 1984, 1989), recently has been found on the adjacent mainland, just north of Prince Rupert. Sampling along the adjacent coastal mainland and on other coastal islands remains so incomplete for insects that most or all of the re- maining proposed endemic insects, including two carabids (Ne- bria charlottae and Nebria louiseae), one cercopid bug (Aphro- phora regina), and one geometrid moth (Xanthorhoe clarkeata Ferguson) may also occur outside the archipelago. Although discovery of populations of supposed endemic forms outside the archipelago weakens the hypothesis of endemism at present, it does not necessarily refute the hypothesis of glacial survival in refugia within the archipelago. Post-glacial coloni- zation of the adjacent mainland or other islands from one or more refugia in the Queen Charlotte Islands is no less plausible than the reverse pathway (i.e., from mainland to islands). In fact the distributions of three non-endemic carabid species (which are included among those classified as West Coastal in distri- bution) suggest this possibility. Nebria haida, Bembidion viator, and B. oblonguloides are known only from the archipelago and the mainland immediately adjacent to it (i.e., the Prince Rupert area). A glacial refugium in the archipelago would have been the most proximate and likely source area for the mainland populations of these three species. A second hypothesis to explain true endemism in the archi- pelago, to the extent that it occurs, is that these species survived glacial periods outside the area, immigrated in post-glacial time, and subsequently became extinct elsewhere. Although this pos- sibility cannot be dismissed, I know of no paleontological evi- dence to support it for any of the supposed endemic plant or animal species. A third hypothesis is that the endemic forms differentiated in post-glacial time within the archipelago, from populations of ancestral species that immigrated to the archipelago following deglaciation. This explanation has been invoked to account for the occurrence of several endemic vertebrates, particularly stick- leback fishes (Moodie and Reimchen 1976a), in the archipelago. Presence on the adjacent mainland of the respective sister spe- cies of these supposed endemic forms would be strong, but not conclusive, evidence in support of this hypothesis. Discovery of such vicariance patterns would suggest that the common ancestors of the species pairs reached the archipelago from the mainland and that subsequent isolation permitted differentia- tion. However, this isolation need not have occurred in post- glacial time, but could have been completed earlier, during ei- ther a glacial or interglacial period. Present occurrence of the sister species on the adjacent mainland could represent reoc- cupation of that area following deglaciation. Unfortunately, knowledge of the phylogenetic reiationships of plant and animal species in the archipelago is woefully lacking, hence the recog- nition of sister species and of their geographical and habitat distributions is generally not possible. An important exception to this generalization, however, is the carabid genus Nebria, which includes the two endemic car- abid species and the possible Glacial-age endemic, Nebria haida, and for which an hypothesis of phylogenetic relationships has been developed (Kavanaugh 1989). These three species are all members of the gregaria infragroup (Kavanaugh 1978), which presently includes five species. Evidence in support of (1) the taxonomic ranking of populations (and groups of populations) in this infragroup as distinct species and (2) the hypothesis of phylogenetic relationships among all those species discussed here will be presented separately in a revision of the Nearctic Nebria fauna (in prep.). The sister species of N. haida is Nebria lituyae Kavanaugh, known only from the Lituya Bay, White Pass, and Juneau areas of southern Alaska. Members of both of these species are restricted to treeless alpine summits and ridges in their respective areas. The sister species of N. /ouiseae is Nebria gregaria Fischer, which is apparently restricted to the Aleutian Islands. The sister taxon of N. charlottae is the species pair N. louiseae + N. gregaria. Members of all three species occur in cobble upper sea beaches, although adults and larvae of N. gregaria are also found under debris on supratidal flats of volcanic ash substrate. All five species in the gregaria infragroup occur in areas for which glacial refugia have been proposed on other biotic and/or geologic grounds. Kavanaugh (1989) re- viewed various hypotheses to account for the present distri- butions of these species, the observed vicariance among species pairs, and the occurrence of both N. /owiseae and N. charlottae in the archipelago (but with allopatric distributions). At present, the conclusions reached in that paper, and summarized here, best account for all available data: (1) Nebria haida, N. louiseae, N. charlottae, and perhaps also Bembidion viator and B. oblon- guloides all probably survived at least the last glacial period in refugia within the archipelago; and (2) Nebria charlottae prob- ably survived more than one glacial period in the archipelago, at least the last in a lowland refugium somewhere on Graham Island. Unfortunately, no fossil specimens have yet been found to conclusively demonstrate the presence of the endemic Nebria or other carabid species in the archipelago during full glacial times; and until such specimens from that period are found, refugial survival of these carabids will remain in question. In fact, no organic deposits of any kind have been found that date from the crucial period between 21,000 and 16,000 years ago (Mathewes 1989a, and pers. comm.,; Blaise et al., in press), so the hypothesis of glacial refugia in the archipelago remains un- supported by conclusive data at present. Nonetheless, what we know about the present habitat and geographical distributions of the proposed glacial survivors can provide clues as to the number, location, and nature of the refugia that they would have required. During the last glacial period, populations of Nebria haida could have survived on the alpine summits, ridges, and head- lands that remained ice-free, as nunataks, above and between small icecaps and the valley and piedmont glaciers that flowed off the Skidegate Plateau and out of the Queen Charlotte Ranges. 96 Such nunataks probably served as a network of small-scale re- fugia, on both Graham and Moresby islands, each with small populations of N. haida that were in genetic contact with each other through the dispersal of individuals. Heusser (1954) de- scribed the surprisingly diverse biotas that survive at present under similar conditions on nunataks in heavily glaciated regions, such as in the Juneau Icefield area of southeastern Alaska. Survival of Nebria charlottae and N. louiseae would have required refugia that included cobble sea beaches, a habitat that would have persisted at least at the bases of headlands between valley glaciers on the west coast of Graham and Moresby islands. Suitable habitat may actually have been more extensive during full glacial times, when portions of the continental shelf were exposed due to lowered sea levels (Fairbridge 1960; Hopkins 1967; Warner, Mathewes, and Clague 1982). Separate lowland refugia are inferred for these two species—somewhere on Gra- ham Island or the adjacent exposed continental shelf for N. charlottae and somewhere in the southern part of the archipelago for N. louiseae. Luternauer et al. (1989) found and described Late Pleistocene terrestrial deposits on the continental shelf northwest of Van- couver Island, at a depth of 95 m below present sea level, which represents a site that was above sea level 10,500 years ago. This discovery suggests that the search for deposits of full glacial age, perhaps from areas that served as refugia, might best be redi- rected to areas of the submerged continental shelf. It 1s also important to note that a depression in sea level of 100 m would expose a broad, dry land connection between the eastern coast of the northern two-thirds of the archipelago and the mainland and inner islands in the Prince Rupert area. The findings of Luternauer et al. (1989) suggest that such a connection existed as recently as perhaps 10,500 years ago or later. It well may have served as the pathway for early post-glacial colonization of the archipelago by immigrants from the south and for col- onization of the adjacent mainland by at least three carabid species from the archipelago. If Bembidion viator and B. oblonguloides also survived in glacial refugia in the archipelago and, as suggested above, col- onized the adjacent mainland in post-glacial time, then the re- fugium(-a) in which they survived should have included lowland marshes or bogs and at least some small patches of coniferous or deciduous forest. The paleobotanical record from northeast- ern Graham Island (Warner et al. 1982; Mathewes 1989a, b) confirms the presence of marshland vegetation and suggests the possibility of stunted spruce in the area as early as 16,000 years ago. Both Bembidion species are known only from Graham Island in the archipelago at present, but the probable location of a glacial refugium for their survival in the area cannot be determined using available data. Introductions with Man Spence and Spence (1988) identified 20 species introduced by humans into western Canada. Two species in the present fauna of the archipelago, Carabus nemoralis Miller, and Trechus ob- tusus Erichson, are native to western Europe, including the Brit- ish Isles, and have been introduced into North America in his- toric times (Lindroth 1957). Both of these species are restricted to synanthropic sites in the archipelago, as well as elsewhere throughout their original and introduced ranges. CALIFORNIA ACADEMY OF SCIENCES Specimens of Carabus nemoralis were first collected in the archipelago in August 1982, at Port Clements on Graham Is- land. All subsequent records have been from the same area. This species was first recorded from North America (New Bruns- wick) in 1870 (Lindroth 1957), from western North America (Washington) in 1909 (Hatch 1953), from Vancouver, British Columbia in 1925, and from Prince Rupert in 1958 (Lindroth 1961-69). In western Canada, it now occurs in scattered, ap- parently disjunct localities in the southern half of British Co- lumbia and in Edmonton, Alberta. All adults of this species are brachypterous, hence flightless. Apparently, the species is ex- tending its range in leaps, no doubt aided by humans, rather than by a gradual, progressive spread by means of its own dis- persal (ambulatory) capabilities. Specimens of this species have been intercepted in shipments of nursery stock (in the bolus of soil around root balls) arriving in Vancouver (Spence and Spence 1988), and are known to have been transported in hay bales. Either of these means could have served as the mode of intro- duction to the archipelago as well. The restriction of C. ne- moralis to one site on Graham Island suggests very recent in- troduction, probably during the late 1970s. The first specimens of Trechus obtusus collected in the ar- chipelago were found 1.4 km south of Masset, again on Graham Island, in August 1983. All subsequent records have been from Masset and its immediate vicinity. Hence, this species is prob- ably also a very recent introduction. Kavanaugh and Erwin (1985) reviewed the history of this species in western North America and concluded that its primary introduction, as well as most of its subsequent spread, had been in nursery stock. Trechus obtusus has not yet been recorded from the Prince Ru- pert area, or anywhere else north of the greater Vancouver area. The latter is a most likely source area for the introduction of this species to the archipelago, whereas either the Vancouver or Prince Rupert areas could have served as sources for the intro- duction of C. nemoralis. Dispersal To and Within the Archipelago Clearly, all but a few of the species represented in the extant carabid fauna immigrated to the archipelago in post-glacial times; but when and how did they reach the islands? Was Hecate Strait a sea gap that had to be crossed, was it covered with glacial ice flowing west off of high peaks of the Coast Range of the main- land, or was it at least partly dry land, connecting the islands with the adjacent mainland? Evidence suggests that the area that is now Hecate Strait presented all of these aspects at one time or another since extensive lowland ice-free areas are first known to have occurred in the archipelago as early as 16,000 years ago (Warner et al. 1982; Clague 1989). Although sea level was prob- ably low enough at this time to expose large parts of the floor of Hecate Strait, perhaps even providing a dry land link to the mainland, the adjacent mainland coast remained ice-covered until less than 13,000 years ago (Clague 1981). Hence, coloni- zation directly from the mainland was probably not possible until after that time. If the magnitude of depression in sea level proposed by Luternauer et al. (1989) for the continental shelf northwest of Vancouver Island at 10,500 years ago occurred also in Hecate Strait at the same time, dry land would have connected the archipelago with the adjacent mainland. By 10,000 years ago, however, the archipelago was completely deglaciated KAVANAUGH—CARABID BEETLES OF THE QUEEN CHARLOTTE ISLANDS 97 and sea level had risen to its present level relative to land (Clague 1989). By about 8,000 years ago, sea level reached a post-glacial high about 15 m above present. It therefore appears that, to have reached the archipelago from the mainland across a land connection, post-glacial immigrants from the south must have done so between 13,000 and 10,000 years ago. It is possible that species extended their ranges northward at an earlier time on the exposed continental shelf to the west of the still glaciated mainland; but depressions more than 200 m below present sea level traverse the shelf in several areas along the route (Barrie and Bornhold 1989; Thomson 1989). It is therefore unlikely that a continuous dry land route has ever been available on the continental shelf in post-glacial time. As noted above, in the discussion of species distributions within the archipelago in relation to brachyptery, there appears to be little or no barrier at present to the dispersal of some flightless carabid beetles between islands. It was noted, in fact, that some species with brachypterous adults occurred on more islands than those with macropterous individuals. It should be noted, however, that members of each of the broadly distributed species are associated with logs—either with rotting logs on the lowland forest floor (e.g., Scaphinotus marginatus, Pterostichus algidus, P. amethystinus, and P. crenicollis), or driftwood logs on sea beaches (e.g., Nebria louiseae). Inter-island dispersal by rafting in or on logs may account for the surprisingly wide dis- tributions of these species in the archipelago. It may also have played a role in initial colonization of the archipelago for some of these same species, but no such mechanism for long-range dispersal across a water gap appears to be required. Although the distributions and relative abundances of various plant spe- cies in the archipelago have changed dramatically during the last 11,000 years (Mathewes 1989a), it seems clear that all 15 of the present habitat types were represented in the archipelago before 10,500 years ago, when dry land probably linked the archipelago to the mainland. Summary It appears that the extant fauna of the Queen Charlotte Islands includes: (1) two species that are western European in origin, recently and secondarily introduced to the archipelago from a source area on the mainland of British Columbia; (2) two-five species that survived at least the last glacial period in one or more refugia in the archipelago, at least three of which expanded their ranges in post-glacial times to include a limited area on the adjacent mainland; and (3) 55-58 species (including two that are probably now extinct in the archipelago) that survived the last glacial period in one or more refugia south of the Cor- dilleran and/or Laurentide icesheets and colonized the archi- pelago in post-glacial times. These immigrants probably reached the archipelago across a dry land connection with the mainland between 13,000 and 10,000 years ago. FAUNAL EVOLUTION Although they have been called the “Canadian Galapagos” (Foster 1985), no adaptive radiations comparable to those we associate with the Galapagos Islands, the Hawaiian Islands, or even the West Indies are evident in the extant biota of the Queen Charlotte Islands, except perhaps among populations of three- spined stickleback fishes (Moodie and Reimchen 1976a, b; Reimchen et al. 1985; Gach and Reimchen 1989). The archi- pelago is remote, diverse in the number and form of its islands, and rich in habitat and species diversity. Yet the area was so recently and drastically affected by Pleistocene climatic changes and events that its biota is predominantly post-glacial in age. In the extant carabid fauna, there is little indication that evo- lutionary change (divergence) has occurred among populations of individual species on different islands, or between mainland populations and their counterparts in the archipelago. Most sur- prisingly, there is apparently not a single instance of the devel- opment of brachyptery among carabids in the archipelago. Al- though many species represented have some or all adults brachypterous, members of respective populations of these spe- cies or their sister species on the mainland or elsewhere are also brachypterous, indicating a trait acquired before, not after, iso- lation in the archipelago. The only known instance of inter-island differentiation noted to date in the carabid fauna of the archipelago is found in Sca- phinotus marginatus. Adults of this species on Reef Island, the most isolated island that has been sampled, are larger on average than those from other islands or the mainland, although a few large individuals are found occasionally in populations through- out the range of the species. The Reef Island population also differs from all others sampled in the proportion of individuals that have a metallic green reflection on their elytra. On Reef Island, 28 out of 30 specimens (93%) studied had green elytral reflection. In all other known populations, individuals with green reflection are either absent or rare, and in no other sample do they represent as much as 10% of the population. In coastal areas, the elytra typically have a vivid, metallic violet reflection. The Reef Island population seems to represent a classic example of Mayr’s (1942) founder principle. This population may be derived from a single large, green, gravid female ancestor that dispersed to the island and founded the population. Carabid Species Diversity and Equilibrium Theory Darlington (1943, 1957), initially through his study of cara- bids of the West Indies, was among the first to recognize a general relationship between the areas of islands and the diversity of species of a particular taxon that occurs on them. Expanding on these and other pioneer studies, including their own (e.g., Wilson 1961), MacArthur and Wilson (1963, 1967) developed a general theory of island biogeography, commonly referred to as the “Equilibrium Theory,” to explain the faunal diversity on is- lands. According to this theory, species diversity reaches an equilibrium when the rate of extinction of species on an island equals the rate of immigration of new species to the island. For a given island, both time required to reach equilibrium and the number of species occurring at equilibrium may be influenced by factors such as its distance from source areas (e.g., mainland faunas) and its area, elevation, and habitat diversity. During the past three decades, numerous studies have tested the general theory and explored the effects of various factors on species diversity on true islands (e.g., Diamond 1969; Wilson and Simberloff 1969) and islands of restricted habitat on con- tinents (e.g., Brown 1971, 1978; Patterson and Atmar 1986; Davis et al. 1988; Dunn and Loehle 1988). Among these factors, the relationship between island area and species diversity has been studied most intensively. An approximation of this rela- tionship is given by the equation S = Cd4* (MacArthur and Wilson 1967), where S is the number of species of a given taxon occurring on the island, C is a coefficient (or the intercept in the log-transformed linear equation), 4 is the area of the island, and zis an exponent of a power function (or the regression slope in the log-transformed linear equation). This equation can be used to describe the relationships between diversity and other factors as well. The Queen Charlotte archipelago should be an ideal area for testing equilibrium theory and studying the effects of area and other factors on species diversity. There are about 150 different islands, varied in size from about 0.01 km? to more than 6,000 km2, in altitude from less than 10 m to more than 1,150 m (Table 1), and in habitat diversity from one or two to 15 different major carabid habitat types (Table 3). The carabid fauna of the archipelago should be ideal for study as well. The fauna is diverse and species occur in all terrestrial habitats, from sea level to the summits of the highest peaks. Unfortunately, the present state of our knowledge of the car- abid fauna of the archipelago is such that its potential for study cannot yet be realized. To date, only 23 islands have been sam- pled at all, and none of these (except perhaps for Graham Island) sufficiently to establish the total number of species present. Al- though we know of two species that have recently colonized the islands (through human introduction) and perhaps as many as three species that occupied the archipelago in the past but do not now occur there, we lack (1) the baseline inventories of the archipelago as a whole, (2) the baseline inventories of individual islands in particular, and (3) the subsequent inventories that we need to calculate rates of immigration and extinction. Such cal- culations must await both adequate baseline (initial) and sub- sequent inventories for each island. All that can be done at present is to use available data for the 23 islands sampled to date (Tables 1, 3) to quantify, at least in a preliminary way, relationships between species diversity and island area, eleva- tion, and habitat diversity in the archipelago. Figures 95-97 illustrate highly significant relationships apparent, through sep- arate regression analyses, between carabid species diversity and each of these three factors, respectively. The z value of 0.354, calculated for the species-area relation- ship, is somewhat higher than the expected 0.27 (MacArthur and Wilson 1967). This may be due to (a) an effect of increased habitat diversity on the largest islands (MacArthur and Wilson 1967), (b) incomplete sampling of carabid diversity on most of the islands (particularly on all but the largest islands), or (c) a combination of these and/or other, unknown factors. Z values of 0.706 and 1.260 for species-elevation and species-habitat diversity relationships, respectively, are difficult to interpret in the absence of comparative figures for other faunas and from other regions. However, with improved sample size and data, the interactions of these factors with area (as they affect species diversity), could be explored more appropriately using methods of principal component analysis. MacArthur (1972) distinguished “oceanic islands” (islands, such as the Galagapos, that have never been connected by a land bridge to mainland areas) from “land bridge islands” (is- lands, such as Trinidad, that have been recently connected to mainland areas by a land bridge and whose colonists, at least some of them, need never have crossed a water gap). He noted that land bridge islands may contain more species than their CALIFORNIA ACADEMY OF SCIENCES equilibrium number (1.e., for their size, if they were oceanic islands) and that their diversity will decrease with time following isolation—rapidly on small islands, very slowly on larger land bridge islands. Clearly, the Queen Charlotte Islands, located on the continental shelf, are “land bridge islands,” most if not all of which were connected to the mainland and/or each other perhaps as recently as 10,500 years ago. This may help to explain their relatively high diversity in relation to the adjacent main- land. OPPORTUNITIES FOR FUTURE RESEARCH ON CARABID BEETLES IN THE ARCHIPELAGO This report has reviewed the current status of our knowledge about the carabid beetle fauna of the Queen Charlotte Islands, its composition, affinities, and origins. Because the number of new species added to the faunal list from most recent collecting efforts has dropped significantly, and because all major habitat types in the archipelago have been sampled to some degree, it is unlikely that many more species presently occurring there remain unrecorded. Knowledge of the overall habitat and geo- graphical distributions of the species represented in the fauna are reasonably well known, with a few exceptions; and these data, along with paleogeographic and paleoenvironmental find- ings, allow us to infer, with some confidence, the affinities and origins of most, if not all, of the fauna. Nonetheless, much remains to be learned about the carabid fauna and its history. The following is a brief, annotated list of projects on which future research could be most fruitful. 1. Distribution of species within the archipelago. — Many parts of the Queen Charlotte Islands remain inadequately sampled. These include: (a) the entire west coast of the archipelago, es- pecially between Lepas Bay and Rennell Sound on Graham Island and almost the entire length of Moresby Island; (b) north- ern portions of the Skidegate Plateau on Graham Island; (c) Langara Island; (d) lowland marsh and bog areas of northwestern and northeastern Graham Island; (e) the entire San Cristoval Range of Moresby Island; (f) all islands not listed in Table 1 — nothing is known about the faunas of these islands, so all records for carabid species occurring on them will represent new infor- mation; and (g) all islands listed in Table 1, except Graham and Moresby islands (but see items [a], [b], [d], and [e] above)— knowledge of the carabid faunas of these islands is still incom- plete, for some, markedly so (e.g., only four species have been recorded from Kunghit Island, one from Maude Island). Col- lecting efforts in these areas will add greatly to our knowledge of the fauna, perhaps uncover unrecorded or undescribed spe- cies, expand our knowledge of the habitat and distributional ranges of the species, and permit better analyses of relationships between species diversity and island area and topographic and habitat diversity. 2. Monitoring of the faunas around townsites and in dis- turbed area.—Continued collecting in and at the margins of synanthropic sites will provide data needed to monitor the spread, if any, of the two known introduced species, Carabus nemoralis and Trechus obtusus, and any possible effects of such spread on the native fauna. Such a project would also be most likely to detect new introductions as they occur. 3. Sampling of the carabid faunas of the adjacent mainland and inshore islands.— Knowledge of the fauna of the adjacent KAVANAUGH—CARABID BEETLES OF THE QUEEN CHARLOTTE ISLANDS 99 100 number of 10 100 1000 10000 area (square km) 100 wo rab) O @ cy wo Ss 10 = (ob) a = =) c 0 10 100 1000 elevation (m) 100 les spec number o (0) | 10 100 number of habitats Ficures 95-97. Relationships between carabid beetle species diversity and three variables for 23 islands in the Queen Charlotte archipelago. Solid dots = individual islands (data from Tables 1, 3), solid diamonds = two islands with equal values, solid triangle = three islands with equal values. FiGure 95. Species diversity in relation to island area. The least-squares regression (solid line) is given by the equation log S (log number of species) = 0.174 + 0.354 log A (log area) [¢ = 3.669, P < 0.01]. Ficure 96. Species diversity in relation to maximum island elevation. The least-squares regression (solid line) is given by the equation log S = —1.108 + 0.706 log A (log elevation) [¢ = 3.370, P <= 0.01]. FiGure 97. Species diversity in relation to habitat diversity. The least-squares regression (solid line) is given by the equation log S = —0.258 + 1.260 log A (log elevation) [¢ = 3.353, P = 0.01). mainland remains inadequate. At present, more species have been recorded from the Queen Charlotte Islands than from the mainland west of the crest of the Coast Range, and it is unlikely that this relationship properly reflects relative diversities of the areas. Little is known about the faunas of the inshore islands along the British Columbia coast or the lower end of the Al- exander Archipelago, just to the north of the Queen Charlotte Islands. Faunas of these areas should be intensively sampled, 100 especially in the habitats known to harbor the apparent Queen Charlotte endemic species (i.e., cobble sea beaches) or proposed refugial survivors (i.e., alpine areas, lowland marshes or bogs, and deciduous and coniferous forests). Hypotheses developed in this report concerning the origins of the carabid fauna of the Queen Charlotte Islands, and the endemic species in particular, can be tested with faunal data from these areas. 4. Paleoenvironmental and paleontological studies. —Clear- ly, additional paleoenvironmental and paleontological data of appropriate age are needed if the existence of one or more glacial refugia in the archipelago is to be convincingly demonstrated. Although the present geographical and habitat distributions of organisms can be used to infer historical events, fossil specimens and associated paleoenvironmental data are required to cor- roborate these interpretations. The search for sites bearing or- ganic deposits of full glacial age should certainly continue on land in the archipelago; but the prospects now seem brightest for the discovery of such sites on presently submerged portions of the continental shelf, particularly within the 100-meter iso- bath around the archipelago, and between it and the adjacent mainland. ACKNOWLEDGMENTS Fieldwork for this study was supported by grants from the American Philosophical Society (Penrose Fund), the Boreal In- stitute for Northern Studies (University of Alberta, Edmonton, Grant-in-Aid), the National Geographic Society (Grant No. 3376-86), and the California Academy of Sciences (G. Lindsay Fund for Field Research and In-house Research Fund). Partial support of publication costs, especially for the inclusion of color plates, was provided by Mr. and Mrs. Peter H. Behr, Mr. and Mrs. Edwin Blue, Mr. and Mrs. Donald B. Dalziel, Dr. and Mrs. George Desalernos, Mr. and Mrs. Herman Eichorn, Mr. Dan. A. Holtman, Mr. Roy A. Holtman, Mr. and Mrs. George Hoyt, Mr. and Mrs. James W. Jordan, Dr. and Mrs. Seymour Jurow, Mr. and Mrs. William S. Mailliard, Mrs. Donald McNeil, Mr. and Mrs. Edward Mentzer, Mrs. Lawrence Metcalf, Mr. A. E. (Monty) Montgomery, Mrs. Francis R. Montgomery, Mr. and Mrs. John B. Nelson, Dr. and Mrs. Gordon W. Repp, Mr. and Mrs. Thomas N. Saunders, Mr. and Mrs. Barton Schackelford, Mr. and Mrs. Eugene A. Shurtleff, Ms. Helen H. Weaver, and an anonymous donor. This generous support 1s gratefully ac- knowledged. Permission to visit and study at several historic village sites was graciously granted by the Skidegate Band Coun- cil. Assistance and companionship in the field were provided by J. Milton Campbell, Daniel H. Mann, Jeffrey L. Kavanaugh, Michael D. Kavanaugh, Rolf W. Mathewes, and Barry G. War- ner. Additional material was collected specifically in support of this project by James S. Ashe, Joseph Belicek, Robert A. Can- nings, Nicholas Gessler, Jack Miller, Marsh Pitman, and Laurie Williams. I thank James S. Ashe, George E. Ball, Robert A. Cannings, Terry L. Erwin, Nicholas Gessler, John D. Lattin, Alan V. Morgan, Alfred F. Newton, Jr., L. L. (Verne) Pechuman, Geoffrey G. E. Scudder, Ales Smetana, William J. Turner, and George E. Wallace, curators in charge of collections from which specimens were borrowed, for providing access to the valuable materials in their care. Very special thanks are due to Tom Hockin and Marge Tighe, Nick and Trisha Gessler, Tom Reimchen and Sheila Douglas, and John and Les McLaughlin, CALIFORNIA ACADEMY OF SCIENCES who generously provided encouragement, advice, logistical sup- port, and sustenance for me and my field associates during our visits to the Queen Charlotte Islands. The scope of this project and the detailed results achieved simply would not have been possible without their exceptional help and support. Scanning electron micrographs were taken by Mary Ann Tenorio; Chris Krueger and Linda Grandke-Kulik provided advice and assis- tance in preparation of computer generated illustrations; Alan E. Leviton provided access to software and advice for data analyses; and Roberta L. Brett, Beverly A. Kavanaugh, Vincent F. Lee, and Norman D. Penny reviewed various drafts of the manuscript and provided many critical comments. The assis- tance of each of these individuals is much appreciated. LITERATURE CITED ANDERSON, R. E. 1988. The Curculionidae of the Queen Charlotte Islands, British Columbia (Insecta; Coleoptera). Canadian Journal of Zoology 66:2406- 2414. ARMIN, C. 1963. Astudy ofthe family Carabidae (Coleoptera) in Boulder County, Colorado. Unpublished Ph.D. Dissertation, University of Colorado, Boulder, Colorado. 476 pp. AsHwortH, A. C. 1979. Quaternary Coleoptera studies in North America: past and present. Pages 395-405 in T. L. Erwin, G. E. Ball, D. R. Whitehead, and A. L. 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APPENDIX A LocaLity DATA FOR CARABIDAE IN THE QUEEN CHARLOTTE ISLANDS Discussions of the geographical, altitudinal, and temporal dis- tributions of carabid species within the archipelago, presented in the preceding text, are based on data from specimens listed here. For each species, distributional data are grouped first by island and then by specific locality. Altitudinal data, month(s) of collection, total number of specimens for each locality, and depositories for specimens are also provided. 104 1. Cicindela oregona LeConte Graham Island: McIntyre Bay (Entry Point to Estrado Lagoon [2-5 m]) [Aug.] (16; CAS), Tlell River (1.2 km N of bridge on Highway 16 [1-3 m]) [July] (6; CAS). Freitag (1965:137) cited additional records for this species from Juskatla and Queen Charlotte City (both on Graham Is- land), but I have not seen specimens from these areas. 2. Scaphinotus angusticollis (Mannerheim) Graham Island: Masset (2; USNM). 3. Scaphinotus marginatus (Fischer) Burnaby Island: bay on north shore (SE of Alder Island [10-20 m]) [Aug.] (3; CAS). Graham Island: Ghost Creek (4.7 km NW of Rennell Sound Road [210 m]) [July] (3; CAS), Ghost Creek drainage (7.3 km NW of Rennell Sound Road [240 m]) [July] (1; CAS), Honna Point [May] (2; AMor), Kiusta village site (0.3 km E of [3-5 m]) [Aug.] (1; CAS), Lepas Bay (northeast shore [5-15 m]) [Aug.] (1; CAS), Masset (and 12 km S of [55 m], 14 km E) [July—Aug.] (7; CAS, CNC), Nebria Peak (northeast slope [700-780 mJ], Lower Nebria Lake [640 m]) [July] (4; CAS), Naikoon Provincial Park (Misty Meadow Campground) [July] (3; FMNH), Port Clements [May, Aug.—Sep.] (8; CNC, QCIM), Slatechuck Mountain (east slope [370-790 m], north slope [760— 980 m}) [July] (35; CAS, CNC, FMNH), Tlell River Park [July] (1; FMNH), Tow Hill Park ({5 m], W of headland) [May, July] (6; CAS, QCIM), Tow Hill Road (19 km E of Masset [10 m]) [July] (S; CAS). Hibben Island: [Aug.] (1; UBC). Huxley Island: southeast shore ({ 10-30 m]) [Aug.] (1; CAS). Kunga Island: north shore (S of Titul Island [6-25 m]) [Aug.] (3; CAS), west shore (on Klue Passage [5—SO m]) [Aug.] (3; CAS). Kunghit Island: Rose Harbour [Aug.] (1; UBC). Louise Island: Mount Kermode (south slope [885-1,080 m]) [Aug.] (1; CAS), Skedans village site ([3-30 m]) [Aug.] (12; CAS). Lyell Island: [July] (1; UBC), Gate Creek (at mouth of [3-10 m]) [Aug.] (12; CAS), Powrivco Point (0.5 km SW of [3-20 m]) [Aug.] (3; CAS), Windy Bay [June] (1; QCIM). Moresby Island: Cumshewa village site ({8— 20 m]) [Aug.] (9; CAS, CNC), Darwin Sound (at Hoya Passage [8-30 m]) [Aug.] (1; CAS), Haswell Bay (1.5 km SW of Hoskins Point [5-30 m]) [Aug.] (6; CAS), Jedway (bay 3 km NE of [6- 50 m]) [Aug.] (8; CAS), Kaisun village site ([S—50 m]) [Aug.] (2; CAS), Mount Moresby (south slope [460 m], west slope at High Goose Lake [640 m]) [July] (82; CAS), Pallant Creek (at Camp Moresby [10 m]) [July] (1; CAS), Skincuttle Inlet (0.8 km SW of Huston Point [5-10 m]) [Aug.] (4; CAS), Takakia Lake ([585 mJ], and slopes [590-670 m] and ridges [730-790 m] E of) [July] (27; CAS). Ramsay Island: north shore (1.5 km E of west end [3-4 m]) [Aug.] (1; CAS). Reef Island: north shore (0.6 km E of west end [20 m]) [Aug.] (30; CASA, QCIM). Talunkwan Island: Heming Head (1 km NW of [10-30 m]) [Aug.] (1; CAS). Tanu Island: Tanu village site ({S5-50 mJ) [Aug.] (9; CAS). Island Unknown: (1; CNC). 4. Cychrus tuberculatus Harris Graham Island: Chown Brook [July] (1; CNC), Masset (7; CNC, USNM), Queen Charlotte City [July] (1; QCIM), Skowkona Creek (7.9 km NW of Queen Charlotte City [120 m]) [July— CALIFORNIA ACADEMY OF SCIENCES Aug.] (8; CAS). Louise Island: Skedans village site ([6-25 m]) {Aug.] (1; CAS). 5. Carabus nemoralis Miller Graham Island: Port Clements (0.1 km W of Highway 16 [15 m]) [July—Nov.] (21; CAS, CNC, QCIM). 6. Carabus taedatus Fabricius Moresby Island: Carabus Peak (north slope [850 m]) [July] (1; CAS). 7. Leistus ferruginosus Mannerheim Graham Island: Kiusta village site (0.3 km E of [3-5 m]) [Aug.] (1; CAS), Skowkona Creek (7.9 km NW of Queen Charlotte City {120 m]) [July] (3; CAS). Moresby Island: Kaisun village site ({3-5 m]) [July] (1; CAS). 8. Nebria charlottae Lindroth Graham Island: Bonanza Creek (0.5 km E of Rennell Sound [10 m]) [July] (1; CAS), Kiusta village site (0.3 km E of [3-5 m]) {July—Aug.] (90; CAS, CNC, QCIM), Masset (1; CNC), Masset Inlet (Estrado Lagoon [2-5 m]) [Aug.] (21; CAS), McIntyre Bay (Entry Point to Estrado Lagoon [2-5 m]) [Aug.] (1; CAS), Tow Hill Park (W of Tow Hill [1-3 m]) [July] (347; CAS, QCIM), Yaku Village site (0.2 km SE of [Aug.] (195; CAS, AMor). 9. Nebria louiseae Kavanaugh Burnaby Island: bay on north shore (SE of Alder Island [3-5 m]) [Aug.] (40; CAS). Hotspring Island: north shore ([3—4 m]) {Aug.] (18; CAS). Lyell Island: Gate Creek (at mouth [3-6 m]) {Aug.] (120; CAS, CNC), Powrivco Point (0.5 km SW of [3-5 m]) [Aug.] (149; CAS). Louise Island: Skedans village site [June, Aug.] (6; BCPM, CAS, CNC). Moresby Island: Haswell Bay (1.5 km SW of Hoskins Point [5 m]) [Aug.] (2; CAS), Kaisun village site [July—Aug.] (315; CAS, CNC). Ramsay Island: north shore (1.5 km E of west end [3-4 m]) [Aug.] (11; CAS). Reef Island: north shore (0.6 km E of west end [5 m]) [Aug.] (68; CAS). Skedans Islands: east shore of west island ({5 m]) [Aug.] (6; CAS). Talunkwan Island: Heming Head (1 km NW of [3-5 m]) {[Aug.] (89; CAS). Tanu Island: Tanu village site ([3—5 m]) [Aug.] (123; CAS, CNC). 10. Nebria haida Kavanaugh Graham Island: Nebria Peak (northeast slope [700-780 m], above Middle Nebria Lake [790-910 m]) [July] (288; CAS, CNC). Moresby Island: Mount Moresby (northwest-facing cirque [9 10— 1,070 mJ], east slope of south peak [910-940], northeast slope of south peak [940-1,080 mJ], west slope at High Goose Lake [640 m]) [July] (193; CAS, CNC), Takakia Lake (ridge E of [730-980 m]) [July] (304; CAS, CNC). 11. Nebria sahlbergii sahlbergii Fischer Burnaby Island: bay on north shore (SE of Alder Island [3-20 m]) [Aug.] (18; CAS). Graham Island ([500 m]) [July] (2; FMNH): Awun Lake (0.8-1.2 km W of [35 m], 3.2 km W [50-60 m]) {Aug.] (18; CAS), Bonanza Creek (0.5 km E of Rennell Sound KAVANAUGH—CARABID BEETLES OF THE QUEEN CHARLOTTE ISLANDS 105 [10-20 m]) [July—Aug.] (7; CAS), ChinukundI Creek (at Highway 16 [3 m]) [July] (6; CAS), Ghost Creek (4.3 km NW of Rennell Sound Road [210 m]) [July] (51; CAS, CNC), Ghost Creek drainage (7.3 km NW of Rennell Sound Road [240 m}) [July] (6; CAS, CNC), Gregory Creek (0.3 km E of Rennell Sound [5— 10 mJ) [Aug.] (26; CAS), Mamin River (1.0 km SE of Juskatla [915 m]) [July] (3; CAS), Nebria Peak (at Lower Nebria Lake [620 m]) [July] (7; CAS), Rennell Sound Road (10 km E of Rennell Sound [270 m]) [July] (17; CAS), Sheilds Bay [May] (7; AMor), Skowkona Creek (7.9 km NW of Queen Charlotte City [120 m]) [July] (4; CAS), Slatechuck Mountain (east slope at Tarundl Creek [300-370 m]) [July] (42; CAS), Tow Hill Park (W of Tow Hill) [May] (1; QCIM), Yakoun River (at Port Clem- ents—Juskatla Road ([30 m]) [July] (1; CNC). Lyell Island: Gate Creek (at mouth [3-6 m] and 1.0 km W of mouth [15 m]) [Aug.] (26; CAS, CNC), Powrivco Bay (at Powrivco Creek [15 m]) [Aug.] (11; CAS). Moresby Island: Alliford Bay [May] (3; AMor), High Goose Creek (west base of Mount Moresby [3-90 m]) [July] (19; CAS), Jedway (bay 3 km NE [6-50 m]) [Aug.] (40; CAS), Mount Moresby (west slope at High Goose Lake [640 m)}) [July] (19; CAS, CNC), Pallant Creek ({15 m]) [July] (29; CAS). Tanu Island: Tanu village site ([5—25 m]) [Aug.] (31; CAS, CNC). 12. Nebria mannerheimii Fischer Graham Island: Bonanza Creek (0.5 km E of Rennell Sound [10 m]) [July—Aug.] (23; CAS), Cape Ball River (1 km N of mouth) [Aug.] (1; QCIM), Chinukundl Creek (at Highway 16 [3 m]) [July] (7; CAS), Deep Creek Beach [May] (1; QCIM), Drizzel Lake (2 km SW [60 m]) [July] (46; CAS), Ghost Creek (4.7 km NW of Rennell Sound Road [210 m]) [July] (3; CAS), Gregory Creek (0.3 km E of Rennell Sound [5-10 m]) [Aug.] (69; CAS), Lepas Bay (northeast shore [3-5 m]) [July, Aug.] (87; CAS, CNC, FMNH), Mamin River (1.0 km SE of Juskatla [15 m]) [July] (66; CAS, CNC), Masset (southeast shore of Delkatla Inlet, and 12 km S [55 m], 16 km S) [July—Sep.] (90; CAS, CUIC, ICCM, MCZ, OSUO, UASM, WSU), Port Clements (1.0 km W of Highway 16 [15 m] and 8 km S of) [July—Aug.] (41; CAS), Rennell Sound (at mouth of Bonanza Creek [3-5 m] and 0.5 km NW of Bonanza Creek [3 m]) [July] (3; CAS), Shields Bay [May] (7; AMor), Tow Hill Park ({1-3 m]) [July] (137; CAS), Yakoun River (at Port Clements—Juskatla Road [30 m]) [July— Aug.] (50; CAS). 13. Nebria diversa LeConte Graham Island (3; CAS, ICCM): Masset (and 7 km E of) [May— Aug.] (80; BCPM, CAS, CNC, CUIC, FMNH, ICCM, MCZ, QCIM, UASM, UBC, UMMZ), McIntyre Bay (0.7 km W of Yakan Point at North Beach [2 m], at mouth of Chown Brook [2-5 m]) [May, July—Aug.] (131; CAS, CNC, QCIM), Rose Point [May] (2; OSUO, UBC), Tlell (and 42 km N, 16 km S, Rich- ardson Ranch) [July—Aug., Nov.] (160; CAS, CNC, QCIM, UASM, UBC), Tlell River (at mouth [1-3 m] and | km S) [July] (36; CAS), Tow Hill Park ({1-3 m], at Tow Hill Road) [July— Aug.] (164; CAS, CNC, OSUO). Moresby Island: Sandspit (1 km S of airstrip [1-3 m]) [July] (17; CAS). 14. Notiophilus sylvaticus Eschscholtz Graham Island: Bonanza Creek (0.5 km E of Rennell Sound [10 m]) [July] (1; CAS), Ghost Creek (4.7 km NW of Rennell Sound Road [210 m]) [July] (1; CNC), Ghost Creek drainage (7.3 km NW of Rennell Sound Road [240 m]) [July] (2; CAS), Kiusta village site (0.2 km E of [8 m]) [Aug.] (1; CAS), Nebria Peak (southeast slope [850 m], Lower Nebria Lake [620 m], northeast slope [700-780 m]) [July] (5; CAS, CNC), Port Clements (0.1 km W of Highway 16 [15 m]) [July] (1; CAS), Slatechuck Moun- tain (east slope [370-790 m], Tarundl Creek [370-580 m)]) [July] (3; CAS, CNC), Tlell River (at end of Richardson Road [15 m]) [Aug.] (4; CNC), Tow Hill [July] (2; FMNH), Tow Hill Road (19 km E of Masset [10 m]) [July] (2; CAS). Kunga Island: north shore (S of Titul Island [6-25 m]) [Aug.] (7; CAS, CNC), west shore (on Klue Passage [5—50 m]) [Aug.] (2; CAS). Louise Island: Mount Kermode (south slope [885—1,080 m]) [Aug.] (1; CAS), Skedans village site ([6-30 m]) [Aug.] (13; CAS). Lyell Island: Gate Creek (at mouth [3-10 m]) [Aug.] (4; CAS, CNC), Powrvico Point (0.5 km SE of) [Aug.] (1; CNC). Moresby Island: Carabus Peak (north slope [850 m]) [July] (1; CAS), Darwin Sound (at Hoya Passage [8-30 m]) [Aug.] (1; CAS), Kaisun village site ({3- 50 m]) [Aug.] (9; CAS, CNC), Mount Moresby (south slope [460 m], west slope at High Goose Lake [640 m]) [July] (14; CAS, CNC), Peel Inlet (at road to Moresby Camp [5 m]) [July] (1; CAS), Takakia Lake ([640 m], and ridges E of [730-790 m]) [July] (3; CAS, CNC). Tanu Island: Tanu village site (stream NW of [10-30 m]) [Aug.] (1; CAS). 15. Elaphrus clairvillei Kirby Graham Island: Elaphrus Bog (19.5 km NW of Queen Charlotte City [150 m]) [July] (3; CAS), Ghost Creek drainage (7.3 km NW of Rennell Sound Road [240 m]) [July] (1; CAS). 16. Elaphrus americanus sylvanus Goulet Graham Island: Bonanza Creek (0.5 km E of Rennell Sound [10 m]) [July] (1; CAS), Chinukundl Creek (5.8 km N on Highway 16 [2 m]) [July] (13; CAS), Elaphrus Bog (19.5 km NW of Queen Charlotte City [150 m]) [July] (32; CAS), Highway 16 (1.9 km N of Chinukundl Creek [10 m]) [Aug.] (4; CAS), Masset (south- east shore of Delkatla Inlet [2 m]) [Aug.] (94; CAS), Skowkona Creek (7.9 km NW of Queen Charlotte City [120 m]) [July] (2; CAS). Moresby Island: Mosquito Lake [July] (4, FMNH). 17. Loricera decempunctata Eschscholtz Graham Island: Bonanza Creek (0.5 km E of Rennel Sound [10 m]) [July] (1; CAS), Chinukundl Creek (5.8 km N on Highway 16 [2 m]) [July] (2; CAS), Elaphrus Bog (19.5 km NW of Queen Charlotte City [150 m]) [July-Aug.] (3; CAS), Ghost Creek drainage (7.3 km NW of Rennell Sound Road [240 m]) [July] (1; CAS), Gregory Creek (0.3 km E of Rennell Sound [5-10 m]) [Aug.] (1; CAS), Highway 16 (1.9 km N of Chinukundl Creek [10 m]) [Aug.] (1; CAS), Kiusta village site [Aug.] (3; CNC), Lepas Bay (northeast shore [3-5 m]) [Aug.] (1; CAS), Mamin River (1 km SE of Juskatla) [July] (1; CNC), Masset (southeast shore of Delkatla Inlet [2 m] and 12 km S on Highway 16 [55 m]) [July—Aug.] (37; CAS), Port Clements (0.1 km W of Highway 16 [15 m]) [July—Aug.] (2; CAS), Tlell (40 km N of) [Aug.] (1; CAS), Tow Hill Park ([5 m]) [July] (4; CAS), Tow Hill Road (5.3 km SW of Tow Hill Park) [July] (1; CNC). Moresby Island: Pallant Creek (at Moresby Camp [3-10 m]) [July] (3; CAS); Peel Inlet (at road to Moresby Camp [5 m]) [July] (1; CAS), Takakia Lake ({585 m]) [July] (1; CAS). 106 18. Dyschirius pacificus Lindroth Island Unknown: “Q.C.I.” (1; CNC). 19. Broscodera insignis (Mannerheim) Graham Island: Kiusta village site [Aug.] (1; CNC), Nebria Peak (Lower Nebna Lake [620-640 m]) [July] (2; CAS), Slatechuck Mountain ({910 mJ) [July] (1; CNC), Yaku village site (0.2 km SE of [3-5 m]) [Aug.] (1; CAS). Moresby Island: Mount Moresby (northeast slope of south peak [940-1 ,080 m], west slope at High Goose Lake [640 m]) [July] (7; CAS), Takakia Lake ({585 m], and slopes [610-670 m] and ridges [730-940 m] E of) [July] (29; CAS, CNC). 20. Zacotus matthewsi LeConte Graham Island: Kiusta village site [July] (1; QCIM). 21. Diplous aterrimus (Dejean) Graham Island: Bonanza Creek (0.5 km E of Rennell Sound [10- 20 m]) [July—Aug.] (40; CAS), Chinukundl Creek (at Highway 16 [3 m]) [July] (25; CAS), Ghost Creek (4.7 km NW of Rennell Sound Road [210 m]) [July] (14; CAS, CNC), Gregory Creek (0.3 km E of Rennell Sound [5-10 m]) [Aug.] (15; CAS), Mamin River (1.0 km SE of Juskatla [15 m]) [July] (3; CAS, CNC), Shields Bay [May] (4; AMor), Skowkona Creek (7.9 km NW of Queen Charlotte City [120 m]) [July] (11; CAS), Tlell River (at end of Richardson Road [15 m]) [July] (3; CAS), Yakoun Lake (Etheline Bay at outlet of Yakoun River [110 m]) [Aug.] (4; CAS), Yakoun River (at Port Clements—Juskatla Road [30 m]) [July] (2; CAS). Lyell Island: Gate Creek (1.0 km W of mouth {15 m]) [Aug.] (39; CAS). Moresby Island: Alliford Bay [May] (1; AMor), High Goose Creek (west base of Mount Moresby [3- 90 m]) [July] (2; CAS), Pallant Creek (at Moresby Camp [3-10 mJ], 0.2 km W of Moresby Camp [20 m]) [July—Aug.] (56; CAS, CNC). 22. Trechus obtusus Erichson Graham Island: Masset (southeast [2 m] and southwest [244 m] shores of Delkatla Inlet and 1.4 km S) [Aug.] (22; CAS, CNC). 23. Trechus ovipennis Motschulsky Graham Island: Chinukundl Creek (5.8 km N on Highway 16 {2 m]) [July] (2; CAS), Masset (southeast shore of Delkatla Inlet [2 m] and 1.4 km §S of) [July—Aug.] (6; CAS, CNC), Rennell Sound (at mouth of Bonanza Creek and 0.5 km NW of mouth {3-5 m]) [July-Aug.] (19; CAS), Tlell River (at end of Richard- son Road [15 m]) [July] (1; CAS), Yaku village site (0.2 km SE of [3-5 m]) [Aug.] (2; CAS). Kunga Island: north shore (S of Titul Island [6-25 m]) [Aug.] (8; CAS, CNC). Louise Island: Skedans village site ({6—30 m] and 1.5 km W of [3-15 m]) [Aug.] (136; CAS, CNC). Moresby Island: Jedway (bay 3 km NE [6- 50 m]) [Aug.] (2; CAS), Kaisun village site ({3—5 m]) [Aug.] (1; CAS), Pallant Creek (0.2 km W of Moresby Camp [20 m]) [Aug.] (11; CAS), Sandspit (1 km § of airstrip [2 m]) [July] (1; CAS). Ramsay Island: north shore (1.5 km E of west end [34 m]) [Aug.] (1; CAS). Talunkwan Island: Heming Head (1 km W of) {Aug.] (1; CNC). CALIFORNIA ACADEMY OF SCIENCES 24. Trechus chalybeus Dejean Graham Island: Ghost Creek (4.7 km NW of Rennell Sound Road [210 m]) [July] (6; CAS), Highway 16 (5.8 km N of Chi- nukundl Creek [2 m]), CAS), 12.8 km S of Tlell River bridge) [July] (5; CAS, CNC), Nebria Peak (Lower Nebria Lake [620 m]) [July] (8; CAS), Skowkona Creek (7.9 km NW of Queen Charlotte City [120 m]) [July] (4; CAS), Slatechuck Mountain (north slope [760-980 m], east slope [430-670 m]) [July] (16; CAS, CNC), Tow Hill Road (19 km E of Masset [10 m]) [July] (46; CAS). Moresby Island: Carabus Peak (north slope [850 m]) [July] (1; CAS), Mount Moresby (west slope at High Goose Lake [640 m]) [July] (91; CAS, CNC), Takakia Lake (585-610 m] and ridges E of [610-670 m]) [July] (23; CAS, CNC). 25. Bembidion zephyrum Fall Graham Island: Lepas Bay (northeast shore [3-5 m]) [Aug.] (144; CAS, CNC). 26. Bembidion inaequale opaciceps Casey Graham Island: Yakoun River (5.5 km S of Port Clements [8 m] and at Port Clements—Juskatla Road [30 m]) [July—Aug.] (5; CAS). 27. Bembidion dyschirinum LeConte No specimens seen. 28. Bembidion castum Casey Graham Island ({S00 m]) [July] (3; FMNH): Chinukundl Creek (at Highway 16 [3 m]) [July] (1; CAS), Cinola Mine [May] (3; AMor), Elaphrus Bog (19.5 km NW of Queen Charlotte City [150 m]) [Aug.] (1; CAS), Ghost Creek drainage (7.3 km NW of Rennell Sound Road [240 m]) [July] (1; CAS), Masset (and 1.4km Sand 14km E) [July] (4; CNC, USNM), Queen Charlotte City ({15 m]) [July] (1; CAS), Skowkona Creek (7.9 km NW of Queen Charlotte City [120 m]) [Aug.] (1; CAS), Tow Hill Road (0.5 km E of Chown Brook bridge [10 m], 19 km E of Masset {10 m]) [July—Aug.] (25; CAS). Kunghit Island: ([490 m]) [July] (1; CAS). Louise Island: Mount Kermode (south slope [885-— 1,080 m]) [Aug.] (1; CAS). Moresby Island: Whiteaves Bay [May] (2; AMor). 29. Bembidion iridescens (LeConte) Graham Island: Bonanza Creek (0.5 km E of Rennell Sound [20 m]) [Aug.] (1; CAS), Elaphrus Bog (19.5 km NW of Queen Charlotte City [150 m]) [Aug.] (1; CAS), Ghost Creek drainage (7.3 km NW of Rennell Sound Road [240 m]) [July] (2; CAS), Highway 16 (5.8 km N of Chinukundl Creek [2 m]) [July] (1; CAS), Port Clements [Aug.] (1; CAS), Queen Charlotte City ({30 m]) [July] (1; CAS), Skowkona Creek (7.9 km NW of Queen Charlotte City [120 m]) [July] (14; CAS), Yakoun River (5.5 km S of Port Clements [8 m], at Port Clements—Juskatla Road (30 m]) [July-Aug.] (3; CAS, CNC). Louise Island: Skedans village site (and 1.5 km W of [3-15 m]) [Aug.] (9; CAS, CNC). Moresby Island: Pallant Creek (at Moresby Camp [3-10 m]) [Mar., July] (16; CAS, QCIM). KAVANAUGH—CARABID BEETLES OF THE QUEEN CHARLOTTE ISLANDS 107 30. Bembidion incertum (Motschulsky) Graham Island: Ghost Creek (4.7 km NW of Rennell Sound Road [210 m]) [July] (2; CAS), Ghost Creek drainage (7.3 km NW of Rennell Sound Road [240 m]) [July] (5; CAS), Nebria Peak (northeast slope [700-850 m], Lower Nebria Lake [620- 640 m], above Middle Nebria Lake [790-910 m]) [July] (96; CAS, CNC), Shields Bay [May] (1; AMor), Skowkona Creek (7.9 km NW of Queen Charlotte City [120 m]) [July] (2; CAS), Slate- chuck Mountain (north slope [820-980 m], northeast slope [760— 980 ml], east slope at Tarundl Creek [300 m]) [July] (24; CAS, CNC, FMNH#). Kunghit Island: ([490 m]) [July] (1; CAS). Louise Island: Mount Kermode (south slope [885—1,080 m]) [Aug.] (7; CAS). Moresby Island: Carabus Peak (north slope [850 m]) [July] (2; CAS), Mount Moresby (east slope of south peak [910- 940 m], northeast slope of south peak [940-1 ,080 m], northwest- facing cirque [910-1,070 m], west slope at High Goose Lake [640 m]) [July] (14; CAS, CNC), Takakia Lake (590-610 m] and ridges E of [610-940 m]) [July] (39; CAS, CNC). 31. Bembidion quadrifoveolatum Mannerheim Graham Island: Awun Lake (ridges W of [460-550 m]) [Aug.] (1; CAS), Bonanza Creek (0.5 km E [10-20 m]) [July] (107; CAS), Chinukundl Creek (at Highway 16 [3 m]) [July] (3; CAS), Drizzel Lake (2 km SW [60 m]) [July] (2; CAS), Elaphrus Bog (19.5 km NW of Queen Charlotte City [150 m]) [Aug.] (11; CAS), Ghost Creek (4.7 km NW of Rennell Sound Road [210 m]) [July] (83; CAS, CNC), Ghost Creek drainage (7.3 km NW of Rennell Sound Road [240 m]) [July] (19; CAS), Gregory Creek (0.3 km E of Rennell Sound [5-10 m]) [Aug.] (14; CAS), High- way 16 (5.8 km N on Highway 16 [2 m]) [July] (1; CAS), Mamin River (1.0 km SE of Juskatla [15 m]) [July] (7; CAS), Masset (southeast shore of Delkatla Inlet [2 m], 12 km S on Highway 16 [55 m]) [July—Aug.] (39; CAS), Naikoon Provincial Park (Misty Meadow Campground) [July] (10; FMNH), Port Clem- ents (0.1 km W of Highway 16 [15 m]) [July] (9; CAS), Rennell Sound (0.5 km N of Bonanza Creek [2 m]) [July] (1; CAS), Rennell Sound Road (10 km E of Rennell Sound [270 m]) [July] (2; CAS), Shields Bay [May] (13; AMor), Skowkona Creek (7.9 km NW of Queen Charlotte City [120 m]) [July] (143; CAS), Slatechuck Mountain (northeast slope [760-980 m], east slope at Tarundl Creek [370 m]) [July] (8; CAS), Tlell River (at end of Richardson Road [15 m]) [July] (15; CAS), Yakoun River (5.5 km S of Port Clements [8 m], at Port Clements—Juskatla Road [30 m]) [July—Aug.] (41; CAS, CNC). Louise Island: Ske- dans village site (1.5 km W [3-15 m]) [Aug.] (29; CAS). Lyell Island: Gate Creek (1.0 km W of mouth [15 m]) [Aug.] (24; CAS, CNC), Powrivco Bay (at Lyell Island Logging Camp [15 m]) [Aug.] (9; CAS). Moresby Island: Alliford Bay [May] (3; AMor), Copper Creek (6 km S of Copper Bay [30 m]) [Aug.] (37; CAS), Gray Bay Road (3 km W of Gray Bay [15 m]) [Aug.] (2; CAS), Kaisun village site ([3—5 m]) [Aug.] (12; CAS), Pallant Creek (at Camp Moresby [10 m] and 0.2 km W [15-20 m]) [July—Aug.] (58; CAS, CNC), Takakia Lake ({585-610 m]) [July] (4; CAS), Whiteaves Bay [May] (1; AMor). Tanu Island: Tanu village site ([5—25 m]) [Aug.] (16; CAS). 32. Bembidion farrarae Hatch Graham Island ({[500 m]) [July] (1; FMNH): Bonanza Creek (0.5 km E of Rennell Sound [10 m]) [July] (1; CAS), Nebria Peak (northeast slope [700-850 m]) [July] (2; CAS), Slatechuck Mountain (northeast slope [760-980 m], east slope at Tarundl Creek [370 m]) [July] (4; CAS). Louise Island: Mount Kermode (south slope [885—1,080 m]) [Aug.] (1; CAS). Moresby Island: Mount Moresby (east slope of south peak [910-940 m]) [July] (78; CAS, CNC), Takakia Lake ([585 m] and ridges E of [980 m]) [July] (2; CAS, CNC). 33. Bembidion viator Casey Graham Island: Awun Lake (ridges W of [460-550 m]) [Aug.] (1; CAS), Highway 16 (5.8 km N of Chinukundl Creek [2 m]) [July] (1; CAS), Tow Hill Park ([{5 m]) [July] (1; CAS). Moresby Island: Mosquito Lake [July] (1; FMNH). 34. Bembidion complanulum (Mannerheim) Graham Island: Nebria Peak (northeast slope [700-850 m], above Middle Nebria Lake [790-910 m]) [July] (25; CAS, CNC), Slate- chuck Mountain [760-980 m]) [July] (3; CAS). Louise Island: Mount Kermode (south slope [885—1,080 m]) [Aug.] (13; CAS). Moresby Island: Mount Moresby (east slope of south peak [9 10- 1,080 m], northwest-facing cirque [910-1,070 m]) [July] (114; CAS, CNC), Takakia Lake (ridges E of [730-940 m]) [July] (76; CAS, CNC). : 35. Bembidion planiusculum Mannerheim Graham Island: Bonanza Creek (0.5 km E of Rennell Sound [10-20 m]) [July—Aug.] (137; CAS), Chinukundl Creek (at High- way 16 [3 m]) [July] (2; CAS), Ghost Creek (4.7 km NW of Rennell Sound Road 210 m)) [July] (28; CAS, CNC), Gregory Creek (0.3 km E of Rennell Sound [5-10 m]) [Aug.] (82; CAS), Mamin River (1.0 km SE of Juskatla [15 m]) [July] (62; CAS, CNC), Shields Bay [May] (6; AMor), Tlell River (at end of Richardson Road [15 m]) [July] (3; CAS), Yakoun Lake (Ethe- line Bay at Yakoun River outlet [110 m]) [Aug.] (13; CAS), Yakoun River (5.5 km S of Port Clements [8 m], at Port Cle- ments—Juskatla Road [30 m]) [July—Aug.] (13; CAS). Lyell Is- land: Gate Creek (1.0 km W of mouth [15 m]) [Aug.] (2; CAS). Moresby Island: High Goose Creek (west base of Mount Mores- by [3-90 m]) [July] (4; CAS), Pallant Creek (at Moresby Camp [3-10 m] and 0.2 km W [15-20 m]) [July-Aug.] (101; CAS, CNC). 36. Bembidion sejunctum semiaureum Fall Graham Island: Queen Charlotte City (10 km W at base of Nipple Mountain) [Aug.] (1; CAS), Tlell (1.0 km S of Tlell River mouth [2 m]) [July] (9; CAS, CNC). 37. Bembidion transversale Dejean Graham Island: Awun Lake (west shore) [Aug.] (9; CAS), High- way 16 (1.9 km N of Chinukundl Creek [10 m]) [Aug.] (1; CAS), Mamin River (1.0 km SE of Juskatla [15 m]) [July] (3; CAS), Naikoon Provincial Park (misty Meadows Campground) [July] (16; FMNH), Tlell River (at end of Richardson Road [15 m]) [July] (13; CAS, CNC), Yakoun Lake (Etheline Bay at Yakoun River outlet [110 m]) [Aug.] (1; CAS), Yakoun River (5.5 km S of Port Clements [8 m], at Port Clements—Juskatla Road [30 m]) [July—Aug.] (7; CAS). Moresby Island: Copper Creek (6 km 108 S of Copper Bay [30 m]) [Aug.] (12; CAS), Mosquito Lake (east shore [80 m]) [July] (2; CAS), Pallant Creek (at Moresby Camp [3-10 m] and 0.2 km W of Moresby Camp [15-20 m]) [July- Aug.] (21; CAS). 38. Bembidion incrematum LeConte Graham Island ({500 m]) [July] (5; FMNH): Bonanza Creek (0.5 km E of Rennell Sound [10 m]) [July] (12; CAS), Cinola Mine [May] (9; AMor), Elaphrus Bog (19.5 km NW of Queen Char- lotte City [150 m]) [July] (12; CAS), Ghost Creek (4.7 km NW of Rennell Sound Road [210 m]) [July] (1; CAS), Highway 16 (1.9 km [10 m] and 5.8 km [2 m] N of ChinukundI Creek) [July— Aug.] (127; CAS), Masset (southwest shore of Delkatla Inlet [2- 4 mJ) [Aug.] (1; CAS), McAuley’s Quarry [May] (1; AMor), Skowkona Creek (7.9 km NW of Queen Charlotte City [120 m]) [July] (32; CAS), Tlell River (at end of Richardson Road [15 m]) [July] (2; CAS), Tow Hill Road (19 km E of Masset [10 m]) [July] (2; CAS, CNC), Yakoun River (at Port Clements—Juskatla Road [30 m]) [July] (4; CAS, CNC). Moresby Island: Copper Creek (6 km S of Copper Bay [30 m]) [Aug.] (12; CAS). 39. Bembidion indistinctum Dejean Graham Island: Chown Brook (at Tow Hill Road [5 m]) [Aug.] (14; CAS), Masset (southeast shore of Delkatla Inlet [2 m]) [Aug.] (2; CAS, CNC), Masset Inlet (Estrado Lagoon [2-5 m]) {Aug.] (135; CAS). Moresby Island: Gray Bay ([4 m]) [Aug.] (22; CAS). 40. Bembidion versicolor (LeConte) Graham Island ({500 m]) [July] (2; FMNH): Bonanza Creek (0.5 km E of Rennell Sound [10 m]) [July] (3; CAS), Cinola Mine [May] (16; AMor), Elaphrus Bog (19.5 km NW of Queen Char- lotte City [150 m]) [July—Aug.] (62; CAS, CNC), Highway 16 (1.9 km [10 m] and 5.8 km [2 m] N of Chinukund Creek) [July- Aug.] (54; CAS), Masset (southeast shore of Delkatla Inlet [2 m] and 12 km S on Highway 16 [55 m]) [July-Aug.] (24; CAS, USNM), McAuley’s Quarry [May] (23, AMor), Nebria Peak (above Middle Nebria Lake [790-910 m]) [July] (1; CAS), Skow- kona Creek (7.9 km NW of Queen Charlotte City [120 m]) [July] (18; CAS), Tow Hill Road (19 km E of Masset [10 m]) [July] (3; CAS), Yakoun River (at Port Clements—Juskatla Road [30 m]) [July] (2; CAS). Moresby Island: Mosquito Lake [July] (2; FMNH). 41. Bembidion fortestriatum (Motschulsky) Graham Island: Agonum Bog (11.1 km NW of Queen Charlotte City [150 m]) [July] (1; CAS), Elaphrus Bog (19.5 km NW of Queen Charlotte City [150 m]) [July] (254; CAS, CNC), Ghost Creek drainage (7.3 km NW of Rennell Sound Road [240] (2; CAS), Highway 16 (1.9 km [10 m] and 5.8 km [2 m] N of Chinukundl Creek) [July—-Aug.] (87; CAS), Masset (southeast shore of Delkatla Inlet [2 m] and 14 km E) [July—Aug.] (7; CNC), McAuley’s Quarry [May] (2; AMor), Port Clements (15 km N [60 m]) [July] (1; CAS), Skowkona Creek (7.9 km NW of Queen Charlotte City [120 m]) [July] (2; CAS), Tow Hill Park ({5 m]) [July] (2; CAS), Tow Hill Road (19 km E of Masset [10 m]) [July] (1; CAS). Moresby Island: Mosquito Lake [July] (24; FMNH). CALIFORNIA ACADEMY OF SCIENCES 42. Bembidion spectabile (Mannerheim) Graham Island: Nebria Peak (northeast slope [700-780 m]) [July] (1; CAS), Port Clements [Apr.] (1; QCIM), Skowkona Creek (7.9 km NW of Queen Charlotte City [120 m]) [July] (1; CNC). Huxley Island: southwest shore ({10-30 m]) [Aug.] (1; CAS). Kunga Island: north shore (S of Titul Island [6-25 m]) [Aug.] (1; CAS). Lyell Island: Gate Creek (at mouth [3-10 m]) [Aug.] (1; CAS). Moresby Island: Takakia Lake ([590-610 m]) [July] (2; CAS). 43. Bembidion oblonguloides Lindroth Graham Island ({500 m]) [July] (1; FMNH): Cinola Mine [May] (3; AMor), Ghost Creek drainage (7.3 km NW of Rennell Sound Road [240 m]) [July] (18; CAS, CNC), Masset (6; USNM), Port Clements (0.1 km W of Highway 16 [15 m]) [July-Aug.] (2; CAS, QCIM), Skowkona Creek (7.9 km NW of Queen Charlotte City [120 m]) [July] (2; CAS), Slatechuck Mountain (east slope [430 m]) [July] (5; CAS), Yaku village site (0.2 km SE [3-5 m]) [Aug.] (1; CAS). 44. Bembidion oblongulum (Mannerheim) Moresby Island: Mount Moresby (west slope at High Goose Lake [640 m]) [July] (2; CAS), Takakia Lake ([610 m]) [July] (1; CAS). 45. Nomius pygmaeus (Dejean) Talunkwan Island: Thurston Harbour [July] (1; CAS). 46. Pterostichus lama (Ménétriés) Graham Island: Cape Ball [Aug.] (3; QCIM), Naikoon Provin- cial Park (at park headquarters [15 m]) [July] (2; CAS), Port Clements [Sep.] (1; QCIM), Queen Charlotte City [July] (1; CNC), Tlell River (1 km S of mouth [2 m]) [July] (1; CAS). Kunga Island: west shore (on KJue Passage [5—5SO m]) [Aug.] (1; CAS). Louise Island: Skedans village site (1.5 km W [3-15 m]) [Aug.]) (1; CAS). Lyell Island: Gate Creek (at mouth [3-10 m]) [Aug.] (3; CAS). Moresby Island: Cumshewa village site ([8—20 m]) {Aug.] (2; CAS). Reef Island: north shore (1.0 km E of west end) [Aug.] (1; CNC). Tanu Island: Tanu village site [Sep.] (1; QCIM). 47. Pterostichus crenicollis LeConte Burnaby Island: bay on north shore (SE of Alder Island [3-5 m]) [Aug.] (2; CAS). Chaatl Island: Chaatl village site ([10-30 m]) [Aug.] (1; CAS). Graham Island: Awun Lake (3.2 km W [50-60 m]) [Aug.] (1; CAS), Deep Creek Beach [May] (1; QCIM), Ghost Creek (4.7 km NW of Rennell Sound Road [210 m]) [July] (1; CAS), Ghost Creek drainage (7.3 km NW of Rennell Sound Road [240 m]) [July] (1; CAS), Highway 16 (1.9 km [10 m] and 5.8 km [2 m] N of ChinukundlI Creek) [July—Aug.] (2; CAS), Kiusta village site (0.3 km E of [3-5 m]) [Aug.] (9; CAS, CNC), Lawn Point [May] (1; AMor), Lepas Bay (northeast shore [3-15 m]) [Aug.] (4; CAS), Mamin River (1.0 km SE of Juskatla [15 m]) [July] (6; CAS, CNC), Masset (southwest shore of Del- katla Inlet [2 m] and 1.4 km S on east shore of Masset Inlet [5 m] and 0.6 km NE on Tow Hill Road [10 m]) [July—Aug.] (7; CAS), Naikoon Provincial Park (at park headquarters [15 m]) KAVANAUGH—CARABID BEETLES OF THE QUEEN CHARLOTTE ISLANDS [July] (1; CAS), Port Clements (0.1 km W of Highway 16 [15 m]) [July—Oct.] (15; CAS, QCIM), Queen Charlotte City (1.5 [15 m] and 24 km W) [July—Aug.] (5; CAS, FMNH), Rennell Sound Road (1.5 km N) [May] (1; AMor), Skowkona Creek (7.9 km NW of Queen Charlotte City [120 m]) [July—Aug.] (90; CAS, CNC), Slatechuck Mountain (east slope [370-790 m]) [July] (1; CAS), Tow Hill Park ([S m] and W of headland) [May, July] (16; CAS, QCIM), Yakoun Lake (Etheline Bay at Yakoun River outlet [110 m]) [Aug.] (1; CAS), Yaku village site (0.2 km E of [3-15 m]) [Aug.] (3; CAS). Huxley Island: southeast shore ({10- 30 m]) [Aug.] (1; CAS). Langara Island: Henslung Cove ([5—-8 m]) [Aug.] (3; CAS). Louise Island: Skedans village site ({6-30 mJ] and 1.5 km W [3-15 m]) [Aug.] (15; CAS, CNC). Lyell Island: Gate Creek (at mouth [3-10 m] and 1.0 km W [15 m]) [Aug.] (7; CAS), Powrivco Creek (near mouth [15 m]) [Aug.] (1; CAS), Powrivco Point (0.5 km SW [3-5 m]) [Aug.] (4; CAS). Moresby Island: Copper Creek (6 km S of Copper Bay [30 m]) [Aug.] (6; CAS), Gray Bay ({3-10 m]) [Aug.] (2; CAS), Jedway (bay 3 km E [6-50 m]) [Aug.] (1; CAS), Kaisun village site ([S—50 m]) [Aug.] (3; CAS), Moresby Camp (3 km §S on road to Peel Inlet [12- 140 mj) [July] (1; CAS), Mosquito Lake [July] (1; FMNH), Pallant Creek (at Moresby Camp [3-10 m] and 0.2 km W [15 m]) [July] (3; CAS), Peel Inlet (at road to Moresby Camp [2 m]) [July] (2; CAS), Sandspit [May] (1; AMor), Skincuttle Inlet (0.8 km SW of Huston Point [5-10 m]) [Aug.] (1; CAS). Ramsay Island: north shore (1.5 km E of west end [3-30 m]) [Aug.] (2; CAS). Reef Island: north shore (0.6 km E of west end [5 m]) [Aug.] (3; CAS). Talunkwan Island: Heming Head (1 km NW (3-5 m]) [Aug.] (1; CAS). Tanu Island: Tanu village site ({S—SO m] and stream NW [10-20 m]) [Aug.—Sep.] (4; CAS, QCIM). 48. Pterostichus algidus LeConte Anthony Island: Ninstints [June] (4, BCPM). Burnaby Island: [July] (1; UBC), bay on north shore (SE of Alder Island [3-5 m, 10-20 m]) [Aug.] (8; CAS), Burnaby Narrows (at cabin) [Sep.] (10; QCIM), Section Cove ({3-6 m]) [Aug.] (4; CAS). Chaatl Island: Chaatl village site ([10-30 m]) [Aug.] (2; CAS). East Copper Island: [Sep.] (8; QCIM). Faraday Island: [July] (1; UBC). Graham Island: Agonum Bog (11.1 km NW of Queen Charlotte City [150 m]) [July] (1; CAS), Cape Ball River (at mouth) [Aug.] (1; QCIM), Chown Brook (at Tow Hill Road [5 m]) [Aug.] (1; CAS), Deep Creek Beach [May] (4; QCIM), Elaphrus Bog (19.5 km NW of Queen Charlotte City [150 m]) [Aug.] (1; CAS), Ghost Creek (4.7 km NW of Rennell Sound Road [210 m]) [July] (7; CAS), Gregory Creek (0.3 km E of Rennell Sound [5-10 m]) [Aug.] (1; CAS), Honna Point [May] (6; AMor), Jungle Beach [July] (1; UBC), Kiusta village site (0.3 km E [3-10 m]) [Aug.] (10; CAS), Lawn Point [May] (8; AMor), Lepas Bay (northeast shore [3-15 m]) [Aug.] (3; CAS), Mamin River (1.0 km SE of Juskatla [15 m]) [July] (8; CAS), Masset (southeast [2 m] and southwest [2-4 m] shores of Delkatla Inlet and 0.6 km NE on Tow Hill Road [10 m], 1.4 km S on east shore of Masset Inlet [5S m]) [May, July—Aug.] (57; CAS, CNC), Masset Inlet (at Es- trado Lagoon [2-5 m]) [Aug.] (2; CAS), McIntyre Bay (Entry Point to Estrado Lagoon [2-5 m], North Beach) [May, Aug.] (3; QCIM), Naikoon Provincial Park (Misty Meadows Camp- ground) [July] (3; FMNH), Port Clements (0.1 km W of Highway 16 [15 m]) [May, July—Nov.] (76; CAS, CNC, QCIM, UBC), 109 Queen Charlotte City (and 1.5 km [15 m] and 24 km W) [July— Sep.] (22; CAS, CNC, UBC), Rennell Sound (at mouth of Bo- nanza Creek [3-10 m]) [July—Aug.] (16; CAS), Shields Bay [May] (14; AMor), Skidegate (Second Beach, 3.2 km NE) [May, July— Aug., Oct.] (7; CAS, QCIM), Skonum Point [May] (6; AMor), Skowkona Creek (7.9 km NW of Queen Charlotte City [120 m]) (July—Aug.] (21; CAS, CNC), Tlell (Wiggins Road) [June—July, Sep.] (6: CNC, QCIM), Tlell River (at end of Richardson Road {15 m]), 12.8 km S of bridge on Highway 16, Tlell River Park) [July—Aug.] (10; CAS, CNC, FMNH), Tow Hill Park ({S m], W of headland) [May, July] (18; AMor, CAS, QCIM), Tow Hill Road (0.5 km E of Chown Brook bridge [10 m], 19 km E of Masset [10 m]) [July—Aug.] (31; CAS), Yakoun Lake (Etheline Bay at Yakoun River outlet [110 m]) [Aug.] (5; CAS), Yakoun River (at Port Clements—Juskatla Road [30 m]) [July] (2; CAS), Yaku village site (0.2 km SE of [3-15 m]) [Aug.] (4; CAS). Hotspring Island: [June] (2; BCPM), north shore ({5S—8 m]) [Aug.] (15; CAS). Huxley Island: southeast shore ([10-30 m]) [Aug.] (18; CAS). Kunga Island: north shore (S of Titul Island [6-25 m]) [Aug.] (4; CAS), west shore (on Klue Passage [5-50 m]) [Aug.] (8; CAS, CNC). Kunghit Island: Rose Harbour [Aug.] (1; UBC). Langara Island: Henslung Cove ([S—8 m]) [Aug.] (1; CAS). Louise Island: Skedans village site ({3-10 m] and 1.5 km W [3- 15 m]) [July—Aug.] (57; CAS, CNC, UBC). Lyell Island: Gate Creek (at mouth [3-10 m] and 1.0 km W of [15 m]) [July—Aug.] (41; CAS, UBC), Powrivco Creek (near mouth [15 m]) [Aug.] (2; CAS), Powrivco Point (0.5 km SW of [3-5 m]) [Aug.] (12; CAS, CNC). Maude Island: Renner Pass Farm [June] (14; QCIM). Moresby Island: Alliford Bay ({10 m]) [July] (13; CAS), Copper Bay [June] (1; UBC), Cumshewa village site ([8—20 m]) [Aug.] (14; CAS, CNC), Darwin Sound (at Hoya Passage [8-30 m]) [Aug.] (1; CAS), Gray Bay ({3-10 m]) [Aug.] (11; CAS), Haswell Bay (1.5 km SW of Hoskins Point [5-30 m]) [Aug.] (1; CAS), Houston Stewart Channel (E of Raspberry Point) [June] (4; UBC), Jedway (bay 3 km NE of [6-50 m]) [Aug.] (21; CAS), Kaisun village site ({3-SO m]) [July-Aug.] (26; CAS, CNC), Louscoone Inlet (E of Etches Point) [June] (4; UBC), Moresby Camp (1-3 km S on road to Peel Inlet [12-140 m]) [July—Aug.] (23; CAS, CNC), Mosquito Lake [July] (1; FMNH), Pallant Creek (at Moresby Camp [3-10 m] and 0.2 km W of [15-20 m]) [July-Sep.] (18; CAS, QCIM), Peel Inlet (at road to Moresby Camp [2 m)) [July] (18; CAS), Sandspit (1 km S of airstrip) (May, July—Aug.] (19; AMor, CAS, QCIM), Skincuttle Inlet (0.8 km SW of Huston Point [5—10 m]) [Aug.] (15; CAS), Whiteaves Bay [May] (4; AMor). Murchison Island: [July] (1; UBC). Ram- say Island: north shore (1.5 km E of west end [3-30 m]) [Aug.] (5; CAS). Reef Island: north shore (1.0 km E of west end) [Aug.] (22; CNC, QCIM). Talunkwan Island: Heming Head (1 km SW of [3-5 m, 10-30 m]) [Aug.] (6; CAS, CNC). Tanu Island: Tanu village site ([3—50 m]) [July-Sep.] (69; CAS, CNC, QCIM, UBC). 49. Pterostichus amethystinus Mannerheim Burnaby Island: bay on north shore (SE of Alder Island [10-20 m]) [Aug.] (1; CAS), Section Cove [Aug.] (1; CNC). Chaatl Is- land: Chaat! village site ({10-30 m]) [Aug.] (2; CAS). Graham Island: [May] (1; UBC), Ghost Creek (4.7 km NW of Rennell Sound Road [210 m]) [July] (5; CAS), Honna Point [May] (4; AMor), Kiusta village site (0.3 km E [5-10 m]) [Aug.] (3; CAS), 110 Mamin River (1.0 km SE of Juskatla [15 m]) [July] (4; CAS), Masset ({10 m] and 0.6 km NE on Tow Hill Road [10 m]) [Aug.] (6; CAS), Naikoon Provincial Park (at park headquarters [15 m]) [July] (2; CAS), Nebria Peak (northeast slope [700-850 m]) [July] (1; CAS), Port Clements (0.1 km W of Highway 16 [15 m]) [July, Oct.] (19; CAS, QCIM), Pure Lake Provincial Park ([65 m]) [Aug.] (2; CAS), Queen Charlotte City (and 1.5 [15 m] and 24 km W) [July—Sep.] (9; CAS, CNC, UBC), Rennell Sound (at mouth of Bonanza Creek [3-10 m] [July, Aug.] (4; CAS, FMNH), Skidegate [Aug.] (1; UBC), Skidegate Inlet [Oct.] (1; UBC), Slatechuck Mountain (east slope [370-790 m] and at Tarundl Creek [300 m]) [July] (4; CAS), Tow Hill Park ([5 m]), beach W of headland [3-8 m]) [July] (3; CAS), Tow Hill Road (19 km E of Masset [10 m]) [July] (5; CAS), Yakoun Lake [Aug.] (1; CAS), Yaku village site (0.2 km E [5-15 m]) [Aug.] (2; CAS). Harrison Island: [June] (1; UBC). Hotspring Island: [July] (1; UBC). Huxley Island: southwest shore ({10-30 m]) [Aug.] (1; CAS). Kunga Island: north shore (S of Titul Island [6-25 m]) {[Aug.] (2; CAS), west shore (on Klue Passage [5—SO m]) [Aug.] (3; CAS). Langara Island: [May, July] (3; UBC). Louise Island: Skedans village site ({6-—30 m] and 1.5 km W [3-15 m]) [Aug.] (3; CAS). Lyell Island: Gate Creek (at mouth [3-10 m]) [Aug.] (8; CAS), Powrivco Point (0.5 km SW [5-20 m]) [Aug.] (1; CAS). Morseby Island: Alliford Bay ({10 m]) [July] (6; CAS), Cum- shewa village site ([{8—20 m]) [Aug.] (5; CAS), Haswell Bay (1.5 km SW of Hoskins Point [5-30 m]) [Aug.] (4; CAS, CNC), Kaisun village site ({3—50 m]) [July—Aug.] (5; CAS), Moresby Camp (3 km S on road to Peel Inlet [12-140 m]) [July] (1; CAS), Pallant Creek (at Moresby Camp [3-10 m] and 0.2 km W of Moresby Camp [15 m]) [July] (3; CAS), Peel Inlet (at road to Moresby Camp) [July] (1; CAS), Sandspit [May, July] (3; AMor, UBC), Skincuttle Inlet (0.8 km SW of Huston Point [5-10 m]) [Aug.] (4: CAS), Whiteaves Bay [May] (3; AMor). Ramsay Is- land: [July] (1; UBC), north shore (0.6 km E of west end [5-30 m]) [Aug.] (1; CAS). Reef Island: north shore (0.6 km E of west end [20 m]) [Aug.] (26; CAS, CNC, QCIM). Tanu Island: Tanu village site (S—50 m] and stream NW of [10-30 m]) [Aug.] (11; CAS, CNC). 50. Pterostichus castaneus (Dejean) Chaatl Island: Chaat] village site ({10-—30 m]) [Aug.] (1; CAS). Graham Island: Ghost Creek (4.7 km NW of Rennell Sound Road [210 m]) [July] (3; CAS), Queen Charlotte City (W of [500 m]) [July] (1; FMNH), Rennell Sound (at mouth of Bonanza Creek [5-10 m]) [Aug.] (1; CAS), Skowkona Creek (7.9 km NW of Queen Charlotte City [120 m]) [July] (1; CAS), Slatechuck Mountain (east slope [370-790 m]) [July] (18; CAS), Tlell River Park [July] (1; FMNH), Yaku village site (0.2 km SE [5-15 m]) {Aug.] (3: CAS). Kunga Island: west shore (on Klue Passage [5— 50 m]) [Aug.] (5; CAS). Langara Island: Henslung Cove ({5-8 m]) [Aug.] (1; CAS). Louise Island: Skedans village site ([6-30 m]) [Aug.] (5; CAS). Lyell Island: Gate Creek (at mouth [3-10 mJ) [Aug.] (11; CAS), Powrivco Point (0.5 km SW [5-20 m]) [Aug.] (1; CAS). Moresby Island: Cumshewa village site ([8—20 m]) [Aug.] (3; CAS, CNC), Haswell Bay (1.5 km SW of Hoskins Point [5-30 m]) [Aug.] (6; CAS), Mount Moresby (south slope [460 m], west slope at High Goose Lake [640 m]) [July] (19; CAS), Pallant Creek (at Moresby Camp [3-10 m] and 0.2 km W of Moresby Camp [15 m]) [July—Sep.] (16; CAS, CNC, QCIM), CALIFORNIA ACADEMY OF SCIENCES Skincuttle Inlet (0.8 km SW of Huston Point [5-10 m}) [Aug.] (3; CAS), Peel Inlet (at road to Moresby Camp [2 m]) [July] (1; CAS), Takakia Lake ([590-610 m]) [July] (8; CAS), Whiteaves Bay [May] (1; AMor). Tanu Island: Tanu village site ([5—50 m] and at stream SW of [10-30 m]) [Aug.—Sep.] (6; CAS, QCIM). 51. Pterostichus adstrictus Eschscholtz Graham Island: Chown Brook (at Tow Hill Road [5 m]) [Aug.] (1; CAS), Deep Creek Beach [May] (4; QCIM), Elaphrus Bog (19.5 km NW of Queen Charlotte City [150 m]) [Aug.] (1; CAS), Ghost Creek (4.7 km NW of Rennell Sound Road [210 m]) [May, July] (4; AMor, CAS), Highway 16 (5.8 km N of Chi- nukundl Creek [2 m]) [July] (1; CAS), Kiusta village site (0.3 km E of [3-5 m]) [Aug.] (1; CAS), Lepas Bay (northeast shore (3-15 m]) [Aug.] (7; CAS), Mamin River (1.0 km SE of Juskatla {15 m]) [July] (1; CAS), Masset (southeast [2 m] and southwest {24 m] shores of Delkatla Inlet and 1.4 km S on east shore of Masset Inlet [5 m]) [July-Aug.] (7; CAS, CNC), Masset Inlet (at Estrado Lagoon [2-5 m]) [Aug.] (1; CAS), McAuley’s Quarry [May] (3; AMor), McIntyre Bay (Entry Point to Estrado Lagoon {2-5 m]) [Aug.] (8; CAS), Naikoon Provincial Park (at park headquarters [15 m]) [July] (2; CAS), Port Clements (0.1 km W of Highway 16 [15 m]) [July—-Sep.] (42; CAS, QCIM), Queen Charlotte City (and 1.5 [15 m] and 24 km W) [July—Aug.] (9; CAS), Shields Bay [May] (3; AMor), Skonum Point [May] (4; AMor), Tow Hill Road (0.5 km E of Chown Brook bridge [10 m]) [Aug.] (13; CAS), Yakoun River (at Port Clements—Juskatla Road [30 m]) [July] (45; CAS, CNC). Moresby Island: Gray Bay ({3-10 m]) [Aug.] (10; CAS), Mount Moresby (west slope at High Goose Lake [640 m]) [July] (1; CAS), Pallant Creek (0.2 km E of Moresby Camp [20 m]) [Aug.] (1; CAS), Sandspit (1.0 km S of airstrip [3-5 m]) [July] (4; CAS). 52. Prterostichus riparius (Dejean) Graham Island: Ghost Creek (4.7 km NW of Rennell Sound Road [210 m]) [July] (6; CAS). 53. Agonum ferruginosum (Dejean) Graham Island: Masset (southeast shore of Delkatla Inlet [2 m]) [Aug.] (2; CAS), Tow Hill Road (5.3 km SW of Tow Hill) [Aug.] (2; CAS, CNC). 54. Agonum belleri (Hatch) Graham Island: Agonum Bog (11.1 km NW of Queen Charlotte City [150 m]) [July—Aug.] (8; CAS), Drizzel Lake ({55 m]) [July] (1; CAS), Elaphrus Bog (19.5 km NW of Queen Charlotte City [150 m}) [July] (1; CNC). 55. Agonum metallescens (LeConte) Graham Island: Elaphrus Bog (19.5 km NW of Queen Charlotte City [150 m]) [July] (14; CAS). 56. Agonum brevicolle Dejean Graham Island: Elaphrus Bog (19.5 km NW of Queen Charlotte City [150 m]) [July] (49; CAS, CNC), Ghost Creek drainage (7.3 km NW of Rennell Sound Road [240 m]) [July] (1; CAS), Ren- KAVANAUGH-CARABID BEETLES OF THE QUEEN CHARLOTTE ISLANDS 111 nell Sound Road (1.5 km N) [May] (2; AMor), Tow Hill Park ({S m]) [July] (1; CAS), Tow Hill Road (5.3 km SW of Tow Hill, 19 km E of Masset [10 m]) [July—Aug.] (22; CAS, CNC). Mores- by Island: Mosquito Lake [July] (1; FMNH). 57. Amara sinuosa (Casey) Moresby Island: Mount Moresby (northeast slope of south peak [940-1,080 m]) [July] (5; CAS). 58. Amara ellipsis (Casey) Graham Island: Tlell (1 km S of Tlell River mouth [3-5 m]) [July] (4; CAS). 59. Amara littoralis Mannerheim Graham Island: Bonanza Creek (0.5 km E of Rennell Sound) [Aug.] (1; CNC), Chinukundl Creek (at Highway 16 [3 m]) [July] (1; CAS), Elaphrus Bog (19.5 km NW of Queen Charlotte City [150 m]) [July] (1; CNC), Highway 16 (5.8 km N of Chinukundl Creek [2 m]) [July] (2; CAS), Honna Point [May] (1; AMor), Masset (southeast shore of Delkatla Inlet [2 m] and 1.4 km S on east shore of Masset Inlet [5 m]) [July—Aug.] (2; CAS), Port Clements (0.1 km W of Highway 16 [15 m]) [July—Aug.] (2; CAS), Queen Charlotte City ({30 m] and 1.5 [15 m] and 24 km W) [July—Sep.] (12; CAS). Lyell Island: Gate Creek (at mouth {3-10 m]) [Aug.] (1; CAS). Moresby Island: Alliford Bay ({10 m]) [May, July] (3; CAS, CNC), Pallant Creek [Apr., June] (5; QCIM), Sandspit (1 km S of airstrip [2 m]) [July] (2; CAS). 60. Harpalus somnulentus Dejean Graham Island: Masset (1.4 km S) [Aug.] (1; CNC), Queen Charlotte City ({[30 m] and 1.5 [15 m] and 24 km W) [July— Aug.] (4; CAS). Moresby Island: Pallant Creek [June] (1; QCIM). 61. Trichocellus cognatus (Gyllenhal) Graham Island: Masset (1.4 km S) [Aug.] (16; CAS, CNC), McIntyre Bay (Entry Point to Estrado Lagoon [2—5 m]) [Aug.] (1; CAS), Queen Charlotte City (24 km W) [Aug.] (1; CAS), Tow Hill Road (0.5 km E of Chown Brook bridge [10 m]) [Aug.] (54; CAS). 62. Bradycellus nigrinus (Dejean) Graham Island: Bonanza Creek (0.5 km E of Rennell Sound [10 m]) [July] (1; CAS), Cinola Mine [May] (1; AMor), Elaphrus Bog (19.5 km NW of Queen Charlotte City [150 m]) [July] (13; CAS, CNC), McAuley’s Quarry [May] (4; AMor), Port Clements (0.1 km W of Highway 16 [15 m]) [Apr., July] (2; CAS, QCIM). Moresby Island: Mosquito Lake [July] (11; FMNH), Pallant Creek [Mar.] (7; QCIM). INDEX TO SCIENTIFIC NAMES CALIFORNIA ACADEMY OF SCIENCES Page references to systematic accounts for taxon names are in bold face and to figures and tables are in Italics. {hies lastocarpa 94 Acarina 6, 55, 57 Agonum 73-74 belleri 11, 15, 26, 35, 50, 74, 81, 88, 110 brevicolle 16, 35, 50, 74-75, 81, 88, 110-111 ferruginosum 35, 50, 74, 81, 88, 110 metallescens 16, 35, 50, 74, 81, 88, 110 sordens group 78 Alnus 63 crispa sinuata 28 rubra 27, 63 Amara 75 ellipsis 13, 36, 53, 75, 81, 88, 111 impuncticollis 75 interstitialis 77 littoralis 13-14, 36, 51, 53, 75-76, 82, 85, 88, 111 scitula 77 sinuosa 24, 36, 53, 75, 81, 88, 111 Amphipoda 5, 95 Anchomenus ferruginosus 74 Anisodactylus 76 Aphrophora regina 5, 95 Aves 5, 83 Bembiduini 33, 62 Bembidion 8, 62-69, 77, 96 castum 34, 45, 63, 80, 87, 106 complanulum 24-25, 34, 46, 65, 81, 87, 107 dyschirinum 34, 45, 63, 80, 83, 87, 106 erasum 63 farrarae 22, 24, 34, 45-46, 64-65, 81, 87, 107 forestriatum 15, 35, 47, 68, 81, 88, 92, 108 tnaequale opaciceps 34, 43, 62-63, 80, 87, 106 incertum 22, 24, 34, 45-47, 64, 80, 87, 107 incrematum 16, 35, 45-47, 67, 81, 88, 108 indistinctum 35, 47, 67, 81, 86, 88, 108 iridescens 34, 44, 63-64, 80, 87, 106 oblonguloides 21, 34, 44, 69, 81, 88, 95-96, 108 oblongulum 18, 28, 34, 44, 69, 78, 81, 88, 108 opaciceps 62 peregrinum 68 planiusculum 17, 35, 45-46, 66, 81, 88, 107 quadrifoveolatum 17, 22, 34, 45, 64, 80, 87, 107 seyunctum semiaureum 13, 35, 4 66, 87, 88, 107 semiaureum 66 spectabile 21, 34, 44, 68-69, 81, 88, 108 transversale 19, 35, 47, 66-67, 81, 88, 107-108 versicolor 16, 20, 35, 47, 67-68, 81, 88, 92, 108 viator 34, 45-46, 65, 81, 86-87, 95— 96, 107 zephyrum 34, 43, 62, 80, 87, 106 Bembidium dyschirinum 63 incrematum 67 planiusculum 66 quadrifoveolatum 64 Bledius 59 Blethisa 57 Bradycellus 77 nigrinus 33, 42-43, 77, 82, 88, 111 Broscini 33, 59 Broscodera 59 insignis 22, 33, 42, 59-60, 80, 87, 106 Bufo boreas boreas 5 Calathus 73 Carabini 31, 49-50 Carabus 51-52 nemoralts 31, 38, 51, 80, 87, 90, 96, 98, 104 taedatus 11, 31, 52-53, 77-78, 80, 87, 92, 94, 104 Celia ellipsis 75 sinuosa 75 Cercopidae 5, 95 Chironomidae 5 Chamaecyparis nootkatensis 28 Cicindelini 31, 36 Cicindela 36-37, 37 oregona 31, 36-37, 39-40, 80, S6- 87, 104 oregona guttifera 36 oregona oregona 36 Coleoptera 5 Collembola 53, 57 Corynocera 5 Curculionidae 6, 83, 9/ Cychrini 31, 40 Cychrus 48-49 angustucollis 41 marginatus 43 marginatus var. fullen 44 tuberculatus 17, 31, 38, 48-49, SO, 87, 104 Diplous 60 aterrimus 35, 48, 60-61, 80, 87, 93, 106 filicornis 93 Diptera 5, 6 Dyschirius 59 pacificus 13, 33, 41, 59, 80, 87, 106 tridentatus 59 Elaphrini 33, 57 Elaphrus Fabricius 1/7, 57-58, 77 americanus 92 americanus sylvanus 16, 33, 37, 41, 58, 80, 87, 105 clairvillet 33, 57-58, 80, 85, 87, 92, 105 Epidemia mariposa charlottensis 5 Europhilus 78 Feronia castanea 71 lama 70 riparia 73 valida 71 Gastropoda 40, 50 Geometridae 5, 95 Harpalini 33, 76 Harpalus 76 cognatus 77 nigrinus 77 somnulentus 14, 33, 42-43, 76, 82, 88, 111 Heteroceridae 59 Heteroptera 6 Hydrachnida 6 Hyla regilla 5 Leistus 53, 55 ferruginosus 17, 31, 38, 55, 80, 87, 104 Lepidoptera 5, 50 Loricerini 33, 58 Loricera 58 decempunctata 16, 33, 41, 58, 80, 86-87, 105 Lycaenidae 6 Lygaeidae 6 Mammalia 5, 83 Misocera insignis 59 Morio pygmaeus 70 Nebruini 31, 53, 55 Nebria 7, 55-57, 77, 83, 95 acuta 83 brevicollis 84 charlottae 1, 7, 12, 33, 39-40, 54, 55, 80, 87, 89, 95-96, 104 crassicornis 83 diversa 13, 31, 39-41, 57, 80, 83, 87, 105 gregaria 7,95 gregaria infragroup 95 gvllenhali castanipes 77, 94 haida 7, 11, 24-25, 33, 39-40, 56, 8O, 83, 87, 95-96, 104 kincaidi 83 lituyae 95 loutseae 1, 7, 12, 39-41, 55, 80, 87, 89, 95-97, 105 mannerheimu 31, 37-39, 56, 80, 83, 87, 105 meanyl 83 pipert 83 sahlbergit sahlbergu 17, 22, 33, 39- 41, 56, 80, 83, 87, 104-105 schwarzt schwarz 7 KAVANAUGH—CARABID BEETLES OF THE QUEEN CHARLOTTE ISLANDS Nomuius 69-70 pygmaeus 35, 48, 70, 78, 79, 81, 88, 108 Notaphus incertus 64 Notiophilini 33, 57 Notiophilus 57 sylvaticus 21, 33, 42, 57, 80, 87, 105 Ochthedromus iridescens 63 versicolor 67 Odonata 78, 83, 9/ Oligochaeta 50 Omala fortestriatum 68 Omus 36 Paramoera carlottensis 5,95 Patrobini 35, 60 Patrobus 60, 78 aterrimus 60 Peryphus complanulus 65 Picea sitchensis 27 Pinus contorta 28 Platynini 35, 73, 76 Platynus 73 belleri 74 metallescens 74 Populus tremuloides 28, 94 Psydrini 35, 69, 78 Psydrus 69 Pterostichini 36, 70, 75-76 Pterostichus 8, 70-73, 77 Page references to names in figures and tables are in italics. alder 63 red J7, 27, 55, 63 Sitka 27, 28 amphibians 5 amphipod, freshwater 5, 95 aspen, quaking 28, 94 bear, black 8 beetles 5 rove 6, 59 tiger. See Cicindela oregona variegated mud-loving 59 birds 5, 83 bugs, seed 6 butterfly, northern copper 5 caribou, Dawson’s 5 caterpillars 50 adstrictus 13-14, 36, 52, 73, 81, 88, 110 algidus 14, 17, 36, 37, 41, 43, 48- 49, 51, 71, 72, 78, 81, 88, 97, 109 amethystinus 18, 21, 36, 51-52, 71, 81, 88, 97, 109-110 castaneus 18, 36, 52, 71, 73, 81, 88, 110 crenicollis 17, 36, 52, 70-71, 81, 88, 97, 108-109 lama 18, 36, 51, 70, 87, 88, 108 luczotii 73 riparius 36, 52, 73, 81, 88, 110 Rangifer tarandus dawsoni 5 Saxifraga ferrugines 5 Scaphinotus 40-41, 44 angusticollis 31, 41-43, 77-78, 80, 83, 87, 92, 104 insulans 44 marginatus 6, 18, 21, 30-31, 38, 43- 44, 47-48, 80, 86-87, 92, 97, 104 Scearitini 33, 58-59 Simuliidae 6, 78, 83, 9/ Sphagnum 9, 11, 15, 26, 28, 74 Staphylinidae 6, 59 Stenolophus 76-77 Tachycellus INDEX TO COMMON NAMES cedar yellow 28 western red 27 dragonflies 83, 9/ earthworms 50 fir, subalpine 94 fishes freshwater 5 stickleback 95, 97 flies, black 6, 83, 97 frog, Pacific tree 5 hellebore, false 2/ hemlock mountain 2/, 23, 28 western 27-28 mammals 5, 83 baudiipennis 77 nigrinus 77 Tachys nanus group 78 Tachyta 62, 78 Thya plicata 27 Trachypachus 78 Trechini 35, 61, 76 Trechus 8, 61-62, 68-69 chalybeus 17, 20, 35, 48-49, 62, 80, 87, 106 oblongulum 69 obtusus 35, 49, 61, 80, 87, 90, 96, 98, 106 ovipennis 35, 49, 61, 80, 87, 106 spectabile 68 Tichocellus 76-77 cognatus 33, 42, 77, 79, 82, 88, 111 Tsuga heterophylla 27 mertensiana 28 Vaccinium 55 Veratrum eschscholtzti 21 NXanthorhoe clarkeata 5, 95 Zabrini 36, 75-76 Zacotus 59-60 matthews! 18, 33, 41-42, 60, 80, 86- 87, 106 matthews matthews! 60 matthewsi subopacus 60 midges 5 mites 55, 57 water 6 moths 5, 95 pine, lodgepole (shore) 28 plants, vascular 5, 83, 9/, 95 reptiles 5 slugs 40, 50 snails 40, 50 spittlebugs 5, 95 springtails 55, 57 spruce, Sitka /8, 27 toad, boreal 5 weevils 6, 83, 9/ 113 a "ww SMITHSONIAN INSTITUTION LIBRARIES wii 9088 0 ISBN 0-940228-17-3